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Biology
OpenStax College Rice University 6100 Main Street MS-380 Houston, Texas 77005
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ISBN-10 ISBN-13 Revision
1938168097 978-1-938168-09-3 BM-1-001-DW
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4EFP @LKQBKQ FP >S>FI>?IB CLO COBB >Q EQQM://@KU.LOD/@LKQBKQ/@LI11448/1.9
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+I<=8:< KF B@FCF>P . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 0E@K 1. /?< C?<D@JKIP F= '@=< C?8GKKA #LK@BMQP LC "FLILDV . . . . . . . . . . . . . . . . . . . C?8GKKA -LIB@RIBP: 4EB "RFIAFKD "IL@HP . . . . . 2.2 7>QBO . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3 #>O?LK . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C?8GK@:8C (8:IFDFC<:LC<J . . . . . . . . . . . . . . . . . . . . 3.1 3VKQEBPFP LC "FLILDF@>I ->@OLJLIB@RIBP . . . . . . . . . . . . . . . 3.2 #>O?LEVAO>QBP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3 ,FMFAP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4 0OLQBFKP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.5 .R@IBF@ !@FAP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 0E@K 2. /?< C<CC C?8GKOVLQF@ #BIIP . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3 %RH>OVLQF@ #BIIP . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.4 4EB %KALJBJ?O>KB 3VPQBJ >KA 0OLQBFKP . . . . . . . . . . . . . . 4.5 4EB #VQLPHBIBQLK . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.6 #LKKB@QFLKP ?BQTBBK #BIIP >KA #BIIRI>O !@QFSFQFBP . . . . . . . . . . C?8GKKA 3QOR@QROB . . . . . . . . . . . . . . . . . . . . . . 5.2 0>PPFSB 4O>KPMLOQ . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3 !@QFSB 4O>KPMLOQ . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.4 "RIH 4O>KPMLOQ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C?8GKKA -BQ>?LIFPJ . . . . . . . . . . . . . . . . . . . . . . . . 6.2 0LQBKQF>I, +FKBQF@, &OBB, >KA !@QFS>QFLK %KBODV . . . . . . . . . . . . 6.3 4EB ,>TP LC 4EBOJLAVK>JF@P . . . . . . . . . . . . . . . . . . . . 6.4 !40: !ABKLPFKB 4OFMELPME>QB . . . . . . . . . . . . . . . . . . . . . 6.5 %KWVJBP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C?8GK<I 7: C<CCLC8I -<JG@I8K@FE . . . . . . . . . . . . . . . . . . . . . . . . 7.1 %KBODV FK ,FSFKD 3VPQBJP . . . . . . . . . . . . . . . . . . . . . . . 7.2 'IV@LIVPFP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.3 /UFA>QFLK LC 0VORS>QB >KA QEB #FQOF@ !@FA #V@IB . . . . . . . . . . . . 7.4 /UFA>QFSB 0ELPMELOVI>QFLK . . . . . . . . . . . . . . . . . . . . . . 7.5 -BQ>?LIFPJ TFQELRQ /UVDBK . . . . . . . . . . . . . . . . . . . . . . 7.6 #LKKB@QFLKP LC #>O?LEVAO>QB, 0OLQBFK, >KA ,FMFA -BQ>?LIF@ 0>QET>VP 7.7 2BDRI>QFLK LC #BIIRI>O 2BPMFO>QFLK . . . . . . . . . . . . . . . . . . . C?8GK@QFLKP LC 0ELQLPVKQEBPFP . . . . . . . . . . 8.3 5PFKD ,FDEQ %KBODV QL ->HB /OD>KF@ -LIB@RIBP . . . . . . . . . . . C?8GKIFKD -LIB@RIBP >KA #BIIRI>O 2B@BMQLOP . . . . . . . . . . . . . 9.2 0OLM>D>QFLK LC QEB 3FDK>I . . . . . . . . . . . . . . . . . . . . . . . 9.3 2BPMLKPB QL QEB 3FDK>I . . . . . . . . . . . . . . . . . . . . . . . . 9.4 3FDK>IFKD FK 3FKDIB-#BIIBA /OD>KFPJP . . . . . . . . . . . . . . . . . C?8GK<I 10: C<CC -<GIF;L:K@FE . . . . . . . . . . . . . . . . . . . . . . . . 10.1 #BII $FSFPFLK . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.2 4EB #BII #V@IB . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.3 #LKQOLI LC QEB #BII #V@IB . . . . . . . . . . . . . . . . . . . . . . . 10.4 #>K@BO >KA QEB #BII #V@IB . . . . . . . . . . . . . . . . . . . . . . 10.5 0OLH>OVLQF@ #BII $FSFPFLK . . . . . . . . . . . . . . . . . . . . . . . 0E@K 3. "<E<K@:J C?8GKI 2BMOLAR@QFLK . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 . . . . . . . . . . . . . 13 13 23 39 40 52 59 71 71 73 82 88 98
. 109 . 109 . 113 . 115 . 123 . 128 . 132 . 141 . 142 . 149 . 157 . 161 . 169 . 170 . 173 . 178 . 181 . 183 . 195 . 196 . 200 . 202 . 205 . 209 . 212 . 214 . 223 . 223 . 228 . 235 . 245 . 246 . 254 . 259 . 262 . 273 . 273 . 277 . 283 . 289 . 291 . 301 . 302 . 310
C?8GKKA QEB ,>TP LC 0OL?>?FIFQV . . . . . . . 12.2 #E>O>@QBOFPQF@P >KA 4O>FQP . . . . . . . . . . . . . . . . . . . . 12.3 ,>TP LC )KEBOFQ>K@B . . . . . . . . . . . . . . . . . . . . . . . C?8GKJ F= $E?<I@K8E:< . . . . . . . . . . 13.1 #EOLJLPLJ>I 4EBLOV >KA 'BKBQF@ ,FKH>DB . . . . . . . . . . . 13.2 #EOLJLPLJ>I ">PFP LC )KEBOFQBA $FPLOABOP . . . . . . . . . . . C?8GKI ">PFP LC -LABOK 5KABOPQ>KAFKD . . . . . . . . . . . . 14.2 $.! 3QOR@QROB >KA 3BNRBK@FKD . . . . . . . . . . . . . . . . . 14.3 ">PF@P LC $.! 2BMIF@>QFLK . . . . . . . . . . . . . . . . . . . . 14.4 $.! 2BMIF@>QFLK FK 0OLH>OVLQBP . . . . . . . . . . . . . . . . . 14.5 $.! 2BMIF@>QFLK FK %RH>OVLQBP . . . . . . . . . . . . . . . . . 14.6 $.! 2BM>FO . . . . . . . . . . . . . . . . . . . . . . . . . . . C?8GKOVLQF@ 4O>KP@OFMQFLK . . . . . . . . . . . . . . . . . . . . . 15.3 %RH>OVLQF@ 4O>KP@OFMQFLK . . . . . . . . . . . . . . . . . . . . . 15.4 2.! 0OL@BPPFKD FK %RH>OVLQBP . . . . . . . . . . . . . . . . . 15.5 2F?LPLJBP >KA 0OLQBFK 3VKQEBPFP . . . . . . . . . . . . . . . . C?8GKQFLK LC 'BKB %UMOBPPFLK . . . . . . . . . . . . . . . . . 16.2 0OLH>OVLQF@ 'BKB 2BDRI>QFLK . . . . . . . . . . . . . . . . . . 16.3 %RH>OVLQF@ %MFDBKBQF@ 'BKB 2BDRI>QFLK . . . . . . . . . . . . . 16.4 %RH>OVLQF@ 4O>KP@OFMQFLK 'BKB 2BDRI>QFLK . . . . . . . . . . . . 16.5 %RH>OVLQF@ 0LPQ-QO>KP@OFMQFLK>I 'BKB 2BDRI>QFLK . . . . . . . . 16.6 %RH>OVLQF@ 4O>KPI>QFLK>I >KA 0LPQ-QO>KPI>QFLK>I 'BKB 2BDRI>QFLK 16.7 #>K@BO >KA 'BKB 2BDRI>QFLK . . . . . . . . . . . . . . . . . . C?8GKP 8E; "<EFD@:J . . . . . . . . . . . . . . . . 17.1 "FLQB@EKLILDV . . . . . . . . . . . . . . . . . . . . . . . . . . 17.2 ->MMFKD 'BKLJBP . . . . . . . . . . . . . . . . . . . . . . . . 17.3 7ELIB-'BKLJB 3BNRBK@FKD . . . . . . . . . . . . . . . . . . . 17.4 !MMIVFKD 'BKLJF@P . . . . . . . . . . . . . . . . . . . . . . . 17.5 'BKLJF@P >KA 0OLQBLJF@P . . . . . . . . . . . . . . . . . . . . 0E@K 4. MFCLK@FE8IP +IF:<JJ<J C?8GK@E F= .G<:@<J . . . . . . . . . . . . 18.1 5KABOPQ>KAFKD %SLIRQFLK . . . . . . . . . . . . . . . . . . . . . 18.2 &LOJ>QFLK LC .BT 3MB@FBP . . . . . . . . . . . . . . . . . . . . 18.3 2B@LKKB@QFLK >KA 2>QBP LC 3MB@F>QFLK . . . . . . . . . . . . . C?8GKQFLK %SLIRQFLK . . . . . . . . . . . . . . . . . . . . . . . 19.2 0LMRI>QFLK 'BKBQF@P . . . . . . . . . . . . . . . . . . . . . . . 19.3 !A>MQFSB %SLIRQFLK . . . . . . . . . . . . . . . . . . . . . . . . C?8GK<E@<J 8E; K?< #@JKFIP F= '@=< . . . . . . . . . . . . 20.1 /OD>KFWFKD ,FCB LK %>OQE . . . . . . . . . . . . . . . . . . . . . 20.2 $BQBOJFKFKD %SLIRQFLK>OV 2BI>QFLKPEFMP . . . . . . . . . . . . . 20.3 0BOPMB@QFSBP LK QEB 0EVILDBKBQF@ 4OBB . . . . . . . . . . . . . 0E@K 5. B@FCF>@:8C D@M<IJ@KP C?8GKI %SLIRQFLK, -LOMELILDV, >KA #I>PPFCF@>QFLK . . . . . . . . . 21.2 6FORP )KCB@QFLKP >KA (LPQP . . . . . . . . . . . . . . . . . . . . 21.3 0OBSBKQFLK >KA 4OB>QJBKQ LC 6FO>I )KCB@QFLKP . . . . . . . . . . 21.4 /QEBO !@BIIRI>O %KQFQFBP: 0OFLKP >KA 6FOLFAP . . . . . . . . . . . C?8GKOVLQF@ $FSBOPFQV . . . . . . . . . . . . . . . . . . . . . . . 22.2 3QOR@QROB LC 0OLH>OVLQBP . . . . . . . . . . . . . . . . . . . . . 22.3 0OLH>OVLQF@ -BQ>?LIFPJ . . . . . . . . . . . . . . . . . . . . . 22.4 ">@QBOF>I $FPB>PBP FK (RJ>KP . . . . . . . . . . . . . . . . . . 22.5 "BKBCF@F>I 0OLH>OVLQBP . . . . . . . . . . . . . . . . . . . . . . C?8GKOVLQF@ /OFDFKP . . . . . . . . . . . . . . . . . . . . . . . . 23.2 #E>O>@QBOFPQF@P LC 0OLQFPQP . . . . . . . . . . . . . . . . . . . . 23.3 'OLRMP LC 0OLQFPQP . . . . . . . . . . . . . . . . . . . . . . . .
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. 319 . 320 . 325 . 337 . 353 . 354 . 358 . 371 . 372 . 374 . 381 . 383 . 386 . 388 . 397 . 397 . 402 . 404 . 408 . 412 . 423 . 424 . 426 . 430 . 432 . 435 . 438 . 439 . 447 . 447 . 458 . 461 . 465 . 468 . . . . . . . . . . . . . . . . . . . . . . . . . . . 477 478 486 494 501 502 505 511 521 521 527 533 545 545 552 559 564 573 574 579 588 591 598 609 610 615 617
23.4 %@LILDV LC 0OLQFPQP . . . . . . . . . . . . . . . . . C?8GK@ . . . . . . . . . . . . . . . . . . . . . . 24.1 #E>O>@QBOFPQF@P LC &RKDF . . . . . . . . . . . . . . 24.2 #I>PPFCF@>QFLKP LC &RKDF . . . . . . . . . . . . . . 24.3 %@LILDV LC &RKDF . . . . . . . . . . . . . . . . . . 24.4 &RKD>I 0>O>PFQBP >KA 0>QELDBKP . . . . . . . . . 24.5 )JMLOQ>K@B LC &RKDF FK (RJ>K ,FCB . . . . . . . . C?8GKOIV 0I>KQ ,FCB . . . . . . . . . . . . . . . . . . . 25.2 'OBBK !ID>B: 0OB@ROPLOP LC ,>KA 0I>KQP . . . . . . 25.3 "OVLMEVQBP . . . . . . . . . . . . . . . . . . . . . 25.4 3BBAIBPP 6>P@RI>O 0I>KQP . . . . . . . . . . . . . C?8GKKQP . . . . . . . . . . . . . . 26.2 'VJKLPMBOJP . . . . . . . . . . . . . . . . . . . 26.3 !KDFLPMBOJP . . . . . . . . . . . . . . . . . . . . 26.4 4EB 2LIB LC 3BBA 0I>KQP . . . . . . . . . . . . . . C?8GKQROBP LC QEB !KFJ>I +FKDALJ . . . . . . . . . . 27.2 &B>QROBP 5PBA QL #I>PPFCV !KFJ>IP . . . . . . . . 27.3 !KFJ>I 0EVILDBKV . . . . . . . . . . . . . . . . . 27.4 4EB %SLIRQFLK>OV (FPQLOV LC QEB !KFJ>I +FKDALJ . C?8GK . . . . . . . . . . . . . . . . . . 28.2 0EVIRJ #KFA>OF> . . . . . . . . . . . . . . . . . . 28.3 3RMBOMEVIRJ ,LMELQOL@ELWL> . . . . . . . . . . . 28.4 3RMBOMEVIRJ %@AVPLWL> . . . . . . . . . . . . . 28.5 3RMBOMEVIRJ $BRQBOLPQLJF> . . . . . . . . . . . . C?8GKQBP . . . . . . . . . . . . . . . . . . . . . 29.2 &FPEBP . . . . . . . . . . . . . . . . . . . . . . . 29.3 !JMEF?F>KP . . . . . . . . . . . . . . . . . . . . 29.4 2BMQFIBP . . . . . . . . . . . . . . . . . . . . . . 29.5 "FOAP . . . . . . . . . . . . . . . . . . . . . . . . 29.6 ->JJ>IP . . . . . . . . . . . . . . . . . . . . . . 29.7 4EB %SLIRQFLK LC 0OFJ>QBP . . . . . . . . . . . . . 0E@K 6. +C8EK .KIL:KLI< 8E; !LE:K@FE C?8GKP . . . . . . . . . . 30.1 4EB 0I>KQ "LAV . . . . . . . . . . . . . . . . . . . 30.2 3QBJP . . . . . . . . . . . . . . . . . . . . . . . 30.3 2LLQP . . . . . . . . . . . . . . . . . . . . . . . . 30.4 ,B>SBP . . . . . . . . . . . . . . . . . . . . . . . 30.5 4O>KPMLOQ LC 7>QBO >KA 3LIRQBP FK 0I>KQP . . . . . 30.6 0I>KQ 3BKPLOV 3VPQBJP >KA 2BPMLKPBP . . . . . . C?8GKI 2BNRFOBJBKQP LC 0I>KQP . . . . . . . . . 31.2 4EB 3LFI . . . . . . . . . . . . . . . . . . . . . . 31.3 .RQOFQFLK>I !A>MQ>QFLKP LC 0I>KQP . . . . . . . . . . C?8GKKA 3QOR@QROB . . . . . 32.2 0LIIFK>QFLK >KA &BOQFIFW>QFLK . . . . . . . . . . . . 32.3 !PBUR>I 2BMOLAR@QFLK . . . . . . . . . . . . . . . 0E@K 7. AE@D8C .KIL:KLI< 8E; !LE:K@FE C?8GKI &LOJ >KA &RK@QFLK . . . . . . . . . . . . . 33.2 !KFJ>I 0OFJ>OV 4FPPRBP . . . . . . . . . . . . . . 33.3 (LJBLPQ>PFP . . . . . . . . . . . . . . . . . . . . C?8GK<JK@M< .PJK<D . 34.1 $FDBPQFSB 3VPQBJP . . . . . . . . . . . . . . . . . 34.2 .RQOFQFLK >KA %KBODV 0OLAR@QFLK . . . . . . . . . . 34.3 $FDBPQFSB 3VPQBJ 0OL@BPPBP . . . . . . . . . . . . 34.4 $FDBPQFSB 3VPQBJ 2BDRI>QFLK . . . . . . . . . . . C?8GK<I 35: /?< )<IMFLJ .PJK<D . . . . . . . . . . . . . .
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633 641 642 648 656 662 666 673 674 680 682 687 699 699 705 709 716 727 728 732 738 740 751 751 755 763 778 789 797 798 802 806 811 817 821 824 839 840 842 850 853 861 868 883 883 887 891 901 901 911 922 933 934 939 950 959 960 970 976 982 989
35.1 .BROLKP >KA 'IF>I #BIIP . . . . . . . . . . . . . . . 35.2 (LT .BROLKP #LJJRKF@>QB . . . . . . . . . . . . . 35.3 4EB #BKQO>I .BOSLRP 3VPQBJ . . . . . . . . . . . . 35.4 4EB 0BOFMEBO>I .BOSLRP 3VPQBJ . . . . . . . . . . . 35.5 .BOSLRP 3VPQBJ $FPLOABOP . . . . . . . . . . . . . C?8GKQLPBKP>QFLK . . . . . . . . . . . . . . . . . . 36.3 4>PQB >KA 3JBII . . . . . . . . . . . . . . . . . . . 36.4 (B>OFKD >KA 6BPQF?RI>O 3BKP>QFLK . . . . . . . . . . 36.5 6FPFLK . . . . . . . . . . . . . . . . . . . . . . . . . C?8GKQFLK LC "LAV 0OL@BPPBP . . . . . . . . . . . . 37.4 2BDRI>QFLK LC (LOJLKB 0OLAR@QFLK . . . . . . . . . 37.5 %KAL@OFKB 'I>KAP . . . . . . . . . . . . . . . . . . C?8GKI 3VPQBJP . . . . . . . . . . . . . . 38.2 "LKB . . . . . . . . . . . . . . . . . . . . . . . . . 38.3 *LFKQP >KA 3HBIBQ>I -LSBJBKQ . . . . . . . . . . . . 38.4 -RP@IB #LKQO>@QFLK >KA ,L@LJLQFLK . . . . . . . . . C?8GKP %U@E>KDB . . . . . . . . . . . . . . 39.2 '>P %U@E>KDB >@OLPP 2BPMFO>QLOV 3ROC>@BP . . . . 39.3 "OB>QEFKD . . . . . . . . . . . . . . . . . . . . . . . 39.4 4O>KPMLOQ LC '>PBP FK (RJ>K "LAFIV &IRFAP . . . . . C?8GKQLOV 3VPQBJ . . . . . . . . . 40.2 #LJMLKBKQP LC QEB "ILLA . . . . . . . . . . . . . . 40.3 ->JJ>IF>K (B>OQ >KA "ILLA 6BPPBIP . . . . . . . . 40.4 "ILLA &ILT >KA "ILLA 0OBPPROB 2BDRI>QFLK . . . . . C?8GKLC8K@FE 8E; O:I<K@FE . . . . . . . 41.1 /PJLOBDRI>QFLK >KA /PJLQF@ ">I>K@B . . . . . . . . 41.2 4EB +FAKBVP >KA /PJLOBDRI>QLOV /OD>KP . . . . . . 41.3 %U@OBQFLK 3VPQBJP . . . . . . . . . . . . . . . . . . 41.4 .FQOLDBKLRP 7>PQBP . . . . . . . . . . . . . . . . . 41.5 (LOJLK>I #LKQOLI LC /PJLOBDRI>QLOV &RK@QFLKP . . . C?8GKQB )JJRKB 2BPMLKPB . . . . . . . . . . . . . . 42.2 !A>MQFSB )JJRKB 2BPMLKPB . . . . . . . . . . . . . 42.3 !KQF?LAFBP . . . . . . . . . . . . . . . . . . . . . . 42.4 $FPORMQFLKP FK QEB )JJRKB 3VPQBJ . . . . . . . . . C?8GKQFLK . . . . . . . . . . . . . . . . . . . . . . 43.3 (RJ>K 2BMOLAR@QFSB !K>QLJV >KA '>JBQLDBKBPFP 43.4 (LOJLK>I #LKQOLI LC (RJ>K 2BMOLAR@QFLK . . . . . 43.5 (RJ>K 0OBDK>K@V >KA "FOQE . . . . . . . . . . . . . 43.6 &BOQFIFW>QFLK >KA %>OIV %J?OVLKF@ $BSBILMJBKQ . . . 43.7 /OD>KLDBKBPFP >KA 6BOQB?O>QB &LOJ>QFLK . . . . . . 0E@K 8. :FCF>P C?8GKP 8E; K?< B@FJG?<I< . . . . . . . . . . . 44.1 4EB 3@LMB LC %@LILDV . . . . . . . . . . . . . . . . 44.2 "FLDBLDO>MEV . . . . . . . . . . . . . . . . . . . . 44.3 4BOOBPQOF>I "FLJBP . . . . . . . . . . . . . . . . . . 44.4 !NR>QF@ "FLJBP . . . . . . . . . . . . . . . . . . . 44.5 #IFJ>QB >KA QEB %CCB@QP LC 'IL?>I #IFJ>QB #E>KDB . C?8GKP . . . . . . 45.1 0LMRI>QFLK $BJLDO>MEV . . . . . . . . . . . . . . . 45.2 ,FCB (FPQLOFBP >KA .>QRO>I 3BIB@QFLK . . . . . . . . . 45.3 %KSFOLKJBKQ>I ,FJFQP QL 0LMRI>QFLK 'OLTQE . . . . . 45.4 0LMRI>QFLK $VK>JF@P >KA 2BDRI>QFLK . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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. 989 . 996 1008 1014 1018 1031 1031 1036 1040 1044 1050 1063 1063 1066 1069 1078 1080 1093 1094 1104 1110 1118 1135 1136 1142 1147 1153 1163 1164 1167 1173 1179 1189 1190 1193 1199 1201 1204 1213 1214 1220 1233 1238 1247 1248 1251 1253 1260 1265 1270 1274 1283 1284 1288 1294 1301 1307 1317 1318 1323 1326 1330
45.5 (RJ>K 0LMRI>QFLK 'OLTQE . . . . . . . . . . . . . . . . . . . . 45.6 #LJJRKFQV %@LILDV . . . . . . . . . . . . . . . . . . . . . . . 45.7 "BE>SFLO>I "FLILDV: 0OLUFJ>QB >KA 5IQFJ>QB #>RPBP LC "BE>SFLO C?8GK<I 46: :FJPJK<DJ . . . . . . . . . . . . . . . . . . . . . . . . . 46.1 %@LILDV LC %@LPVPQBJP . . . . . . . . . . . . . . . . . . . . . 46.2 %KBODV &ILT QEOLRDE %@LPVPQBJP . . . . . . . . . . . . . . . . 46.3 "FLDBL@EBJF@>I #V@IBP . . . . . . . . . . . . . . . . . . . . . C?8GKP 8E; B@F;@M<IJ@KP . . . . . . . . . . 47.1 4EB "FLAFSBOPFQV #OFPFP . . . . . . . . . . . . . . . . . . . . . . 47.2 4EB )JMLOQ>K@B LC "FLAFSBOPFQV QL (RJ>K ,FCB . . . . . . . . . . 47.3 4EOB>QP QL "FLAFSBOPFQV . . . . . . . . . . . . . . . . . . . . . . 47.4 0OBPBOSFKD "FLAFSBOPFQV . . . . . . . . . . . . . . . . . . . . . . A AGG<E;@O . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . $E;<O . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . .
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1334 1338 1350 1367 1367 1376 1380 1397 1398 1407 1410 1418 1427 1458
02%&!#% 4/ ")/,/'9
02%&!#% 4/ ")/,/'9
8CJAMKC RM B0636.@, ?L 0NCL4R?V $MJJCEC PCQMSPAC. 5FGQ RCVR@MMI F?Q @CCL APC?RCB UGRF QCTCP?J EM?JQ GL KGLB: ?AACQQG@GJGRW, ASQRMKGX?RGML, ?LB QRSBCLR CLE?ECKCLRq?JJ UFGJC CLAMSP?EGLE QAGCLAC QRSBCLRQ RMU?PB FGEF JCTCJQ MD ?A?BCKGA QAFMJ?PQFGN. *LQRPSARMPQ ?LB QRSBCLRQ ?JGIC UGJJ DGLB RF?R RFGQ RCVR@MMI MDDCPQ ? QRPMLE DMSLB?RGML GL @GMJMEW GL ?L ?AACQQG@JC DMPK?R.
A9FLK *G<E.K8O CFCC<><

0NCL4R?V $MJJCEC GQ ? LML-NPMDGR MPE?LGX?RGML AMKKGRRCB RM GKNPMTGLE QRSBCLR ?AACQQ RM OS?JGRW JC?PLGLE K?RCPG?JQ. 0SP DPCC RCVR@MMIQ ?PC BCTCJMNCB ?LB NCCP-PCTGCUCB @W CBSA?RMPQ RM CLQSPC RFCW ?PC PC?B?@JC, ?AASP?RC, ?LB KCCR RFC QAMNC ?LB QCOSCLAC PCOSGPCKCLRQ MD RMB?WsQ AMJJCEC AMSPQCQ. 6LJGIC RP?BGRGML?J RCVR@MMIQ, 0NCL4R?V $MJJCEC PCQMSPACQ JGTC MLJGLC ?LB ?PC MULCB @W RFC AMKKSLGRW MD CBSA?RMPQ SQGLE RFCK. 5FPMSEF MSP N?PRLCPQFGNQ UGRF AMKN?LGCQ ?LB DMSLB?RGMLQ AMKKGRRCB RM PCBSAGLE AMQRQ DMP QRSBCLRQ, 0NCL4R?V $MJJCEC GQ UMPIGLE RM GKNPMTC ?AACQQ RM FGEFCP CBSA?RGML DMP ?JJ. 0NCL4R?V $MJJCEC GQ ?L GLGRG?RGTC MD 3GAC 6LGTCPQGRW ?LB GQ K?BC NMQQG@JC RFPMSEF RFC ECLCPMSQ QSNNMPR MD QCTCP?J NFGJ?LRFPMNGA DMSLB?RGMLQ.
A9FLK *G<E.K8O CFCC<><WJ -<JFLI:<J

0NCL4R?V $MJJCEC PCQMSPACQ NPMTGBC OS?JGRW ?A?BCKGA GLQRPSARGML. 5FPCC ICW DC?RSPCQ QCR MSP K?RCPG?JQ ?N?PR DPMK MRFCPQ: RFCW A?L @C ASQRMKGXCB @W GLQRPSARMPQ DMP C?AF AJ?QQ, RFCW ?PC ? tJGTGLEu PCQMSPAC RF?R EPMUQ MLJGLC RFPMSEF AMLRPG@SRGMLQ DPMK QAGCLAC CBSA?RMPQ, ?LB RFCW ?PC ?T?GJ?@JC DPCC MP DMP KGLGK?J AMQR.
CLJKFD@Q8K@FE
0NCL4R?V $MJJCEC JC?PLGLE PCQMSPACQ ?PC BCQGELCB RM @C ASQRMKGXCB DMP C?AF AMSPQC. 0SP RCVR@MMIQ NPMTGBC ? QMJGB DMSLB?RGML ML UFGAF GLQRPSARMPQ A?L @SGJB, ?LB MSP PCQMSPACQ ?PC AMLACGTCB ?LB UPGRRCL UGRF DJCVG@GJGRW GL KGLB. *LQRPSARMPQ A?L QCJCAR RFC QCARGMLQ KMQR PCJCT?LR RM RFCGP ASPPGASJ? ?LB APC?RC ? RCVR@MMI RF?R QNC?IQ BGPCARJW RM RFC LCCBQ MD RFCGP AJ?QQCQ ?LB QRSBCLR @MBW. 5C?AFCPQ ?PC CLAMSP?ECB RM CVN?LB ML CVGQRGLE CV?KNJCQ @W ?BBGLE SLGOSC AMLRCVR TG? ECMEP?NFGA?JJW JMA?JGXCB ?NNJGA?RGMLQ ?LB RMNGA?J AMLLCARGMLQ. B0636.@ A?L @C C?QGJW ASQRMKGXCB SQGLE MSP MLJGLC NJ?RDMPK. 4GKNJW QCJCAR RFC AMLRCLR KMQR PCJCT?LR RM WMSP ASPPCLR QCKCQRCP ?LB APC?RC ? RCVR@MMI RF?R QNC?IQ BGPCARJW RM RFC LCCBQ MD WMSP AJ?QQ. B0636.@ GQ MPE?LGXCB ?Q ? AMJJCARGML MD QCARGMLQ RF?R A?L @C PC?PP?LECB, KMBGDGCB, ?LB CLF?LACB RFPMSEF JMA?JGXCB CV?KNJCQ MP RM GLAMPNMP?RC ? QNCAGDGA RFCKC MD WMSP AMSPQC. 5FGQ ASQRMKGX?RGML DC?RSPC UGJJ FCJN @PGLE @GMJMEW RM JGDC DMP WMSP QRSBCLRQ ?LB UGJJ CLQSPC RF?R WMSP RCVR@MMI RPSJW PCDJCARQ RFC EM?JQ MD WMSP AMSPQC.
CLI8K@FE
5M @PM?BCL ?AACQQ ?LB CLAMSP?EC AMKKSLGRW ASP?RGML, B0636.@ GQ tMNCL QMSPACu JGACLQCB SLBCP ? $PC?RGTC $MKKMLQ "RRPG@SRGML ($$-#:) JGACLQC. 5FC QAGCLRGDGA AMKKSLGRW GQ GLTGRCB RM QS@KGR CV?KNJCQ, CKCPEGLE PCQC?PAF, ?LB MRFCP DCCB@?AI RM CLF?LAC ?LB QRPCLERFCL RFC K?RCPG?J ?LB ICCN GR ASPPCLR ?LB PCJCT?LR DMP RMB?WsQ QRSBCLRQ. 4S@KGR WMSP QSEECQRGMLQ RM GLDM!MNCLQR?VAMJJCEC.MPE, ?LB AFCAI GL ML CBGRGML QR?RSQ, ?JRCPL?RC TCPQGMLQ, CPP?R?, ?LB LCUQ ML RFC 4R?V%?QF ?R FRRN://MNCLQR?VAMJJCEC.MPE.
CFJK
0SP RCVR@MMIQ ?PC ?T?GJ?@JC DMP DPCC MLJGLC, ?LB GL JMU-AMQR NPGLR ?LB C-@MMI CBGRGMLQ.
A9FLK B@FCF>P
B0636.@ GQ BCQGELCB DMP KSJRG-QCKCQRCP @GMJMEW AMSPQCQ DMP QAGCLAC K?HMPQ. *R GQ EPMSLBCB ML ?L CTMJSRGML?PW @?QGQ ?LB GLAJSBCQ CVAGRGLE DC?RSPCQ RF?R FGEFJGEFR A?PCCPQ GL RFC @GMJMEGA?J QAGCLACQ ?LB CTCPWB?W ?NNJGA?RGMLQ MD RFC AMLACNRQ ?R F?LB. 5M KCCR RFC LCCBQ MD RMB?WsQ GLQRPSARMPQ ?LB QRSBCLRQ, QMKC AMLRCLR F?Q @CCL QRP?RCEGA?JJW AMLBCLQCB UFGJC K?GLR?GLGLE RFC MTCP?JJ QAMNC ?LB AMTCP?EC MD RP?BGRGML?J RCVRQ DMP RFGQ AMSPQC. *LQRPSARMPQ A?L ASQRMKGXC RFC @MMI, ?B?NRGLE GR RM RFC ?NNPM?AF RF?R UMPIQ
02%&!#% 4/ ")/,/'9
@CQR GL RFCGP AJ?QQPMMK. B0636.@ ?JQM GLAJSBCQ ?L GLLMT?RGTC ?PR NPMEP?K RF?R GLAMPNMP?RCQ APGRGA?J RFGLIGLE ?LB AJGAICP OSCQRGMLQ RM FCJN QRSBCLRQ SLBCPQR?LBq?LB ?NNJWqICW AMLACNRQ.
CFM<I8>< 8E; .:FG<

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p RFC QFCCP @PC?BRF MD RFC RMNGA?J AMTCP?EC. 8C NPMTGBC ? RFMPMSEF RPC?RKCLR MD @GMJMEWsQ DMSLB?RGML?J AMLACNRQ UFGJC AMLBCLQGLE QCJCARCB RMNGAQ GL PCQNMLQC RM RFC K?PICRsQ PCOSCQR DMP ? RCVR@MMI UGRF ? QAMNC RF?R GQ K?L?EC?@JC DMP GLQRPSARMPQ ?LB QRSBCLRQ ?JGIC. 8C ?JQM QRPGTC RM K?IC @GMJMEW, ?Q ? BGQAGNJGLC, GLRCPCQRGLE ?LB ?AACQQG@JC RM QRSBCLRQ. *L ?BBGRGML RM ? AMKNPCFCLQGTC AMTCP?EC MD AMPC AMLACNRQ ?LB DMSLB?RGML?J PCQC?PAF, UC F?TC GLAMPNMP?RCB DC?RSPCQ RF?R BP?U JC?PLCPQ GLRM RFC BGQAGNJGLC GL KC?LGLEDSJ U?WQ. 5FC NCB?EMEGA?J AFMGACQ, AF?NRCP ?PP?LECKCLRQ, ?LB JC?PLGLE M@HCARGTC DSJDGJJKCLR UCPC BCTCJMNCB ?LB TCRRCB UGRF RFC DCCB@?AI MD ?LMRFCP MLC FSLBPCB PCTGCUCPQ, UFM RFMPMSEFJW PC?B RFC K?RCPG?J ?LB MDDCPCB BCR?GJCB APGRGA?J AMKKCLR?PW. 6LGR 1: The Chemistry of Life. 0SP MNCLGLE SLGR GLRPMBSACQ QRSBCLRQ RM RFC QAGCLACQ, GLAJSBGLE RFC QAGCLRGDGA KCRFMB ?LB RFC DSLB?KCLR?J AMLACNRQ MD AFCKGQRPW ?LB NFWQGAQ RF?R NPMTGBC ? DP?KCUMPI UGRFGL UFGAF JC?PLCPQ AMKNPCFCLB @GMJMEGA?J NPMACQQCQ. 6LGR 2: The Cell. 4RSBCLRQ UGJJ E?GL QMJGB SLBCPQR?LBGLE MD RFC QRPSARSPCQ, DSLARGMLQ, ?LB NPMACQQCQ MD RFC KMQR @?QGA SLGR MD JGDC: RFC ACJJ. 6LGR 3: Genetics. 0SP AMKNPCFCLQGTC ECLCRGAQ SLGR R?ICQ JC?PLCPQ DPMK RFC C?PJGCQR CVNCPGKCLRQ RF?R PCTC?JCB RFC @?QGQ MD ECLCRGAQ RFPMSEF RFC GLRPGA?AGCQ MD %/" RM ASPPCLR ?NNJGA?RGMLQ GL RFC CKCPEGLE QRSBGCQ MD @GMRCAFLMJMEW ?LB ECLMKGAQ. 6LGR 4: Evolutionary Processes. 5FC AMPC AMLACNRQ MD CTMJSRGML ?PC BGQASQQCB GL RFGQ SLGR UGRF CV?KNJCQ GJJSQRP?RGLE CTMJSRGML?PW NPMACQQCQ. "BBGRGML?JJW, RFC CTMJSRGML?PW @?QGQ MD @GMJMEW PC?NNC?PQ RFPMSEFMSR RFC RCVR@MMI GL ECLCP?J BGQASQQGML ?LB GQ PCGLDMPACB RFPMSEF QNCAG?J A?JJ-MSR DC?RSPCQ FGEFJGEFRGLE QNCAGDGA CTMJSRGML-@?QCB RMNGAQ. 6LGR 5: Biological Diversity. 5FC BGTCPQGRW MD JGDC GQ CVNJMPCB UGRF BCR?GJCB QRSBW MD T?PGMSQ MPE?LGQKQ ?LB BGQASQQGML MD CKCPEGLE NFWJMECLCRGA PCJ?RGMLQFGNQ. 5FGQ SLGR KMTCQ DPMK TGPSQCQ RM JGTGLE MPE?LGQKQ JGIC @?ARCPG?, BGQASQQCQ RFC MPE?LGQKQ DMPKCPJW EPMSNCB ?Q NPMRGQRQ, ?LB BCTMRCQ KSJRGNJC AF?NRCPQ RM NJ?LR ?LB ?LGK?J JGDC. 6LGR 6: Plant Structure and Function. 0SP NJ?LR SLGR RFMPMSEFJW AMTCPQ RFC DSLB?KCLR?J ILMUJCBEC MD NJ?LR JGDC CQQCLRG?J RM ?L GLRPMBSARMPW @GMJMEW AMSPQC. 6LGR 7: Animal Structure and Function. "L GLRPMBSARGML RM RFC DMPK ?LB DSLARGML MD RFC ?LGK?J @MBW GQ DMJJMUCB @W AF?NRCPQ ML QNCAGDGA @MBW QWQRCKQ ?LB NPMACQQCQ. 5FGQ SLGR RMSAFCQ ML RFC @GMJMEW MD ?JJ MPE?LGQKQ UFGJC K?GLR?GLGLE ?L CLE?EGLE DMASQ ML FSK?L ?L?RMKW ?LB NFWQGMJMEW RF?R FCJNQ QRSBCLRQ AMLLCAR RM RFC RMNGAQ. 6LGR 8: Ecology. &AMJMEGA?J AMLACNRQ ?PC @PM?BJW AMTCPCB GL RFGQ SLGR, UGRF DC?RSPCQ FGEFJGEFRGLE JMA?JGXCB, PC?J-UMPJB GQQSCQ MD AMLQCPT?RGML ?LB @GMBGTCPQGRW.
+<;8>F>@:8C !FLE;8K@FE 8E; !<8KLI<J

B0636.@ GQ EPMSLBCB ML ? QMJGB QAGCLRGDGA @?QC ?LB BCQGELCB RM FCJN QRSBCLRQ SLBCPQR?LB RFC AMLACNRQ ?R F?LB. 5FPMSEFMSR RFC RCVR, MLC A?L CVNJMPC DC?RSPCQ RF?R CLE?EC RFC QRSBCLRQ GL QAGCLRGDGA GLOSGPW @W R?IGLE QCJCARCB RMNGAQ ? QRCN DSPRFCP. 0SP DC?RSPCQ GLAJSBC: Evolution Connection DC?RSPCQ SNFMJB RFC GKNMPR?LAC MD CTMJSRGML RM ?JJ @GMJMEGA?J QRSBW RFPMSEF BGQASQQGMLQ JGIC t5FC &TMJSRGML MD .CR?@MJGA 1?RFU?WQu ?LB t"JE?C ?LB &TMJSRGML?PW 1?RFQ RM 1FMRMQWLRFCQGQ.u Scientific Method Connection A?JJ-MSRQ U?JI QRSBCLRQ RFPMSEF ?ARS?J MP RFMSEFR CVNCPGKCLRQ RF?R CJSAGB?RC RFC QRCNQ MD RFC QAGCLRGDGA NPMACQQ ?Q ?NNJGCB RM RFC RMNGA. 'C?RSPCQ GLAJSBC t%CRCPKGLGLE RFC 5GKC 4NCLR GL $CJJ $WAJC 4R?ECQu ?LB t5CQRGLE RFC )WNMRFCQGQ MD *LBCNCLBCLR "QQMPRKCLR.u Career Connection DC?RSPCQ NPCQCLR GLDMPK?RGML ML ? T?PGCRW MD A?PCCPQ GL RFC @GMJMEGA?J QAGCLACQ, GLRPMBSAGLE QRSBCLRQ RM RFC CBSA?RGML?J PCOSGPCKCLRQ ?LB B?W-RM-B?W UMPI JGDC MD ? T?PGCRW MD NPMDCQQGMLQ, QSAF ?Q KGAPM@GMJMEGQR, CAMJMEGQR, LCSPMJMEGQR, ?LB DMPCLQGA QAGCLRGQR.
10
02%&!#% 4/ ")/,/'9
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!)/+39/,/) +'843/30LC RFGLE GQ AMKKML RM ?JJ DMPKQ MD QAGCLAC: ?L SJRGK?RC EM?J tRM ILMU.u $SPGMQGRW ?LB GLOSGPW ?PC RFC BPGTGLE DMPACQ DMP RFC BCTCJMNKCLR MD QAGCLAC. 4AGCLRGQRQ QCCI RM SLBCPQR?LB RFC UMPJB ?LB RFC U?W GR MNCP?RCQ. 5M BM RFGQ, RFCW SQC RUM KCRFMBQ MD JMEGA?J RFGLIGLE: GLBSARGTC PC?QMLGLE ?LB BCBSARGTC PC?QMLGLE. Inductive reasoning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tJGEFRQ SNu BSPGLE RFGQ ?ARGTGRW GQ RFCL NPCBGARCB RM @C RFC N?PR AMLRPMJJGLE RFC PCQNMLQC RM RFC QCJCARCB QRGKSJSQ, GL RFGQ A?QC, GK?ECQ MD DMMB. 5FC tJGEFRGLE SNu MD RFC T?PGMSQ ?PC?Q MD RFC @P?GL GQ A?SQCB @W CVACQQ ?@QMPNRGML MD P?BGM?ARGTC QSE?P BCPGT?RGTCQ @W ?ARGTC ?PC?Q MD RFC @P?GL. 5FC PCQSJR?LR GLAPC?QC GL P?BGM?ARGTGRW GQ M@QCPTCB @W ? QA?LLCP. 5FCL, PCQC?PAFCPQ A?L QRGKSJ?RC RF?R N?PR MD RFC @P?GL RM QCC GD QGKGJ?P PCQNMLQCQ PCQSJR. %CBSARGTC PC?QMLGLE MP BCBSARGML GQ RFC RWNC MD JMEGA SQCB GL FWNMRFCQGQ-@?QCB QAGCLAC. *L BCBSARGTC PC?QML, RFC N?RRCPL MD RFGLIGLE KMTCQ GL RFC MNNMQGRC BGPCARGML ?Q AMKN?PCB RM GLBSARGTC PC?QMLGLE. Deductive reasoning GQ ? DMPK MD JMEGA?J RFGLIGLE RF?R SQCQ ? ECLCP?J NPGLAGNJC MP J?U RM DMPCA?QR QNCAGDGA PCQSJRQ. 'PMK RFMQC ECLCP?J NPGLAGNJCQ, ? QAGCLRGQR A?L CVRP?NMJ?RC ?LB NPCBGAR RFC QNCAGDGA PCQSJRQ RF?R UMSJB @C T?JGB ?Q JMLE ?Q RFC ECLCP?J NPGLAGNJCQ ?PC T?JGB. 4RSBGCQ GL AJGK?RC AF?LEC A?L GJJSQRP?RC RFGQ RWNC MD PC?QMLGLE. 'MP CV?KNJC, QAGCLRGQRQ K?W NPCBGAR RF?R GD RFC AJGK?RC @CAMKCQ U?PKCP GL ? N?PRGASJ?P PCEGML, RFCL RFC BGQRPG@SRGML MD NJ?LRQ ?LB ?LGK?JQ QFMSJB AF?LEC. 5FCQC NPCBGARGMLQ F?TC @CCL K?BC ?LB RCQRCB, ?LB K?LW QSAF AF?LECQ F?TC @CCL DMSLB, QSAF ?Q RFC KMBGDGA?RGML MD ?P?@JC ?PC?Q DMP ?EPGASJRSPC, UGRF AF?LEC @?QCB ML RCKNCP?RSPC ?TCP?ECQ. #MRF RWNCQ MD JMEGA?J RFGLIGLE ?PC PCJ?RCB RM RFC RUM K?GL N?RFU?WQ MD QAGCLRGDGA QRSBW: BCQAPGNRGTC QAGCLAC ?LB FWNMRFCQGQ-@?QCB QAGCLAC. Descriptive (or discovery) science, UFGAF GQ SQS?JJW GLBSARGTC, ?GKQ RM M@QCPTC, CVNJMPC, ?LB BGQAMTCP, UFGJC hypothesis-based science, UFGAF GQ SQS?JJW BCBSARGTC, @CEGLQ UGRF ? QNCAGDGA OSCQRGML MP NPM@JCK ?LB ? NMRCLRG?J ?LQUCP MP QMJSRGML RF?R A?L @C RCQRCB. 5FC @MSLB?PW @CRUCCL RFCQC RUM DMPKQ MD QRSBW GQ MDRCL @JSPPCB, ?LB KMQR QAGCLRGDGA CLBC?TMPQ AMK@GLC @MRF ?NNPM?AFCQ. 5FC DSXXW @MSLB?PW @CAMKCQ ?NN?PCLR UFCL RFGLIGLE ?@MSR FMU C?QGJW M@QCPT?RGML A?L JC?B RM QNCAGDGA OSCQRGMLQ. 'MP CV?KNJC, ? ECLRJCK?L GL RFC 1940Q M@QCPTCB RF?R RFC @SPP QCCBQ RF?R QRSAI RM FGQ AJMRFCQ ?LB FGQ BMEsQ DSP F?B ? RGLW FMMI QRPSARSPC. 0L AJMQCP GLQNCARGML, FC BGQAMTCPCB RF?R RFC @SPPQs EPGNNGLE BCTGAC U?Q KMPC PCJG?@JC RF?L ? XGNNCP. )C CTCLRS?JJW BCTCJMNCB ? AMKN?LW ?LB NPMBSACB RFC FMMI-?LB-JMMN D?QRCLCP NMNSJ?PJW ILMUL RMB?W ?Q 7CJAPM. %CQAPGNRGTC QAGCLAC ?LB FWNMRFCQGQ-@?QCB QAGCLAC ?PC GL AMLRGLSMSQ BG?JMESC.
/?< .:@<EK@=@: (<K?F;

#GMJMEGQRQ QRSBW RFC JGTGLE UMPJB @W NMQGLE OSCQRGMLQ ?@MSR GR ?LB QCCIGLE QAGCLAC-@?QCB PCQNMLQCQ. 5FGQ ?NNPM?AF GQ AMKKML RM MRFCP QAGCLACQ ?Q UCJJ ?LB GQ MDRCL PCDCPPCB RM ?Q RFC QAGCLRGDGA KCRFMB. 5FC QAGCLRGDGA KCRFMB U?Q SQCB CTCL GL ?LAGCLR RGKCQ, @SR GR U?Q DGPQR BMASKCLRCB @W &LEJ?LBsQ 4GP 'P?LAGQ #?AML (1561p1626) (Figure 1.5), UFM QCR SN GLBSARGTC KCRFMBQ DMP QAGCLRGDGA GLOSGPW. 5FC QAGCLRGDGA KCRFMB GQ LMR CVAJSQGTCJW SQCB @W @GMJMEGQRQ @SR A?L @C ?NNJGCB RM ?JKMQR ?JJ DGCJBQ MD QRSBW ?Q ? JMEGA?J, P?RGML?J NPM@JCK-QMJTGLE KCRFMB.
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!@>LI< 1.5 3FO &O>K@FP ">@LK (1561t1626) FP @OBAFQBA TFQE ?BFKD QEB CFOPQ QL ABCFKB QEB P@FBKQFCF@ JBQELA. (@OBAFQ: 0>RI S>K 3LJBO)
5FC QAGCLRGDGA NPMACQQ RWNGA?JJW QR?PRQ UGRF ?L M@QCPT?RGML (MDRCL ? NPM@JCK RM @C QMJTCB) RF?R JC?BQ RM ? OSCQRGML. -CRsQ RFGLI ?@MSR ? QGKNJC NPM@JCK RF?R QR?PRQ UGRF ?L M@QCPT?RGML ?LB ?NNJW RFC QAGCLRGDGA KCRFMB RM QMJTC RFC NPM@JCK. 0LC .MLB?W KMPLGLE, ? QRSBCLR ?PPGTCQ ?R AJ?QQ ?LB OSGAIJW BGQAMTCPQ RF?R RFC AJ?QQPMMK GQ RMM U?PK. 5F?R GQ ?L M@QCPT?RGML RF?R ?JQM BCQAPG@CQ ? NPM@JCK: RFC AJ?QQPMMK GQ RMM U?PK. 5FC QRSBCLR RFCL ?QIQ ? OSCQRGML: t8FW GQ RFC AJ?QQPMMK QM U?PK u P74548/3- ' H>549.+8/8 3CA?JJ RF?R ? FWNMRFCQGQ GQ ? QSEECQRCB CVNJ?L?RGML RF?R A?L @C RCQRCB. 5M QMJTC ? NPM@JCK, QCTCP?J FWNMRFCQCQ K?W @C NPMNMQCB. 'MP CV?KNJC, MLC FWNMRFCQGQ KGEFR @C, t5FC AJ?QQPMMK GQ U?PK @CA?SQC LM MLC RSPLCB ML RFC ?GP AMLBGRGMLGLE.u #SR RFCPC AMSJB @C MRFCP PCQNMLQCQ RM RFC OSCQRGML, ?LB RFCPCDMPC MRFCP FWNMRFCQCQ K?W @C NPMNMQCB. " QCAMLB FWNMRFCQGQ KGEFR @C, t5FC AJ?QQPMMK GQ U?PK @CA?SQC RFCPC GQ ? NMUCP D?GJSPC, ?LB QM RFC ?GP AMLBGRGMLGLE BMCQLsR UMPI.u 0LAC ? FWNMRFCQGQ F?Q @CCL QCJCARCB, RFC QRSBCLR A?L K?IC ? NPCBGARGML. " NPCBGARGML GQ QGKGJ?P RM ? FWNMRFCQGQ @SR GR RWNGA?JJW F?Q RFC DMPK?R t*D . . . RFCL . . . .u 'MP CV?KNJC, RFC NPCBGARGML DMP RFC DGPQR FWNMRFCQGQ KGEFR @C, tI- RFC QRSBCLR RSPLQ ML RFC ?GP AMLBGRGMLGLE, ;/,5 RFC AJ?QQPMMK UGJJ LM JMLECP @C RMM U?PK.u "+89/3- ' H>549.+8/8 " T?JGB FWNMRFCQGQ KSQR @C RCQR?@JC. *R QFMSJB ?JQM @C falsifiable, KC?LGLE RF?R GR A?L @C BGQNPMTCL @W CVNCPGKCLR?J PCQSJRQ. *KNMPR?LRJW, QAGCLAC BMCQ LMR AJ?GK RM tNPMTCu ?LWRFGLE @CA?SQC QAGCLRGDGA SLBCPQR?LBGLEQ ?PC ?JU?WQ QS@HCAR RM KMBGDGA?RGML UGRF DSPRFCP GLDMPK?RGML. 5FGQ QRCNqMNCLLCQQ RM BGQNPMTGLE GBC?QqGQ UF?R BGQRGLESGQFCQ QAGCLACQ DPMK LML-QAGCLACQ. 5FC NPCQCLAC MD RFC QSNCPL?RSP?J, DMP GLQR?LAC, GQ LCGRFCP RCQR?@JC LMP D?JQGDG?@JC. 5M RCQR ? FWNMRFCQGQ, ? PCQC?PAFCP UGJJ AMLBSAR MLC MP KMPC CVNCPGKCLRQ BCQGELCB RM CJGKGL?RC MLC MP KMPC MD RFC FWNMRFCQCQ. &?AF CVNCPGKCLR UGJJ F?TC MLC MP KMPC T?PG?@JCQ ?LB MLC MP KMPC AMLRPMJQ. " variable GQ ?LW N?PR MD RFC CVNCPGKCLR RF?R A?L T?PW MP AF?LEC BSPGLE RFC CVNCPGKCLR. 5FC control group AMLR?GLQ CTCPW DC?RSPC MD RFC CVNCPGKCLR?J EPMSN CVACNR GR GQ LMR EGTCL RFC K?LGNSJ?RGML RF?R GQ FWNMRFCQGXCB ?@MSR. 5FCPCDMPC, GD RFC PCQSJRQ MD RFC CVNCPGKCLR?J EPMSN BGDDCP DPMK RFC AMLRPMJ EPMSN, RFC BGDDCPCLAC KSQR @C BSC RM RFC FWNMRFCQGXCB K?LGNSJ?RGML, P?RFCP RF?L QMKC MSRQGBC D?ARMP. -MMI DMP RFC T?PG?@JCQ ?LB AMLRPMJQ GL RFC CV?KNJCQ RF?R DMJJMU. 5M RCQR RFC DGPQR FWNMRFCQGQ, RFC QRSBCLR UMSJB DGLB MSR GD RFC ?GP AMLBGRGMLGLE GQ ML. *D RFC ?GP AMLBGRGMLGLE GQ RSPLCB ML @SR BMCQ LMR UMPI, RFCPC QFMSJB @C ?LMRFCP PC?QML, ?LB RFGQ FWNMRFCQGQ QFMSJB @C PCHCARCB. 5M RCQR RFC QCAMLB FWNMRFCQGQ, RFC QRSBCLR AMSJB AFCAI GD RFC JGEFRQ GL RFC AJ?QQPMMK ?PC DSLARGML?J. *D QM, RFCPC GQ LM NMUCP D?GJSPC ?LB RFGQ FWNMRFCQGQ QFMSJB @C PCHCARCB. &?AF FWNMRFCQGQ QFMSJB @C RCQRCB @W A?PPWGLE MSR ?NNPMNPG?RC CVNCPGKCLRQ. #C ?U?PC RF?R PCHCARGLE MLC FWNMRFCQGQ BMCQ LMR BCRCPKGLC UFCRFCP MP LMR RFC MRFCP FWNMRFCQCQ A?L @C ?AACNRCB; GR QGKNJW CJGKGL?RCQ MLC FWNMRFCQGQ RF?R GQ LMR T?JGB (Figure 1.6). 6QGLE RFC QAGCLRGDGA KCRFMB, RFC FWNMRFCQCQ RF?R ?PC GLAMLQGQRCLR UGRF CVNCPGKCLR?J B?R? ?PC PCHCARCB. 8FGJC RFGQ tU?PK AJ?QQPMMKu CV?KNJC GQ @?QCB ML M@QCPT?RGML?J PCQSJRQ, MRFCP FWNMRFCQCQ ?LB CVNCPGKCLRQ KGEFR F?TC AJC?PCP AMLRPMJQ. 'MP GLQR?LAC, ? QRSBCLR KGEFR ?RRCLB AJ?QQ ML .MLB?W ?LB PC?JGXC QFC F?B BGDDGASJRW AMLACLRP?RGLE ML RFC JCARSPC. 0LC M@QCPT?RGML RM CVNJ?GL RFGQ MAASPPCLAC KGEFR
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@C, t8FCL * C?R @PC?ID?QR @CDMPC AJ?QQ, * ?K @CRRCP ?@JC RM N?W ?RRCLRGML.u 5FC QRSBCLR AMSJB RFCL BCQGEL ?L CVNCPGKCLR UGRF ? AMLRPMJ RM RCQR RFGQ FWNMRFCQGQ. *L FWNMRFCQGQ-@?QCB QAGCLAC, QNCAGDGA PCQSJRQ ?PC NPCBGARCB DPMK ? ECLCP?J NPCKGQC. 5FGQ RWNC MD PC?QMLGLE GQ A?JJCB BCBSARGTC PC?QMLGLE: BCBSARGML NPMACCBQ DPMK RFC ECLCP?J RM RFC N?PRGASJ?P. #SR RFC PCTCPQC MD RFC NPMACQQ GQ ?JQM NMQQG@JC: QMKCRGKCQ, QAGCLRGQRQ PC?AF ? ECLCP?J AMLAJSQGML DPMK ? LSK@CP MD QNCAGDGA M@QCPT?RGMLQ. 5FGQ RWNC MD PC?QMLGLE GQ A?JJCB GLBSARGTC PC?QMLGLE, ?LB GR NPMACCBQ DPMK RFC N?PRGASJ?P RM RFC ECLCP?J. *LBSARGTC ?LB BCBSARGTC PC?QMLGLE ?PC MDRCL SQCB GL R?LBCK RM ?BT?LAC QAGCLRGDGA ILMUJCBEC (Figure 1.7).
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!@>LI< 1.6 4EB P@FBKQFCF@ JBQELA @LKPFPQP LC > PBOFBP LC TBII-ABCFKBA PQBMP. )C > EVMLQEBPFP FP KLQ PRMMLOQBA ?V BUMBOFJBKQ>I A>Q>, > KBT EVMLQEBPFP @>K ?B MOLMLPBA.
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wQ TLOH TEBK MIRDDBA FKQL FQ. @. -V QL>PQBO ALBPKwQ QL>PQ JV ?OB>A. A. ) MIRD JV @LCCBB J>HBO FKQL QEB LRQIBQ. B. -V @LCCBBJ>HBO TLOHP. C. 7EV ALBPKwQ JV QL>PQBO TLOH
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!@>LI< 1.7 3@FBKQFPQP RPB QTL QVMBP LC OB>PLKFKD, FKAR@QFSB >KA ABAR@QFSB OB>PLKFKD, QL >AS>K@B P@FBKQFCF@ HKLTIBADB. !P FP QEB @>PB FK QEFP BU>JMIB, QEB @LK@IRPFLK COLJ FKAR@QFSB OB>PLKFKD @>K LCQBK ?B@LJB QEB MOBJFPB CLO FKAR@QFSB OB>PLKFKD.
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sR LCACQQ?PGJW QAGCLRGDGA GL L?RSPC.
/NF /PG<J F= .:@<E:<: B8J@: .:@<E:< 8E; AGGC@<; .:@<E:<

5FC QAGCLRGDGA AMKKSLGRW F?Q @CCL BC@?RGLE DMP RFC J?QR DCU BCA?BCQ ?@MSR RFC T?JSC MD BGDDCPCLR RWNCQ MD QAGCLAC. *Q GR T?JS?@JC RM NSPQSC QAGCLAC DMP RFC Q?IC MD QGKNJW E?GLGLE ILMUJCBEC, MP BMCQ QAGCLRGDGA ILMUJCBEC MLJW F?TC UMPRF GD UC A?L ?NNJW GR RM QMJTGLE ? QNCAGDGA NPM@JCK MP RM @CRRCPGLE MSP JGTCQ 5FGQ OSCQRGML DMASQCQ ML RFC BGDDCPCLACQ @CRUCCL RUM RWNCQ MD QAGCLAC: @?QGA QAGCLAC ?LB ?NNJGCB QAGCLAC. Basic science MP tNSPCu QAGCLAC QCCIQ RM CVN?LB ILMUJCBEC PCE?PBJCQQ MD RFC QFMPR-RCPK ?NNJGA?RGML MD RF?R ILMUJCBEC. *R GQ LMR DMASQCB ML BCTCJMNGLE ? NPMBSAR MP ? QCPTGAC MD GKKCBG?RC NS@JGA MP AMKKCPAG?J T?JSC. 5FC GKKCBG?RC EM?J MD @?QGA QAGCLAC GQ ILMUJCBEC DMP ILMUJCBECsQ Q?IC, RFMSEF RFGQ BMCQ LMR KC?L RF?R, GL RFC CLB, GR K?W LMR PCQSJR GL ? NP?ARGA?J ?NNJGA?RGML.
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38
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46
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48
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49
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4MKC ?RMKQ ?PC KMPC QR?@JC UFCL RFCW E?GL MP JMQC ?L CJCARPML (MP NMQQG@JW RUM) ?LB DMPK GMLQ. 5FGQ DGJJQ RFCGP MSRCPKMQR CJCARPML QFCJJ ?LB K?ICQ RFCK CLCPECRGA?JJW KMPC QR?@JC. #CA?SQC RFC LSK@CP MD CJCARPMLQ BMCQ LMR COS?J RFC LSK@CP MD NPMRMLQ, C?AF GML F?Q ? LCR AF?PEC. Cations ?PC NMQGRGTC GMLQ RF?R ?PC DMPKCB @W JMQGLE CJCARPMLQ. /CE?RGTC GMLQ ?PC DMPKCB @W E?GLGLE CJCARPMLQ ?LB ?PC A?JJCB ?LGMLQ. Anions ?PC BCQGEL?RCB @W RFCGP CJCKCLR?J L?KC @CGLE ?JRCPCB RM CLB GL t-GBCu: RFC ?LGML MD AFJMPGLC GQ A?JJCB AFJMPGBC, ?LB RFC ?LGML MD QSJDSP GQ A?JJCB QSJDGBC, DMP CV?KNJC. 5FGQ KMTCKCLR MD CJCARPMLQ DPMK MLC CJCKCLR RM ?LMRFCP GQ PCDCPPCB RM ?Q electron transfer. "Q Figure 2.11 GJJSQRP?RCQ, QMBGSK (/?) MLJW F?Q MLC CJCARPML GL GRQ MSRCP CJCARPML QFCJJ. *R R?ICQ JCQQ CLCPEW DMP QMBGSK RM BML?RC RF?R MLC CJCARPML RF?L GR BMCQ RM ?AACNR QCTCL KMPC CJCARPMLQ RM DGJJ RFC MSRCP QFCJJ. *D QMBGSK JMQCQ ?L CJCARPML, GR LMU F?Q 11 NPMRMLQ, 11 LCSRPMLQ, ?LB MLJW 10 CJCARPMLQ, JC?TGLE GR UGRF ?L MTCP?JJ AF?PEC MD +1. *R GQ LMU PCDCPPCB RM ?Q ? QMBGSK GML. $FJMPGLC ($J) GL GRQ JMUCQR CLCPEW QR?RC (A?JJCB RFC EPMSLB QR?RC) F?Q QCTCL CJCARPMLQ GL GRQ MSRCP QFCJJ. "E?GL, GR GQ KMPC CLCPEW-CDDGAGCLR DMP AFJMPGLC RM E?GL MLC CJCARPML RF?L RM JMQC QCTCL. 5FCPCDMPC, GR RCLBQ RM E?GL ?L CJCARPML RM APC?RC ?L GML UGRF 17 NPMRMLQ, 17 LCSRPMLQ, ?LB 18 CJCARPMLQ, EGTGLE GR ? LCR LCE?RGTC (p1) AF?PEC. *R GQ LMU PCDCPPCB RM ?Q ? AFJMPGBC GML. *L RFGQ CV?KNJC, QMBGSK UGJJ BML?RC GRQ MLC CJCARPML RM CKNRW GRQ QFCJJ, ?LB AFJMPGLC UGJJ ?AACNR RF?R CJCARPML RM DGJJ GRQ QFCJJ. #MRF GMLQ LMU Q?RGQDW RFC MARCR PSJC ?LB F?TC AMKNJCRC MSRCPKMQR QFCJJQ. #CA?SQC RFC LSK@CP MD CJCARPMLQ GQ LM JMLECP COS?J RM RFC LSK@CP MD NPMRMLQ, C?AF GQ LMU ?L GML ?LB F?Q ? +1 (QMBGSK A?RGML) MP p1 (AFJMPGBC ?LGML) AF?PEC. /MRC RF?R RFCQC RP?LQ?ARGMLQ A?L LMPK?JJW MLJW R?IC NJ?AC QGKSJR?LCMSQJW: GL MPBCP DMP ? QMBGSK ?RMK RM JMQC ?L CJCARPML, GR KSQR @C GL RFC NPCQCLAC MD ? QSGR?@JC PCAGNGCLR JGIC ? AFJMPGLC ?RMK.
50
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51
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52
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
2.2 R 28K<I
#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC NPMNCPRGCQ MD U?RCP RF?R ?PC APGRGA?J RM K?GLR?GLGLE JGDC z &VNJ?GL UFW U?RCP GQ ?L CVACJJCLR QMJTCLR z 1PMTGBC CV?KNJCQ MD U?RCPsQ AMFCQGTC ?LB ?BFCQGTC NPMNCPRGCQ z %GQASQQ RFC PMJC MD ?AGBQ, @?QCQ, ?LB @SDDCPQ GL FMKCMQR?QGQ
#(!04%2 2 X 4(% #(%-)#!, &/5.$!4)/. /& ,)&%
53
8FW BM QAGCLRGQRQ QNCLB RGKC JMMIGLE DMP U?RCP ML MRFCP NJ?LCRQ 8FW GQ U?RCP QM GKNMPR?LR *R GQ @CA?SQC U?RCP GQ CQQCLRG?J RM JGDC ?Q UC ILMU GR. 8?RCP GQ MLC MD RFC KMPC ?@SLB?LR KMJCASJCQ ?LB RFC MLC KMQR APGRGA?J RM JGDC ML &?PRF. "NNPMVGK?RCJW 70 NCPACLR MD RFC FSK?L @MBW GQ K?BC SN MD U?RCP. 8GRFMSR GR, JGDC ?Q UC ILMU GR QGKNJW UMSJB LMR CVGQR. 5FC NMJ?PGRW MD RFC U?RCP KMJCASJC ?LB GRQ PCQSJRGLE FWBPMECL @MLBGLE K?IC U?RCP ? SLGOSC QS@QR?LAC UGRF QNCAG?J NPMNCPRGCQ RF?R ?PC GLRGK?RCJW RGCB RM RFC NPMACQQCQ MD JGDC. -GDC MPGEGL?JJW CTMJTCB GL ? U?RCPW CLTGPMLKCLR, ?LB KMQR MD ?L MPE?LGQKsQ ACJJSJ?P AFCKGQRPW ?LB KCR?@MJGQK MAASP GLQGBC RFC U?RCPW AMLRCLRQ MD RFC ACJJsQ AWRMNJ?QK. 4NCAG?J NPMNCPRGCQ MD U?RCP ?PC GRQ FGEF FC?R A?N?AGRW ?LB FC?R MD T?NMPGX?RGML, GRQ ?@GJGRW RM BGQQMJTC NMJ?P KMJCASJCQ, GRQ AMFCQGTC ?LB ?BFCQGTC NPMNCPRGCQ, ?LB GRQ BGQQMAG?RGML GLRM GMLQ RF?R JC?BQ RM RFC ECLCP?RGML MD N). 6LBCPQR?LBGLE RFCQC AF?P?ARCPGQRGAQ MD U?RCP FCJNQ RM CJSAGB?RC GRQ GKNMPR?LAC GL K?GLR?GLGLE JGDC.
28K<IWJ +FC8I@KP
0LC MD U?RCPsQ GKNMPR?LR NPMNCPRGCQ GQ RF?R GR GQ AMKNMQCB MD NMJ?P KMJCASJCQ: RFC FWBPMECL ?LB MVWECL UGRFGL U?RCP KMJCASJCQ ()20) DMPK NMJ?P AMT?JCLR @MLBQ. 8FGJC RFCPC GQ LM LCR AF?PEC RM ? U?RCP KMJCASJC, RFC NMJ?PGRW MD U?RCP APC?RCQ ? QJGEFRJW NMQGRGTC AF?PEC ML FWBPMECL ?LB ? QJGEFRJW LCE?RGTC AF?PEC ML MVWECL, AMLRPG@SRGLE RM U?RCPsQ NPMNCPRGCQ MD ?RRP?ARGML. 8?RCPsQ AF?PECQ ?PC ECLCP?RCB @CA?SQC MVWECL GQ KMPC CJCARPMLCE?RGTC RF?L FWBPMECL, K?IGLE GR KMPC JGICJW RF?R ? QF?PCB CJCARPML UMSJB @C DMSLB LC?P RFC MVWECL LSAJCSQ RF?L RFC FWBPMECL LSAJCSQ, RFSQ ECLCP?RGLE RFC N?PRG?J LCE?RGTC AF?PEC LC?P RFC MVWECL. "Q ? PCQSJR MD U?RCPsQ NMJ?PGRW, C?AF U?RCP KMJCASJC ?RRP?ARQ MRFCP U?RCP KMJCASJCQ @CA?SQC MD RFC MNNMQGRC AF?PECQ @CRUCCL U?RCP KMJCASJCQ, DMPKGLE FWBPMECL @MLBQ. 8?RCP ?JQM ?RRP?ARQ MP GQ ?RRP?ARCB RM MRFCP NMJ?P KMJCASJCQ ?LB GMLQ. " NMJ?P QS@QR?LAC RF?R GLRCP?ARQ PC?BGJW UGRF MP BGQQMJTCQ GL U?RCP GQ PCDCPPCB RM ?Q hydrophilic (FWBPM- = tU?RCPu; -NFGJGA = tJMTGLEu). *L AMLRP?QR, LML-NMJ?P KMJCASJCQ QSAF ?Q MGJQ ?LB D?RQ BM LMR GLRCP?AR UCJJ UGRF U?RCP, ?Q QFMUL GL Figure 2.13 ?LB QCN?P?RC DPMK GR P?RFCP RF?L BGQQMJTC GL GR, ?Q UC QCC GL Q?J?B BPCQQGLEQ AMLR?GLGLE MGJ ?LB TGLCE?P (?L ?AGBGA U?RCP QMJSRGML). 5FCQC LMLNMJ?P AMKNMSLBQ ?PC A?JJCB hydrophobic (FWBPM- = tU?RCPu; -NFM@GA = tDC?PGLEu).
!@>LI< 2.13 /FI >KA T>QBO AL KLQ JFU. !P QEFP J>@OL FJ>DB LC LFI >KA T>QBO PELTP, LFI ALBP KLQ AFPPLISB FK T>QBO ?RQ CLOJP AOLMIBQP FKPQB>A. 4EFP FP ARB QL FQ ?BFKD > KLKMLI>O @LJMLRKA. (@OBAFQ: '>RQ>J $LDO>).
28K<IWJ .K8K<J: "8J, '@HL@;, 8E; .FC@;

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
54
#(!04%2 2 X 4(% #(%-)#!, &/5.$!4)/. /& ,)&%
@PC?I RFC FWBPMECL @MLBQ) RF?R K?ICQ GAC JCQQ BCLQC RF?L JGOSGB U?RCP, ? NFCLMKCLML LMR QCCL GL RFC QMJGBGDGA?RGML MD MRFCP JGOSGBQ. 8?RCPsQ JMUCP BCLQGRW GL GRQ QMJGB DMPK GQ BSC RM RFC U?W FWBPMECL @MLBQ ?PC MPGCLRCB ?Q GR DPCCXCQ: RFC U?RCP KMJCASJCQ ?PC NSQFCB D?PRFCP ?N?PR AMKN?PCB RM JGOSGB U?RCP. 8GRF KMQR MRFCP JGOSGBQ, QMJGBGDGA?RGML UFCL RFC RCKNCP?RSPC BPMNQ GLAJSBCQ RFC JMUCPGLE MD IGLCRGA CLCPEW @CRUCCL KMJCASJCQ, ?JJMUGLE RFCK RM N?AI CTCL KMPC RGEFRJW RF?L GL JGOSGB DMPK ?LB EGTGLE RFC QMJGB ? EPC?RCP BCLQGRW RF?L RFC JGOSGB. 5FC JMUCP BCLQGRW MD GAC, GJJSQRP?RCB ?LB NGARSPCB GL Figure 2.14, ?L ?LMK?JW, A?SQCQ GR RM DJM?R ?R RFC QSPD?AC MD JGOSGB U?RCP, QSAF ?Q GL ?L GAC@CPE MP GL RFC GAC AS@CQ GL ? EJ?QQ MD GAC U?RCP. *L J?ICQ ?LB NMLBQ, GAC UGJJ DMPK ML RFC QSPD?AC MD RFC U?RCP APC?RGLE ?L GLQSJ?RGLE @?PPGCP RF?R NPMRCARQ RFC ?LGK?JQ ?LB NJ?LR JGDC GL RFC NMLB DPMK DPCCXGLE. 8GRFMSR RFGQ J?WCP MD GLQSJ?RGLE GAC, NJ?LRQ ?LB ?LGK?JQ JGTGLE GL RFC NMLB UMSJB DPCCXC GL RFC QMJGB @JMAI MD GAC ?LB AMSJB LMR QSPTGTC. 5FC BCRPGKCLR?J CDDCAR MD DPCCXGLE ML JGTGLE MPE?LGQKQ GQ A?SQCB @W RFC CVN?LQGML MD GAC PCJ?RGTC RM JGOSGB U?RCP. 5FC GAC APWQR?JQ RF?R DMPK SNML DPCCXGLE PSNRSPC RFC BCJGA?RC KCK@P?LCQ CQQCLRG?J DMP RFC DSLARGML MD JGTGLE ACJJQ, GPPCTCPQG@JW B?K?EGLE RFCK. $CJJQ A?L MLJW QSPTGTC DPCCXGLE GD RFC U?RCP GL RFCK GQ RCKNMP?PGJW PCNJ?ACB @W ?LMRFCP JGOSGB JGIC EJWACPMJ.
!@>LI< 2.14 (VAOLDBK ?LKAFKD J>HBP F@B IBPP ABKPB QE>K IFNRFA T>QBO. 4EB (>) I>QQF@B PQOR@QROB LC F@B J>HBP FQ IBPP ABKPB QE>K QEB COBBIV CILTFKD JLIB@RIBP LC IFNRFA T>QBO, BK>?IFKD FQ QL (?) CIL>Q LK T>QBO. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V *>KB 7EFQKBV, FJ>DB @OB>QBA RPFKD 6FPR>I -LIB@RI>O $VK>JF@P ;1< (6-$) PLCQT>OB ; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V #>OILP 0LKQB)
$JGAI here (http://openstaxcollege.org/l/ice_lattice2) RM QCC ? 3-% ?LGK?RGML MD RFC QRPSARSPC MD ?L GAC J?RRGAC. (*K?EC APCBGR: +?LC 8FGRLCW. *K?EC APC?RCB SQGLE 7GQS?J .MJCASJ?P %WL?KGAQ 7.% <2= QMDRU?PC. )
28K<IWJ #@>? #<8K C8G8:@KP

8?RCPsQ FGEF FC?R A?N?AGRW GQ ? NPMNCPRW A?SQCB @W FWBPMECL @MLBGLE ?KMLE U?RCP KMJCASJCQ. 8?RCP F?Q RFC FGEFCQR specific heat capacity MD ?LW JGOSGBQ. 4NCAGDGA FC?R GQ BCDGLCB ?Q RFC ?KMSLR MD FC?R MLC EP?K MD ? QS@QR?LAC KSQR ?@QMP@ MP JMQC RM AF?LEC GRQ RCKNCP?RSPC @W MLC BCEPCC $CJQGSQ. 'MP U?RCP, RFGQ ?KMSLR GQ MLC calorie. *R RFCPCDMPC R?ICQ U?RCP ? JMLE RGKC RM FC?R ?LB JMLE RGKC RM AMMJ. *L D?AR, RFC QNCAGDGA FC?R A?N?AGRW MD U?RCP GQ ?@MSR DGTC RGKCQ KMPC RF?L RF?R MD Q?LB. 5FGQ CVNJ?GLQ UFW RFC J?LB AMMJQ D?QRCP RF?L RFC QC?. %SC RM GRQ FGEF FC?R A?N?AGRW, U?RCP GQ SQCB @W U?PK @JMMBCB ?LGK?JQ RM KMPC
1. 8. )SKNFPCW 8., ". %?JIC, ?LB ,. 4AFSJRCL, t7.%q7GQS?J .MJCASJ?P %WL?KGAQ,u J6<95(3 6- M63,*<3(9 G9(7/0*: 14 (1996): 33-38.
2. 8. )SKNFPCW 8., ". %?JIC, ?LB ,. 4AFSJRCL, t7.%q7GQS?J .MJCASJ?P %WL?KGAQ,u J6<95(3 6- M63,*<3(9 G9(7/0*: 14 (1996): 33-38.
#(!04%2 2 X 4(% #(%-)#!, &/5.$!4)/. /& ,)&%
55
CTCLJW BGQNCPQC FC?R GL RFCGP @MBGCQ: GR ?ARQ GL ? QGKGJ?P K?LLCP RM ? A?PsQ AMMJGLE QWQRCK, RP?LQNMPRGLE FC?R DPMK U?PK NJ?ACQ RM AMMJ NJ?ACQ, A?SQGLE RFC @MBW RM K?GLR?GL ? KMPC CTCL RCKNCP?RSPC.
28K<IWJ #<8K F= 18GFI@Q8K@FE

8?RCP ?JQM F?Q ? FGEF heat of vaporization, RFC ?KMSLR MD CLCPEW PCOSGPCB RM AF?LEC MLC EP?K MD ? JGOSGB QS@QR?LAC RM ? E?Q. " AMLQGBCP?@JC ?KMSLR MD FC?R CLCPEW (586 A?J) GQ PCOSGPCB RM ?AAMKNJGQF RFGQ AF?LEC GL U?RCP. 5FGQ NPMACQQ MAASPQ ML RFC QSPD?AC MD U?RCP. "Q JGOSGB U?RCP FC?RQ SN, FWBPMECL @MLBGLE K?ICQ GR BGDDGASJR RM QCN?P?RC RFC JGOSGB U?RCP KMJCASJCQ DPMK C?AF MRFCP, UFGAF GQ PCOSGPCB DMP GR RM CLRCP GRQ E?QCMSQ NF?QC (QRC?K). "Q ? PCQSJR, U?RCP ?ARQ ?Q ? FC?R QGLI MP FC?R PCQCPTMGP ?LB PCOSGPCQ KSAF KMPC FC?R RM @MGJ RF?L BMCQ ? JGOSGB QSAF ?Q CRF?LMJ (EP?GL ?JAMFMJ), UFMQC FWBPMECL @MLBGLE UGRF MRFCP CRF?LMJ KMJCASJCQ GQ UC?ICP RF?L U?RCPsQ FWBPMECL @MLBGLE. &TCLRS?JJW, ?Q U?RCP PC?AFCQ GRQ @MGJGLE NMGLR MD 100\ $CJQGSQ (212\ '?FPCLFCGR), RFC FC?R GQ ?@JC RM @PC?I RFC FWBPMECL @MLBQ @CRUCCL RFC U?RCP KMJCASJCQ, ?LB RFC IGLCRGA CLCPEW (KMRGML) @CRUCCL RFC U?RCP KMJCASJCQ ?JJMUQ RFCK RM CQA?NC DPMK RFC JGOSGB ?Q ? E?Q. &TCL UFCL @CJMU GRQ @MGJGLE NMGLR, U?RCPsQ GLBGTGBS?J KMJCASJCQ ?AOSGPC CLMSEF CLCPEW DPMK MRFCP U?RCP KMJCASJCQ QSAF RF?R QMKC QSPD?AC U?RCP KMJCASJCQ A?L CQA?NC ?LB T?NMPGXC: RFGQ NPMACQQ GQ ILMUL ?Q evaporation. 5FC D?AR RF?R FWBPMECL @MLBQ LCCB RM @C @PMICL DMP U?RCP RM CT?NMP?RC KC?LQ RF?R ? QS@QR?LRG?J ?KMSLR MD CLCPEW GQ SQCB GL RFC NPMACQQ. "Q RFC U?RCP CT?NMP?RCQ, CLCPEW GQ R?ICL SN @W RFC NPMACQQ, AMMJGLE RFC CLTGPMLKCLR UFCPC RFC CT?NMP?RGML GQ R?IGLE NJ?AC. *L K?LW JGTGLE MPE?LGQKQ, GLAJSBGLE GL FSK?LQ, RFC CT?NMP?RGML MD QUC?R, UFGAF GQ 90 NCPACLR U?RCP, ?JJMUQ RFC MPE?LGQK RM AMMJ QM RF?R FMKCMQR?QGQ MD @MBW RCKNCP?RSPC A?L @C K?GLR?GLCB.
28K<IWJ .FCM<EK +IFG<IK@<J

4GLAC U?RCP GQ ? NMJ?P KMJCASJC UGRF QJGEFRJW NMQGRGTC ?LB QJGEFRJW LCE?RGTC AF?PECQ, GMLQ ?LB NMJ?P KMJCASJCQ A?L PC?BGJW BGQQMJTC GL GR. 5FCPCDMPC, U?RCP GQ PCDCPPCB RM ?Q ? solvent, ? QS@QR?LAC A?N?@JC MD BGQQMJTGLE MRFCP NMJ?P KMJCASJCQ ?LB GMLGA AMKNMSLBQ. 5FC AF?PECQ ?QQMAG?RCB UGRF RFCQC KMJCASJCQ UGJJ DMPK FWBPMECL @MLBQ UGRF U?RCP, QSPPMSLBGLE RFC N?PRGAJC UGRF U?RCP KMJCASJCQ. 5FGQ GQ PCDCPPCB RM ?Q ? sphere of hydration, MP ? FWBP?RGML QFCJJ, ?Q GJJSQRP?RCB GL Figure 2.15 ?LB QCPTCQ RM ICCN RFC N?PRGAJCQ QCN?P?RCB MP BGQNCPQCB GL RFC U?RCP 8FCL GMLGA AMKNMSLBQ ?PC ?BBCB RM U?RCP, RFC GLBGTGBS?J GMLQ PC?AR UGRF RFC NMJ?P PCEGMLQ MD RFC U?RCP KMJCASJCQ ?LB RFCGP GMLGA @MLBQ ?PC BGQPSNRCB GL RFC NPMACQQ MD dissociation. %GQQMAG?RGML MAASPQ UFCL ?RMKQ MP EPMSNQ MD ?RMKQ @PC?I MDD DPMK KMJCASJCQ ?LB DMPK GMLQ. $MLQGBCP R?@JC Q?JR (/?$J, MP QMBGSK AFJMPGBC): UFCL /?$J APWQR?JQ ?PC ?BBCB RM U?RCP, RFC KMJCASJCQ MD /?$J BGQQMAG?RC GLRM /?+ ?LB $Jp GMLQ, ?LB QNFCPCQ MD FWBP?RGML DMPK ?PMSLB RFC GMLQ, GJJSQRP?RCB GL Figure 2.15. 5FC NMQGRGTCJW AF?PECB QMBGSK GML GQ QSPPMSLBCB @W RFC N?PRG?JJW LCE?RGTC AF?PEC MD RFC U?RCP KMJCASJCsQ MVWECL. 5FC LCE?RGTCJW AF?PECB AFJMPGBC GML GQ QSPPMSLBCB @W RFC N?PRG?JJW NMQGRGTC AF?PEC MD RFC FWBPMECL ML RFC U?RCP KMJCASJC.
!@>LI< 2.15 7EBK Q>?IB P>IQ (.>#I) FP JFUBA FK T>QBO, PMEBOBP LC EVAO>QFLK >OB CLOJBA >OLRKA QEB FLKP.
28K<IWJ CF?<J@M< 8E; A;?<J@M< +IFG<IK@<J

)?TC WMS CTCP DGJJCB ? EJ?QQ MD U?RCP RM RFC TCPW RMN ?LB RFCL QJMUJW ?BBCB ? DCU KMPC BPMNQ #CDMPC GR MTCPDJMUQ, RFC U?RCP DMPKQ ? BMKC-JGIC QF?NC ?@MTC RFC PGK MD RFC EJ?QQ. 5FGQ U?RCP A?L QR?W ?@MTC RFC EJ?QQ @CA?SQC MD RFC NPMNCPRW MD cohesion. *L AMFCQGML, U?RCP KMJCASJCQ ?PC ?RRP?ARCB RM C?AF MRFCP (@CA?SQC MD FWBPMECL @MLBGLE), ICCNGLE RFC KMJCASJCQ RMECRFCP ?R RFC JGOSGB-E?Q (U?RCP-?GP) GLRCPD?AC, ?JRFMSEF RFCPC GQ LM KMPC PMMK GL RFC EJ?QQ. $MFCQGML ?JJMUQ DMP RFC BCTCJMNKCLR MD surface tension, RFC A?N?AGRW MD ? QS@QR?LAC RM UGRFQR?LB @CGLE PSNRSPCB UFCL NJ?ACB SLBCP RCLQGML MP QRPCQQ. 5FGQ GQ ?JQM UFW U?RCP DMPKQ BPMNJCRQ UFCL NJ?ACB ML ? BPW QSPD?AC P?RFCP RF?L @CGLE DJ?RRCLCB MSR @W EP?TGRW. 8FCL ? QK?JJ QAP?N MD N?NCP GQ NJ?ACB MLRM RFC BPMNJCR MD U?RCP, RFC N?NCP DJM?RQ ML RMN MD RFC U?RCP BPMNJCR CTCL RFMSEF N?NCP GQ BCLQCP (FC?TGCP) RF?L RFC U?RCP. $MFCQGML ?LB QSPD?AC RCLQGML ICCN RFC FWBPMECL @MLBQ MD U?RCP KMJCASJCQ GLR?AR ?LB QSNNMPR
56
#(!04%2 2 X 4(% #(%-)#!, &/5.$!4)/. /& ,)&%
RFC GRCK DJM?RGLE ML RFC RMN. *RsQ CTCL NMQQG@JC RM tDJM?Ru ? LCCBJC ML RMN MD ? EJ?QQ MD U?RCP GD GR GQ NJ?ACB ECLRJW UGRFMSR @PC?IGLE RFC QSPD?AC RCLQGML, ?Q QFMUL GL Figure 2.16.
!@>LI< 2.16 4EB TBFDEQ LC QEB KBBAIB FP MRIIFKD QEB PROC>@B ALTKT>OA; >Q QEB P>JB QFJB, QEB PROC>@B QBKPFLK FP MRIIFKD FQ RM, PRPMBKAFKD FQ LK QEB PROC>@B LC QEB T>QBO >KA HBBMFKD FQ COLJ PFKHFKD. .LQF@B QEB FKABKQ>QFLK FK QEB T>QBO >OLRKA QEB KBBAIB. (@OBAFQ: #LOV :>KHBO)
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58
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5FC SLGOSC NPMNCPRGCQ MD U?RCP RF?R AMLRPG@SRC RM RFGQ A?N?AGRW RM @?J?LAC N)q?Q UCJJ ?Q U?RCPsQ MRFCP AF?P?ARCPGQRGAQq?PC CQQCLRG?J RM QSQR?GLGLE JGDC ML &?PRF.
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60
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5FC RFPCC-BGKCLQGML?J NJ?ACKCLR MD ?RMKQ ?LB AFCKGA?J @MLBQ UGRFGL MPE?LGA KMJCASJCQ GQ ACLRP?J RM SLBCPQR?LBGLE RFCGP AFCKGQRPW. .MJCASJCQ RF?R QF?PC RFC Q?KC AFCKGA?J DMPKSJ? @SR BGDDCP GL RFC NJ?ACKCLR (QRPSARSPC) MD RFCGP ?RMKQ ?LB/MP AFCKGA?J @MLBQ ?PC ILMUL ?Q isomers. Structural isomers (JGIC @SR?LC ?LB GQM@SRCLC QFMUL GL Figure 2.24a) BGDDCP GL RFC NJ?ACKCLR MD RFCGP AMT?JCLR @MLBQ: @MRF KMJCASJCQ F?TC DMSP A?P@MLQ ?LB RCL FWBPMECLQ ($4)10), @SR RFC BGDDCPCLR ?PP?LECKCLR MD RFC ?RMKQ UGRFGL RFC KMJCASJCQ JC?BQ RM BGDDCPCLACQ GL RFCGP AFCKGA?J NPMNCPRGCQ. 'MP CV?KNJC, BSC RM RFCGP BGDDCPCLR AFCKGA?J NPMNCPRGCQ, @SR?LC GQ QSGRCB DMP SQC ?Q ? DSCJ DMP AGE?PCRRC JGEFRCPQ ?LB RMPAFCQ, UFCPC?Q GQM@SRCLC GQ QSGRCB DMP SQC ?Q ? PCDPGECP?LR ?LB ? NPMNCJJ?LR GL QNP?W A?LQ. Geometric isomers, ML RFC MRFCP F?LB, F?TC QGKGJ?P NJ?ACKCLRQ MD RFCGP AMT?JCLR @MLBQ @SR BGDDCP GL FMU RFCQC @MLBQ ?PC K?BC RM RFC QSPPMSLBGLE ?RMKQ, CQNCAG?JJW GL A?P@ML-RM-A?P@ML BMS@JC @MLBQ. *L RFC QGKNJC KMJCASJC @SRCLC ($4)8), RFC RUM KCRFWJ EPMSNQ ($)3) A?L @C ML CGRFCP QGBC MD RFC BMS@JC AMT?JCLR @MLB ACLRP?J RM RFC KMJCASJC, ?Q GJJSQRP?RCB GL Figure 2.24b. 8FCL RFC A?P@MLQ ?PC @MSLB ML RFC Q?KC QGBC MD RFC BMS@JC @MLB, RFGQ GQ RFC *0: AMLDGESP?RGML; GD RFCW ?PC ML MNNMQGRC QGBCQ MD RFC BMS@JC @MLB, GR GQ ? ;9(5: AMLDGESP?RGML. *L RFC ;9(5: AMLDGESP?RGML, RFC A?P@MLQ DMPK ? KMPC MP JCQQ JGLC?P QRPSARSPC, UFCPC?Q RFC A?P@MLQ GL RFC *0: AMLDGESP?RGML K?IC ? @CLB (AF?LEC GL BGPCARGML) MD RFC A?P@ML @?AI@MLC.
62
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!@>LI< 2.24 -LIB@RIBP QE>Q E>SB QEB P>JB KRJ?BO >KA QVMB LC >QLJP >OO>KDBA AFCCBOBKQIV >OB @>IIBA FPLJBOP. (>) 3QOR@QRO>I FPLJBOP E>SB > AFCCBOBKQ @LS>IBKQ >OO>KDBJBKQ LC >QLJP. (?) 'BLJBQOF@ FPLJBOP E>SB > AFCCBOBKQ >OO>KDBJBKQ LC >QLJP >OLRKA > ALR?IB ?LKA. (@) %K>KQFLJBOP >OB JFOOLO FJ>DBP LC B>@E LQEBO.
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igure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63
?RMKQ ?PC A?JJCB Q?RSP?RCB D?RQ, KC?LGLE RF?R RFCW AMLR?GL ?JJ RFC FWBPMECL ?RMKQ ?T?GJ?@JC. 4?RSP?RCB D?RQ ?PC ? QMJGB ?R PMMK RCKNCP?RSPC ?LB SQS?JJW MD ?LGK?J MPGEGL.
!@>LI< 2.25 4EBPB PM>@B-CFIIFKD JLABIP PELT > *0: (LIBF@ >@FA) >KA > ;9(5: (BIF>AF@ >@FA) C>QQV >@FA. .LQF@B QEB ?BKA FK QEB JLIB@RIB @>RPB ?V QEB *0: @LKCFDRO>QFLK.
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Enantiomers ?PC KMJCASJCQ RF?R QF?PC RFC Q?KC AFCKGA?J QRPSARSPC ?LB AFCKGA?J @MLBQ @SR BGDDCP GL RFC RFPCC-BGKCLQGML?J NJ?ACKCLR MD ?RMKQ QM RF?R RFCW ?PC KGPPMP GK?ECQ. "Q QFMUL GL Figure 2.26, ?L ?KGLM ?AGB ?J?LGLC CV?KNJC, RFC RUM QRPSARSPCQ ?PC LML-QSNCPGKNMQ?@JC. *L L?RSPC, MLJW RFC --DMPKQ MD ?KGLM ?AGBQ ?PC SQCB RM K?IC NPMRCGLQ. 4MKC % DMPKQ MD ?KGLM ?AGBQ ?PC QCCL GL RFC ACJJ U?JJQ MD @?ARCPG?, @SR LCTCP GL RFCGP NPMRCGLQ. 4GKGJ?PJW, RFC %-DMPK MD EJSAMQC GQ RFC K?GL NPMBSAR MD NFMRMQWLRFCQGQ ?LB RFC --DMPK MD RFC KMJCASJC GQ P?PCJW QCCL GL L?RSPC.
!@>LI< 2.26 $->I>KFKB >KA ,->I>KFKB >OB BU>JMIBP LC BK>KQFLJBOP LO JFOOLO FJ>DBP. /KIV QEB ,CLOJP LC >JFKL >@FAP >OB RPBA QL J>HB MOLQBFKP.
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Functional groups ?PC EPMSNQ MD ?RMKQ RF?R MAASP UGRFGL KMJCASJCQ ?LB AMLDCP QNCAGDGA AFCKGA?J NPMNCPRGCQ RM RFMQC KMJCASJCQ. 5FCW ?PC DMSLB ?JMLE RFC tA?P@ML @?AI@MLCu MD K?APMKMJCASJCQ. 5FGQ A?P@ML @?AI@MLC GQ DMPKCB @W AF?GLQ ?LB/MP PGLEQ MD A?P@ML ?RMKQ UGRF RFC MAA?QGML?J QS@QRGRSRGML MD ?L CJCKCLR QSAF ?Q LGRPMECL MP MVWECL. .MJCASJCQ UGRF MRFCP CJCKCLRQ GL RFCGP A?P@ML @?AI@MLC ?PC substituted hydrocarbons. 5FC DSLARGML?J EPMSNQ GL ? K?APMKMJCASJC ?PC SQS?JJW ?RR?AFCB RM RFC A?P@ML @?AI@MLC ?R MLC MP QCTCP?J BGDDCPCLR NJ?ACQ ?JMLE GRQ AF?GL ?LB/MP PGLE QRPSARSPC. &?AF MD RFC DMSP RWNCQ MD K?APMKMJCASJCQqNPMRCGLQ, JGNGBQ, A?P@MFWBP?RCQ, ?LB LSAJCGA ?AGBQqF?Q GRQ MUL AF?P?ARCPGQRGA QCR MD DSLARGML?J EPMSNQ RF?R AMLRPG@SRCQ EPC?RJW RM GRQ BGDDCPGLE AFCKGA?J NPMNCPRGCQ ?LB GRQ DSLARGML GL JGTGLE MPE?LGQKQ. " DSLARGML?J EPMSN A?L N?PRGAGN?RC GL QNCAGDGA AFCKGA?J PC?ARGMLQ. 4MKC MD RFC GKNMPR?LR DSLARGML?J EPMSNQ GL @GMJMEGA?J KMJCASJCQ ?PC QFMUL GL Figure 2.27; RFCW GLAJSBC: FWBPMVWJ, KCRFWJ, A?P@MLWJ, A?P@MVWJ, ?KGLM, NFMQNF?RC, ?LB QSJDFWBPWJ. 5FCQC EPMSNQ NJ?W ?L GKNMPR?LR PMJC GL RFC DMPK?RGML MD KMJCASJCQ JGIC %/", NPMRCGLQ, A?P@MFWBP?RCQ, ?LB JGNGBQ. 'SLARGML?J EPMSNQ ?PC SQS?JJW AJ?QQGDGCB ?Q
64
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
!@>LI< 2.27 4EB CRK@QFLK>I DOLRMP PELTK EBOB >OB CLRKA FK J>KV AFCCBOBKQ ?FLILDF@>I JLIB@RIBP.
)WBPMECL @MLBQ @CRUCCL DSLARGML?J EPMSNQ (UGRFGL RFC Q?KC KMJCASJC MP @CRUCCL BGDDCPCLR KMJCASJCQ) ?PC GKNMPR?LR RM RFC DSLARGML MD K?LW K?APMKMJCASJCQ ?LB FCJN RFCK RM DMJB NPMNCPJW GLRM ?LB K?GLR?GL RFC ?NNPMNPG?RC QF?NC DMP DSLARGMLGLE. )WBPMECL @MLBQ ?PC ?JQM GLTMJTCB GL T?PGMSQ PCAMELGRGML NPMACQQCQ, QSAF ?Q %/" AMKNJCKCLR?PW @?QC N?GPGLE ?LB RFC @GLBGLE MD ?L CLXWKC RM GRQ QS@QRP?RC, ?Q GJJSQRP?RCB GL Figure 2.28.
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65
!@>LI< 2.28 (VAOLDBK ?LKAP @LKKB@Q QTL PQO>KAP LC $.! QLDBQEBO QL @OB>QB QEB ALR?IB-EBIFU PQOR@QROB.
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sQ GQMRMNCQ 8KFD@: ELD9<I RMR?J LSK@CP MD NPMRMLQ GL ?L ?RMK 98C8E:<; :?<D@:8C <HL8K@FE QR?RCKCLR MD ? AFCKGA?J PC?ARGML UGRF RFC LSK@CP MD C?AF RWNC MD ?RMK COS?JGXCB DMP @MRF RFC NPMBSARQ ?LB PC?AR?LRQ 98J< KMJCASJC RF?R BML?RCQ FWBPMVGBC GMLQ MP MRFCPUGQC @GLBQ CVACQQ FWBPMECL GMLQ ?LB BCAPC?QCQ RFC AMLACLRP?RGML MD FWBPMECL GMLQ GL ? QMJSRGML 9L==<I QS@QR?LAC RF?R NPCTCLRQ ? AF?LEC GL N) @W ?@QMP@GLE MP PCJC?QGLE FWBPMECL MP FWBPMVGBC GMLQ :8CFI@< ?KMSLR MD FC?R PCOSGPCB RM AF?LEC RFC RCKNCP?RSPC MD MLC EP?K MD U?RCP @W MLC BCEPCC $CJQGSQ :8G@CC8IP 8:K@FE MAASPQ @CA?SQC U?RCP KMJCASJCQ ?PC ?RRP?ARCB RM AF?PECQ ML RFC GLLCP QSPD?ACQ MD L?PPMU RS@SJ?P QRPSARSPCQ QSAF ?Q EJ?QQ RS@CQ, BP?UGLE RFC U?RCP KMJCASJCQ RM RFC QGBCQ MD RFC RS@CQ :8K@FE NMQGRGTC GML RF?R GQ DMPKCB @W ?L ?RMK JMQGLE MLC MP KMPC CJCARPMLQ :?<D@:8C 9FE; GLRCP?ARGML @CRUCCL RUM MP KMPC MD RFC Q?KC MP BGDDCPCLR ?RMKQ RF?R PCQSJRQ GL RFC DMPK?RGML MD KMJCASJCQ :?<D@:8C I<8:K@FE NPMACQQ JC?BGLE RM RFC PC?PP?LECKCLR MD ?RMKQ GL KMJCASJCQ :?<D@:8C I<8:K@M@KP RFC ?@GJGRW RM AMK@GLC ?LB RM AFCKGA?JJW @MLB UGRF C?AF MRFCP :F?<J@FE GLRCPKMJCASJ?P DMPACQ @CRUCCL U?RCP KMJCASJCQ A?SQCB @W RFC NMJ?P L?RSPC MD U?RCP; PCQNMLQG@JC DMP QSPD?AC RCLQGML :FDGFLE; QS@QR?LAC AMKNMQCB MD KMJCASJCQ AMLQGQRGLE MD ?RMKQ MD ?R JC?QR RUM BGDDCPCLR CJCKCLRQ :FM8C<EK 9FE; RWNC MD QRPMLE @MLB DMPKCB @CRUCCL RUM MD RFC Q?KC MP BGDDCPCLR CJCKCLRQ; DMPKQ UFCL CJCARPMLQ ?PC QF?PCB @CRUCCL ?RMKQ ;@JJF:@8K@FE PCJC?QC MD ?L GML DPMK ? KMJCASJC QSAF RF?R RFC MPGEGL?J KMJCASJC LMU AMLQGQRQ MD ?L GML ?LB RFC AF?PECB PCK?GLQ MD RFC MPGEGL?J, QSAF ?Q UFCL U?RCP BGQQMAG?RCQ GLRM )+ ?LB 0)<C<:KIFCPK< GML LCACQQ?PW DMP LCPTC GKNSJQC AMLBSARGML, KSQAJC AMLRP?ARGMLQ ?LB U?RCP @?J?LAC <C<:KIFE :FE=@>LI8K@FE ?PP?LECKCLR MD CJCARPMLQ GL ?L ?RMKs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p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Carbohydrates A?L @C PCNPCQCLRCB @W RFC QRMGAFGMKCRPGA DMPKSJ? ($)20)5, UFCPC L GQ RFC LSK@CP MD A?P@MLQ GL RFC KMJCASJC. *L MRFCP UMPBQ, RFC P?RGM MD A?P@ML RM FWBPMECL RM MVWECL GQ 1:2:1 GL A?P@MFWBP?RC KMJCASJCQ. 5FGQ DMPKSJ? ?JQM CVNJ?GLQ RFC MPGEGL MD RFC RCPK tA?P@MFWBP?RCu: RFC AMKNMLCLRQ ?PC A?P@ML (tA?P@Mu) ?LB RFC AMKNMLCLRQ MD U?RCP (FCLAC, tFWBP?RCu). $?P@MFWBP?RCQ ?PC AJ?QQGDGCB GLRM RFPCC QS@RWNCQ: KMLMQ?AAF?PGBCQ, BGQ?AAF?PGBCQ, ?LB NMJWQ?AAF?PGBCQ. M4348')).'7/*+8 Monosaccharides (KMLM- = tMLCu; Q?AAF?P- = tQUCCRu) ?PC QGKNJC QSE?PQ, RFC KMQR AMKKML MD UFGAF GQ EJSAMQC. *L KMLMQ?AAF?PGBCQ, RFC LSK@CP MD A?P@MLQ SQS?JJW P?LECQ DPMK RFPCC RM QCTCL. .MQR KMLMQ?AAF?PGBC L?KCQ CLB UGRF RFC QSDDGV -MQC. *D RFC QSE?P F?Q ?L ?JBCFWBC EPMSN (RFC DSLARGML?J EPMSN UGRF RFC QRPSARSPC 3-$)0), GR GQ ILMUL ?Q ?L ?JBMQC, ?LB GD GR F?Q ? ICRMLC EPMSN (RFC DSLARGML?J EPMSN UGRF RFC QRPSARSPC 3$(=0)3'), GR GQ ILMUL ?Q ? ICRMQC. %CNCLBGLE ML RFC LSK@CP MD A?P@MLQ GL RFC QSE?P, RFCW ?JQM K?W @C ILMUL ?Q RPGMQCQ (RFPCC A?P@MLQ), NCLRMQCQ (DGTC A?P@MLQ), ?LB MP FCVMQCQ (QGV A?P@MLQ). 4CC Figure 3.4 DMP ?L GJJSQRP?RGML MD RFC KMLMQ?AAF?PGBCQ.
74
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igure 3.5).
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75
!@>LI< 3.5 'IR@LPB, D>I>@QLPB, >KA COR@QLPB >OB >II EBULPBP. 4EBV >OB PQOR@QRO>I FPLJBOP, JB>KFKD QEBV E>SB QEB P>JB @EBJF@>I CLOJRI> (#6(12/6) ?RQ > AFCCBOBKQ >OO>KDBJBKQ LC >QLJP.
7E>Q HFKA LC PRD>OP >OB QEBPB, >IALPB LO HBQLPB (JSAMQC, E?J?ARMQC, ?LB DPSARMQC ?PC GQMKCPGA KMLMQ?AAF?PGBCQ (FCVMQCQ), KC?LGLE RFCW F?TC RFC Q?KC AFCKGA?J DMPKSJ? @SR F?TC QJGEFRJW BGDDCPCLR QRPSARSPCQ. (JSAMQC ?LB E?J?ARMQC ?PC ?JBMQCQ, ?LB DPSARMQC GQ ? ICRMQC. .MLMQ?AAF?PGBCQ A?L CVGQR ?Q ? JGLC?P AF?GL MP ?Q PGLE-QF?NCB KMJCASJCQ; GL ?OSCMSQ QMJSRGMLQ RFCW ?PC SQS?JJW DMSLB GL PGLE DMPKQ (Figure 3.6). (JSAMQC GL ? PGLE DMPK A?L F?TC RUM BGDDCPCLR ?PP?LECKCLRQ MD RFC FWBPMVWJ EPMSN (0)) ?PMSLB RFC ?LMKCPGA A?P@ML (A?P@ML 1 RF?R @CAMKCQ ?QWKKCRPGA GL RFC NPMACQQ MD PGLE DMPK?RGML). *D RFC FWBPMVWJ EPMSN GQ @CJMU A?P@ML LSK@CP 1 GL RFC QSE?P, GR GQ Q?GB RM @C GL RFC ?JNF? (C) NMQGRGML, ?LB GD GR GQ ?@MTC RFC NJ?LC, GR GQ Q?GB RM @C GL RFC @CR? (D) NMQGRGML.
76
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!@>LI< 3.6 &FSB >KA PFU @>O?LK JLKLP>@@E>OFABP BUFPQ FK BNRFIF?OFRJ ?BQTBBK IFKB>O >KA OFKD CLOJP. 7EBK QEB OFKD CLOJP, QEB PFAB @E>FK FQ @ILPBP LK FP IL@HBA FKQL >K C LO D MLPFQFLK. &OR@QLPB >KA OF?LPB >IPL CLOJ OFKDP, >IQELRDE QEBV CLOJ CFSB-JBJ?BOBA OFKDP >P LMMLPBA QL QEB PFU-JBJ?BOBA OFKD LC DIR@LPB.
D/8')).'7/*+8 Disaccharides (BG- = tRUMu) DMPK UFCL RUM KMLMQ?AAF?PGBCQ SLBCPEM ? BCFWBP?RGML PC?ARGML (?JQM ILMUL ?Q ? AMLBCLQ?RGML PC?ARGML MP BCFWBP?RGML QWLRFCQGQ). %SPGLE RFGQ NPMACQQ, RFC FWBPMVWJ EPMSN MD MLC KMLMQ?AAF?PGBC AMK@GLCQ UGRF RFC FWBPMECL MD ?LMRFCP KMLMQ?AAF?PGBC, PCJC?QGLE ? KMJCASJC MD U?RCP ?LB DMPKGLE ? AMT?JCLR @MLB. " AMT?JCLR @MLB DMPKCB @CRUCCL ? A?P@MFWBP?RC KMJCASJC ?LB ?LMRFCP KMJCASJC (GL RFGQ A?QC, @CRUCCL RUM KMLMQ?AAF?PGBCQ) GQ ILMUL ?Q ? glycosidic bond (Figure 3.7). (JWAMQGBGA @MLBQ (?JQM A?JJCB EJWAMQGBGA JGLI?ECQ) A?L @C MD RFC ?JNF? MP RFC @CR? RWNC.
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77
!@>LI< 3.7 3R@OLPB FP CLOJBA TEBK > JLKLJBO LC DIR@LPB >KA > JLKLJBO LC COR@QLPB >OB GLFKBA FK > ABEVAO>QFLK OB>@QFLK QL CLOJ > DIV@LPFAF@ ?LKA. )K QEB MOL@BPP, > T>QBO JLIB@RIB FP ILPQ. "V @LKSBKQFLK, QEB @>O?LK >QLJP FK > JLKLP>@@E>OFAB >OB KRJ?BOBA COLJ QEB QBOJFK>I @>O?LK @ILPBPQ QL QEB @>O?LKVI DOLRM. )K PR@OLPB, > DIV@LPFAF@ IFKH>DB FP CLOJBA ?BQTBBK @>O?LK 1 FK DIR@LPB >KA @>O?LK 2 FK COR@QLPB.
$MKKML BGQ?AAF?PGBCQ GLAJSBC J?ARMQC, K?JRMQC, ?LB QSAPMQC (Figure 3.8). -?ARMQC GQ ? BGQ?AAF?PGBC AMLQGQRGLE MD RFC KMLMKCPQ EJSAMQC ?LB E?J?ARMQC. *R GQ DMSLB L?RSP?JJW GL KGJI. .?JRMQC, MP K?JR QSE?P, GQ ? BGQ?AAF?PGBC DMPKCB @W ? BCFWBP?RGML PC?ARGML @CRUCCL RUM EJSAMQC KMJCASJCQ. 5FC KMQR AMKKML BGQ?AAF?PGBC GQ QSAPMQC, MP R?@JC QSE?P, UFGAF GQ AMKNMQCB MD RFC KMLMKCPQ EJSAMQC ?LB DPSARMQC.
78
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!@>LI< 3.8 #LJJLK AFP>@@E>OFABP FK@IRAB J>IQLPB (DO>FK PRD>O), I>@QLPB (JFIH PRD>O), >KA PR@OLPB (Q>?IB PRD>O).
P41>8')).'7/*+8 " JMLE AF?GL MD KMLMQ?AAF?PGBCQ JGLICB @W EJWAMQGBGA @MLBQ GQ ILMUL ?Q ? polysaccharide (NMJW= tK?LWu). 5FC AF?GL K?W @C @P?LAFCB MP SL@P?LAFCB, ?LB GR K?W AMLR?GL BGDDCPCLR RWNCQ MD KMLMQ?AAF?PGBCQ. 5FC KMJCASJ?P UCGEFR K?W @C 100,000 B?JRMLQ MP KMPC BCNCLBGLE ML RFC LSK@CP MD KMLMKCPQ HMGLCB. 4R?PAF, EJWAMECL, ACJJSJMQC, ?LB AFGRGL ?PC NPGK?PW CV?KNJCQ MD NMJWQ?AAF?PGBCQ. 4R?PAF GQ RFC QRMPCB DMPK MD QSE?PQ GL NJ?LRQ ?LB GQ K?BC SN MD ? KGVRSPC MD ?KWJMQC ?LB ?KWJMNCARGL (@MRF NMJWKCPQ MD EJSAMQC). 1J?LRQ ?PC ?@JC RM QWLRFCQGXC EJSAMQC, ?LB RFC CVACQQ EJSAMQC, @CWMLB RFC NJ?LRs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igure 3.9, ?KWJMQC GQ QR?PAF DMPKCB @W SL@P?LAFCB AF?GLQ MD EJSAMQC KMLMKCPQ (MLJW C 1-4 JGLI?ECQ), UFCPC?Q ?KWJMNCARGL GQ ? @P?LAFCB NMJWQ?AAF?PGBC (C 1-6 JGLI?ECQ ?R RFC @P?LAF NMGLRQ).
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79
!@>LI< 3.9 !JVILPB >KA >JVILMB@QFK >OB QTL AFCCBOBKQ CLOJP LC PQ>O@E. !JVILPB FP @LJMLPBA LC RK?O>K@EBA @E>FKP LC DIR@LPB JLKLJBOP @LKKB@QBA ?V C 1,4 DIV@LPFAF@ IFKH>DBP. !JVILMB@QFK FP @LJMLPBA LC ?O>K@EBA @E>FKP LC DIR@LPB JLKLJBOP @LKKB@QBA ?V C 1,4 >KA C 1,6 DIV@LPFAF@ IFKH>DBP. "B@>RPB LC QEB T>V QEB PR?RKFQP >OB GLFKBA, QEB DIR@LPB @E>FKP E>SB > EBIF@>I PQOR@QROB. 'IV@LDBK (KLQ PELTK) FP PFJFI>O FK PQOR@QROB QL >JVILMB@QFK ?RQ JLOB EFDEIV ?O>K@EBA.
Glycogen GQ RFC QRMP?EC DMPK MD EJSAMQC GL FSK?LQ ?LB MRFCP TCPRC@P?RCQ ?LB GQ K?BC SN MD KMLMKCPQ MD EJSAMQC. (JWAMECL GQ RFC ?LGK?J COSGT?JCLR MD QR?PAF ?LB GQ ? FGEFJW @P?LAFCB KMJCASJC SQS?JJW QRMPCB GL JGTCP ?LB KSQAJC ACJJQ. 8FCLCTCP @JMMB EJSAMQC JCTCJQ BCAPC?QC, EJWAMECL GQ @PMICL BMUL RM PCJC?QC EJSAMQC GL ? NPMACQQ ILMUL ?Q EJWAMECLMJWQGQ. Cellulose GQ RFC KMQR ?@SLB?LR L?RSP?J @GMNMJWKCP. 5FC ACJJ U?JJ MD NJ?LRQ GQ KMQRJW K?BC MD ACJJSJMQC; RFGQ NPMTGBCQ QRPSARSP?J QSNNMPR RM RFC ACJJ. 8MMB ?LB N?NCP ?PC KMQRJW ACJJSJMQGA GL L?RSPC. $CJJSJMQC GQ K?BC SN MD EJSAMQC KMLMKCPQ RF?R ?PC JGLICB @W D 1-4 EJWAMQGBGA @MLBQ (Figure 3.10).
80
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!@>LI< 3.10 )K @BIIRILPB, DIR@LPB JLKLJBOP >OB IFKHBA FK RK?O>K@EBA @E>FKP ?V D 1-4 DIV@LPFAF@ IFKH>DBP. "B@>RPB LC QEB T>V QEB DIR@LPB PR?RKFQP >OB GLFKBA, BSBOV DIR@LPB JLKLJBO FP CIFMMBA OBI>QFSB QL QEB KBUQ LKB OBPRIQFKD FK > IFKB>O, CF?OLRP PQOR@QROB.
"Q QFMUL GL Figure 3.10, CTCPW MRFCP EJSAMQC KMLMKCP GL ACJJSJMQC GQ DJGNNCB MTCP, ?LB RFC KMLMKCPQ ?PC N?AICB RGEFRJW ?Q CVRCLBCB JMLE AF?GLQ. 5FGQ EGTCQ ACJJSJMQC GRQ PGEGBGRW ?LB FGEF RCLQGJC QRPCLERFq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igure 3.11). 5FGQ CVMQICJCRML GQ K?BC MD RFC @GMJMEGA?J K?APMKMJCASJC chitin, UFGAF GQ ? NMJWQ?AAF?PGBC-AMLR?GLGLE LGRPMECL. *R GQ K?BC MD PCNC?RGLE SLGRQ MD /-?ACRWJ-D-B-EJSAMQ?KGLC, ? KMBGDGCB QSE?P. $FGRGL GQ ?JQM ? K?HMP AMKNMLCLR MD DSLE?J ACJJ U?JJQ; DSLEG ?PC LCGRFCP ?LGK?JQ LMP NJ?LRQ ?LB DMPK ? IGLEBMK MD RFCGP MUL GL RFC BMK?GL &SI?PW?.
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"PC A?P@MFWBP?RCQ EMMB DMP WMS 1CMNJC UFM UGQF RM JMQC UCGEFR ?PC MDRCL RMJB RF?R A?P@MFWBP?RCQ ?PC @?B DMP RFCK ?LB QFMSJB @C ?TMGBCB. 4MKC BGCRQ AMKNJCRCJW DMP@GB A?P@MFWBP?RC AMLQSKNRGML, AJ?GKGLE RF?R ? JMU-A?P@MFWBP?RC BGCR FCJNQ NCMNJC RM JMQC UCGEFR D?QRCP. )MUCTCP, A?P@MFWBP?RCQ F?TC @CCL ?L GKNMPR?LR N?PR MD RFC FSK?L BGCR DMP RFMSQ?LBQ MD WC?PQ; ?PRGD?ARQ DPMK ?LAGCLR AGTGJGX?RGMLQ QFMU RFC NPCQCLAC MD UFC?R, PGAC, ?LB AMPL GL MSP ?LACQRMPQs QRMP?EC ?PC?Q. $?P@MFWBP?RCQ QFMSJB @C QSNNJCKCLRCB UGRF NPMRCGLQ, TGR?KGLQ, ?LB D?RQ RM @C N?PRQ MD ? UCJJ@?J?LACB BGCR. $?JMPGC-UGQC, ? EP?K MD A?P@MFWBP?RC NPMTGBCQ 4.3 ,A?J. 'MP AMKN?PGQML, D?RQ NPMTGBC 9 ,A?J/E, ? JCQQ BCQGP?@JC P?RGM. $?P@MFWBP?RCQ AMLR?GL QMJS@JC ?LB GLQMJS@JC CJCKCLRQ; RFC GLQMJS@JC N?PR GQ ILMUL ?Q DG@CP, UFGAF GQ KMQRJW ACJJSJMQC. 'G@CP F?Q K?LW SQCQ; GR NPMKMRCQ PCESJ?P @MUCJ KMTCKCLR @W ?BBGLE @SJI, ?LB GR PCESJ?RCQ RFC P?RC MD AMLQSKNRGML MD @JMMB EJSAMQC. 'G@CP ?JQM FCJNQ RM PCKMTC CVACQQ AFMJCQRCPMJ DPMK RFC @MBW: DG@CP @GLBQ RM RFC AFMJCQRCPMJ GL RFC QK?JJ GLRCQRGLC, RFCL ?RR?AFCQ RM RFC AFMJCQRCPMJ ?LB NPCTCLRQ RFC AFMJCQRCPMJ N?PRGAJCQ DPMK CLRCPGLE RFC @JMMBQRPC?K, ?LB RFCL AFMJCQRCPMJ CVGRQ RFC @MBW TG? RFC DCACQ. 'G@CP-PGAF BGCRQ ?JQM F?TC ? NPMRCARGTC PMJC GL PCBSAGLE RFC MAASPPCLAC MD AMJML A?LACP. *L ?BBGRGML, ? KC?J AMLR?GLGLE UFMJC EP?GLQ ?LB TCECR?@JCQ EGTCQ ? DCCJGLE MD DSJJLCQQ. "Q ?L GKKCBG?RC QMSPAC MD CLCPEW, EJSAMQC GQ @PMICL BMUL BSPGLE RFC NPMACQQ MD ACJJSJ?P PCQNGP?RGML, UFGAF NPMBSACQ "51, RFC CLCPEW ASPPCLAW MD RFC ACJJ. 8GRFMSR RFC AMLQSKNRGML MD A?P@MFWBP?RCQ, RFC ?T?GJ?@GJGRW MD tGLQR?LR CLCPEWu UMSJB @C PCBSACB. &JGKGL?RGLE A?P@MFWBP?RCQ DPMK RFC BGCR GQ LMR RFC @CQR U?W RM JMQC UCGEFR. " JMU-A?JMPGC BGCR RF?R GQ PGAF GL UFMJC EP?GLQ, DPSGRQ, TCECR?@JCQ, ?LB JC?L KC?R, RMECRFCP UGRF NJCLRW MD CVCPAGQC ?LB NJCLRW MD U?RCP, GQ RFC KMPC QCLQG@JC U?W RM JMQC UCGEFR.
'MP ?L ?BBGRGML?J NCPQNCARGTC ML A?P@MFWBP?RCQ, CVNJMPC t#GMKMJCASJCQ: RFC $?P@MFWBP?RCQu RFPMSEF RFGQ interactive animation (http://openstaxcollege.org/l/carbohydrates) .
82
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC DMSP K?HMP RWNCQ MD JGNGBQ z &VNJ?GL RFC PMJC MD D?RQ GL QRMPGLE CLCPEW z %GDDCPCLRG?RC @CRUCCL Q?RSP?RCB ?LB SLQ?RSP?RCB D?RRW ?AGBQ z %CQAPG@C NFMQNFMJGNGBQ ?LB RFCGP PMJC GL ACJJQ z %CDGLC RFC @?QGA QRPSARSPC MD ? QRCPMGB ?LB QMKC DSLARGMLQ MD QRCPMGBQ z &VNJ?GL RFC FMU AFMJCQRCPMJ FCJNQ RM K?GLR?GL RFC DJSGB L?RSPC MD RFC NJ?QK? KCK@P?LC Lipids GLAJSBC ? BGTCPQC EPMSN MD AMKNMSLBQ RF?R ?PC J?PECJW LMLNMJ?P GL L?RSPC. 5FGQ GQ @CA?SQC RFCW ?PC FWBPMA?P@MLQ RF?R GLAJSBC KMQRJW LMLNMJ?P A?P@MLpA?P@ML MP A?P@MLpFWBPMECL @MLBQ. /ML-NMJ?P KMJCASJCQ ?PC FWBPMNFM@GA (tU?RCP DC?PGLEu), MP GLQMJS@JC GL U?RCP. -GNGBQ NCPDMPK K?LW BGDDCPCLR DSLARGMLQ GL ? ACJJ. $CJJQ QRMPC CLCPEW DMP JMLE-RCPK SQC GL RFC DMPK MD D?RQ. -GNGBQ ?JQM NPMTGBC GLQSJ?RGML DPMK RFC CLTGPMLKCLR DMP NJ?LRQ ?LB ?LGK?JQ (Figure 3.12). 'MP CV?KNJC, RFCW FCJN ICCN ?OS?RGA @GPBQ ?LB K?KK?JQ BPW UFCL DMPKGLE ? NPMRCARGTC J?WCP MTCP DSP MP DC?RFCPQ @CA?SQC MD RFCGP U?RCPPCNCJJ?LR FWBPMNFM@GA L?RSPC. -GNGBQ ?PC ?JQM RFC @SGJBGLE @JMAIQ MD K?LW FMPKMLCQ ?LB ?PC ?L GKNMPR?LR AMLQRGRSCLR MD ?JJ ACJJSJ?P KCK@P?LCQ. -GNGBQ GLAJSBC D?RQ, MGJQ, U?VCQ, NFMQNFMJGNGBQ, ?LB QRCPMGBQ.
!@>LI< 3.12 (VAOLMEL?F@ IFMFAP FK QEB CRO LC >NR>QF@ J>JJ>IP, PR@E >P QEFP OFSBO LQQBO, MOLQB@Q QEBJ COLJ QEB BIBJBKQP. (@OBAFQ: +BK "LPJ>)
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" D?R KMJCASJC AMLQGQRQ MD RUM K?GL AMKNMLCLRQqEJWACPMJ ?LB D?RRW ?AGBQ. (JWACPMJ GQ ?L MPE?LGA AMKNMSLB (?JAMFMJ) UGRF RFPCC A?P@MLQ, DGTC FWBPMECLQ, ?LB RFPCC FWBPMVWJ (0)) EPMSNQ. '?RRW ?AGBQ F?TC ? JMLE AF?GL MD FWBPMA?P@MLQ RM UFGAF ? A?P@MVWJ EPMSN GQ ?RR?AFCB, FCLAC RFC L?KC tD?RRW ?AGB.u 5FC LSK@CP MD A?P@MLQ GL RFC D?RRW ?AGB K?W P?LEC DPMK 4 RM 36; KMQR AMKKML ?PC RFMQC AMLR?GLGLE 12p18 A?P@MLQ. *L ? D?R KMJCASJC, RFC D?RRW ?AGBQ ?PC ?RR?AFCB RM C?AF MD RFC RFPCC A?P@MLQ MD RFC EJWACPMJ KMJCASJC UGRF ?L CQRCP @MLB RFPMSEF ?L MVWECL ?RMK (Figure 3.13).
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83
!@>LI< 3.13 4OF>@VIDIV@BOLI FP CLOJBA ?V QEB GLFKFKD LC QEOBB C>QQV >@FAP QL > DIV@BOLI ?>@H?LKB FK > ABEVAO>QFLK OB>@QFLK. 4EOBB JLIB@RIBP LC T>QBO >OB OBIB>PBA FK QEB MOL@BPP.
%SPGLE RFGQ CQRCP @MLB DMPK?RGML, RFPCC U?RCP KMJCASJCQ ?PC PCJC?QCB. 5FC RFPCC D?RRW ?AGBQ GL RFC RPG?AWJEJWACPMJ K?W @C QGKGJ?P MP BGQQGKGJ?P. '?RQ ?PC ?JQM A?JJCB triacylglycerols MP triglycerides @CA?SQC MD RFCGP AFCKGA?J QRPSARSPC. 4MKC D?RRW ?AGBQ F?TC AMKKML L?KCQ RF?R QNCAGDW RFCGP MPGEGL. 'MP CV?KNJC, N?JKGRGA ?AGB, ? saturated fatty acid, GQ BCPGTCB DPMK RFC N?JK RPCC. "P?AFGBGA ?AGB GQ BCPGTCB DPMK A9(*/0: /@76.,(, RFC QAGCLRGDGA L?KC DMP EPMSLBLSRQ MP NC?LSRQ. '?RRW ?AGBQ K?W @C Q?RSP?RCB MP SLQ?RSP?RCB. *L ? D?RRW ?AGB AF?GL, GD RFCPC ?PC MLJW QGLEJC @MLBQ @CRUCCL LCGEF@MPGLE A?P@MLQ GL RFC FWBPMA?P@ML AF?GL, RFC D?RRW ?AGB GQ Q?GB RM @C Q?RSP?RCB. 4?RSP?RCB D?RRW ?AGBQ ?PC Q?RSP?RCB UGRF FWBPMECL; GL MRFCP UMPBQ, RFC LSK@CP MD FWBPMECL ?RMKQ ?RR?AFCB RM RFC A?P@ML QICJCRML GQ K?VGKGXCB. 4RC?PGA ?AGB GQ ?L CV?KNJC MD ? Q?RSP?RCB D?RRW ?AGB ( Figure 3.14)
!@>LI< 3.14 3QB>OF@ >@FA FP > @LJJLK P>QRO>QBA C>QQV >@FA.
8FCL RFC FWBPMA?P@ML AF?GL AMLR?GLQ ? BMS@JC @MLB, RFC D?RRW ?AGB GQ Q?GB RM @C unsaturated. 0JCGA ?AGB GQ ?L CV?KNJC MD ?L SLQ?RSP?RCB D?RRW ?AGB (Figure 3.15).
84
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!@>LI< 3.15 /IBF@ >@FA FP > @LJJLK RKP>QRO>QBA C>QQV >@FA.
.MQR SLQ?RSP?RCB D?RQ ?PC JGOSGB ?R PMMK RCKNCP?RSPC ?LB ?PC A?JJCB MGJQ. *D RFCPC GQ MLC BMS@JC @MLB GL RFC KMJCASJC, RFCL GR GQ ILMUL ?Q ? KMLMSLQ?RSP?RCB D?R (C.E., MJGTC MGJ), ?LB GD RFCPC GQ KMPC RF?L MLC BMS@JC @MLB, RFCL GR GQ ILMUL ?Q ? NMJWSLQ?RSP?RCB D?R (C.E., A?LMJ? MGJ). 8FCL ? D?RRW ?AGB F?Q LM BMS@JC @MLBQ, GR GQ ILMUL ?Q ? Q?RSP?RCB D?RRW ?AGB @CA?SQC LM KMPC FWBPMECL K?W @C ?BBCB RM RFC A?P@ML ?RMKQ MD RFC AF?GL. " D?R K?W AMLR?GL QGKGJ?P MP BGDDCPCLR D?RRW ?AGBQ ?RR?AFCB RM EJWACPMJ. -MLE QRP?GEFR D?RRW ?AGBQ UGRF QGLEJC @MLBQ RCLB RM ECR N?AICB RGEFRJW ?LB ?PC QMJGB ?R PMMK RCKNCP?RSPC. "LGK?J D?RQ UGRF QRC?PGA ?AGB ?LB N?JKGRGA ?AGB (AMKKML GL KC?R) ?LB RFC D?R UGRF @SRWPGA ?AGB (AMKKML GL @SRRCP) ?PC CV?KNJCQ MD Q?RSP?RCB D?RQ. .?KK?JQ QRMPC D?RQ GL QNCAG?JGXCB ACJJQ A?JJCB ?BGNMAWRCQ, UFCPC EJM@SJCQ MD D?R MAASNW KMQR MD RFC ACJJsQ TMJSKC. *L NJ?LRQ, D?R MP MGJ GQ QRMPCB GL K?LW QCCBQ ?LB GQ SQCB ?Q ? QMSPAC MD CLCPEW BSPGLE QCCBJGLE BCTCJMNKCLR. 6LQ?RSP?RCB D?RQ MP MGJQ ?PC SQS?JJW MD NJ?LR MPGEGL ?LB AMLR?GL *0: SLQ?RSP?RCB D?RRW ?AGBQ. C0: ?LB ;9(5: GLBGA?RC RFC AMLDGESP?RGML MD RFC KMJCASJC ?PMSLB RFC BMS@JC @MLB. *D FWBPMECLQ ?PC NPCQCLR GL RFC Q?KC NJ?LC, GR GQ PCDCPPCB RM ?Q ? AGQ D?R; GD RFC FWBPMECL ?RMKQ ?PC ML RUM BGDDCPCLR NJ?LCQ, GR GQ PCDCPPCB RM ?Q ? trans fat. 5FC *0: BMS@JC @MLB A?SQCQ ? @CLB MP ? tIGLIu RF?R NPCTCLRQ RFC D?RRW ?AGBQ DPMK N?AIGLE RGEFRJW, ICCNGLE RFCK JGOSGB ?R PMMK RCKNCP?RSPC (Figure 3.16). 0JGTC MGJ, AMPL MGJ, A?LMJ? MGJ, ?LB AMB JGTCP MGJ ?PC CV?KNJCQ MD SLQ?RSP?RCB D?RQ. 6LQ?RSP?RCB D?RQ FCJN RM JMUCP @JMMB AFMJCQRCPMJ JCTCJQ UFCPC?Q Q?RSP?RCB D?RQ AMLRPG@SRC RM NJ?OSC DMPK?RGML GL RFC ?PRCPGCQ.
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85
!@>LI< 3.16 3>QRO>QBA C>QQV >@FAP E>SB EVAOL@>O?LK @E>FKP @LKKB@QBA ?V PFKDIB ?LKAP LKIV. 5KP>QRO>QBA C>QQV >@FAP E>SB LKB LO JLOB ALR?IB ?LKAP. %>@E ALR?IB ?LKA J>V ?B FK > *0: LO ;9(5: @LKCFDRO>QFLK. )K QEB *0: @LKCFDRO>QFLK, ?LQE EVAOLDBKP >OB LK QEB P>JB PFAB LC QEB EVAOL@>O?LK @E>FK. )K QEB ;9(5: @LKCFDRO>QFLK, QEB EVAOLDBKP >OB LK LMMLPFQB PFABP. ! *0: ALR?IB ?LKA @>RPBP > HFKH FK QEB @E>FK.
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t@?Bu AFMJCQRCPMJ, UFGAF GL RSPL K?W JC?B RM NJ?OSC BCNMQGRGML GL RFC ?PRCPGCQ, PCQSJRGLE GL FC?PR BGQC?QC. .?LW D?QR DMMB PCQR?SP?LRQ F?TC PCACLRJW @?LLCB RFC SQC MD RP?LQ D?RQ, ?LB DMMB J?@CJQ ?PC PCOSGPCB RM BGQNJ?W RFC RP?LQ D?R AMLRCLR. O2+-' F'99> A)/*8 &QQCLRG?J D?RRW ?AGBQ ?PC D?RRW ?AGBQ PCOSGPCB @SR LMR QWLRFCQGXCB @W RFC FSK?L @MBW. $MLQCOSCLRJW, RFCW F?TC RM @C QSNNJCKCLRCB RFPMSEF GLECQRGML TG? RFC BGCR. Omega-3 D?RRW ?AGBQ (JGIC RF?R QFMUL GL Figure 3.17) D?JJ GLRM RFGQ A?RCEMPW ?LB ?PC MLC MD MLJW RUM ILMUL DMP FSK?LQ (RFC MRFCP @CGLE MKCE?-6 D?RRW ?AGB). 5FCQC ?PC NMJWSLQ?RSP?RCB D?RRW ?AGBQ ?LB ?PC A?JJCB MKCE?-3 @CA?SQC RFC RFGPB A?P@ML DPMK RFC CLB MD RFC FWBPMA?P@ML AF?GL GQ AMLLCARCB RM GRQ LCGEF@MPGLE A?P@ML @W ? BMS@JC @MLB.
86
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!@>LI< 3.17 !IME>-IFKLIBKF@ >@FA FP >K BU>JMIB LC >K LJBD>-3 C>QQV >@FA. )Q E>P QEOBB *0: ALR?IB ?LKAP >KA, >P > OBPRIQ, > @ROSBA PE>MB. &LO @I>OFQV, QEB @>O?LKP >OB KLQ PELTK. %>@E PFKDIV ?LKABA @>O?LK E>P QTL EVAOLDBKP >PPL@F>QBA TFQE FQ, >IPL KLQ PELTK.
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tD?RRWu DMMBQ JC?BQ RM UCGEFR E?GL. )MUCTCP, D?RQ BM F?TC GKNMPR?LR DSLARGMLQ. .?LW TGR?KGLQ ?PC D?R QMJS@JC, ?LB D?RQ QCPTC ?Q ? JMLE-RCPK QRMP?EC DMPK MD D?RRW ?AGBQ: ? QMSPAC MD CLCPEW. 5FCW ?JQM NPMTGBC GLQSJ?RGML DMP RFC @MBW. 5FCPCDMPC, tFC?JRFWu D?RQ GL KMBCP?RC ?KMSLRQ QFMSJB @C AMLQSKCB ML ? PCESJ?P @?QGQ.
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Wax AMTCPQ RFC DC?RFCPQ MD QMKC ?OS?RGA @GPBQ ?LB RFC JC?D QSPD?ACQ MD QMKC NJ?LRQ. #CA?SQC MD RFC FWBPMNFM@GA L?RSPC MD U?VCQ, RFCW NPCTCLR U?RCP DPMK QRGAIGLE ML RFC QSPD?AC (Figure 3.18). 8?VCQ ?PC K?BC SN MD JMLE D?RRW ?AGB AF?GLQ CQRCPGDGCB RM JMLE-AF?GL ?JAMFMJQ.
!@>LI< 3.18 7>UV @LSBOFKDP LK PLJB IB>SBP >OB J>AB LC IFMFAP. (@OBAFQ: 2LDBO 'OFCCFQE)
+?FJG?FC@G@;J
Phospholipids ?PC K?HMP AMLQRGRSCLRQ MD RFC NJ?QK? KCK@P?LC, RFC MSRCPKMQR J?WCP MD ?LGK?J ACJJQ. -GIC D?RQ, RFCW ?PC AMKNMQCB MD D?RRW ?AGB AF?GLQ ?RR?AFCB RM ? EJWACPMJ MP QNFGLEMQGLC @?AI@MLC. *LQRC?B MD RFPCC D?RRW ?AGBQ ?RR?AFCB ?Q GL RPGEJWACPGBCQ, FMUCTCP, RFCPC ?PC RUM D?RRW ?AGBQ DMPKGLE BG?AWJEJWACPMJ, ?LB RFC RFGPB A?P@ML MD RFC EJWACPMJ @?AI@MLC GQ MAASNGCB @W ? KMBGDGCB NFMQNF?RC EPMSN (Figure 3.19). " NFMQNF?RC EPMSN ?JMLC ?RR?AFCB RM ? BG?EJWACPMJ BMCQ LMR OS?JGDW ?Q ? NFMQNFMJGNGB; GR GQ NFMQNF?RGB?RC
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87
(BG?AWJEJWACPMJ 3-NFMQNF?RC), RFC NPCASPQMP MD NFMQNFMJGNGBQ. 5FC NFMQNF?RC EPMSN GQ KMBGDGCB @W ?L ?JAMFMJ. 1FMQNF?RGBWJAFMJGLC ?LB NFMQNF?RGBWJQCPGLC ?PC RUM GKNMPR?LR NFMQNFMJGNGBQ RF?R ?PC DMSLB GL NJ?QK? KCK@P?LCQ.
!@>LI< 3.19 ! MELPMELIFMFA FP > JLIB@RIB TFQE QTL C>QQV >@FAP >KA > JLAFCFBA MELPME>QB DOLRM >QQ>@EBA QL > DIV@BOLI ?>@H?LKB. 4EB MELPME>QB J>V ?B JLAFCFBA ?V QEB >AAFQFLK LC @E>ODBA LO MLI>O @EBJF@>I DOLRMP. 4TL @EBJF@>I DOLRMP QE>Q J>V JLAFCV QEB MELPME>QB, @ELIFKB >KA PBOFKB, >OB PELTK EBOB. "LQE @ELIFKB >KA PBOFKB >QQ>@E QL QEB MELPME>QB DOLRM >Q QEB MLPFQFLK I>?BIBA 2 SF> QEB EVAOLUVI DOLRM FKAF@>QBA FK DOBBK.
" NFMQNFMJGNGB GQ ?L ?KNFGN?RFGA KMJCASJC, KC?LGLE GR F?Q ? FWBPMNFM@GA ?LB ? FWBPMNFGJGA N?PR. 5FC D?RRW ?AGB AF?GLQ ?PC FWBPMNFM@GA ?LB A?LLMR GLRCP?AR UGRF U?RCP, UFCPC?Q RFC NFMQNF?RC-AMLR?GLGLE EPMSN GQ FWBPMNFGJGA ?LB GLRCP?ARQ UGRF U?RCP (Figure 3.20).
!@>LI< 3.20 4EB MELPMELIFMFA ?FI>VBO FP QEB J>GLO @LJMLKBKQ LC >II @BIIRI>O JBJ?O>KBP. 4EB EVAOLMEFIF@ EB>A DOLRMP LC QEB MELPMELIFMFAP C>@B QEB >NRBLRP PLIRQFLK. 4EB EVAOLMEL?F@ Q>FIP >OB PBNRBPQBOBA FK QEB JFAAIB LC QEB ?FI>VBO.
5FC FC?B GQ RFC FWBPMNFGJGA N?PR, ?LB RFC R?GJ AMLR?GLQ RFC FWBPMNFM@GA D?RRW ?AGBQ. *L ? KCK@P?LC, ? @GJ?WCP MD NFMQNFMJGNGBQ DMPKQ RFC K?RPGV MD RFC QRPSARSPC, RFC D?RRW ?AGB R?GJQ MD NFMQNFMJGNGBQ D?AC GLQGBC, ?U?W DPMK U?RCP, UFCPC?Q RFC NFMQNF?RC EPMSN D?ACQ RFC MSRQGBC, ?OSCMSQ QGBC (Figure 3.20). 1FMQNFMJGNGBQ ?PC PCQNMLQG@JC DMP RFC BWL?KGA L?RSPC MD RFC NJ?QK? KCK@P?LC. *D ? BPMN MD NFMQNFMJGNGBQ GQ NJ?ACB GL U?RCP, GR QNMLR?LCMSQJW DMPKQ ? QRPSARSPC ILMUL ?Q ? KGACJJC, UFCPC RFC FWBPMNFGJGA NFMQNF?RC FC?BQ D?AC RFC MSRQGBC ?LB RFC D?RRW ?AGBQ D?AC RFC GLRCPGMP MD RFGQ QRPSARSPC.
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6LJGIC RFC NFMQNFMJGNGBQ ?LB D?RQ BGQASQQCB C?PJGCP, steroids F?TC ? DSQCB PGLE QRPSARSPC. "JRFMSEF RFCW BM LMR PCQCK@JC RFC MRFCP JGNGBQ, RFCW ?PC EPMSNCB UGRF RFCK @CA?SQC RFCW ?PC ?JQM FWBPMNFM@GA?LB GLQMJS@JC GL U?RCP. "JJ QRCPMGBQ F?TC DMSP JGLICB A?P@ML PGLEQ ?LB QCTCP?J MD RFCK, JGIC AFMJCQRCPMJ, F?TC
88
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? QFMPR R?GJ (Figure 3.21). .?LW QRCPMGBQ ?JQM F?TC RFC p0) DSLARGML?J EPMSN, UFGAF NSRQ RFCK GL RFC ?JAMFMJ AJ?QQGDGA?RGML (QRCPMJQ).
!@>LI< 3.21 3QBOLFAP PR@E >P @ELIBPQBOLI >KA @LOQFPLI >OB @LJMLPBA LC CLRO CRPBA EVAOL@>O?LK OFKDP.
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
'MP ?L ?BBGRGML?J NCPQNCARGTC ML JGNGBQ, CVNJMPC RFC GLRCP?ARGTC ?LGK?RGML Biomolecules: The Lipids (http://openstaxcollege.org/l/lipids)
3.4 R +IFK<@EJ
#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC DSLARGMLQ NPMRCGLQ NCPDMPK GL RFC ACJJ ?LB GL RGQQSCQ z %GQASQQ RFC PCJ?RGMLQFGN @CRUCCL ?KGLM ?AGBQ ?LB NPMRCGLQ z &VNJ?GL RFC DMSP JCTCJQ MD NPMRCGL MPE?LGX?RGML z %CQAPG@C RFC U?WQ GL UFGAF NPMRCGL QF?NC ?LB DSLARGML ?PC JGLICB
#(!04%2 3 X ")/,/')#!, -!#2/-/,%#5,%3
89
Proteins ?PC MLC MD RFC KMQR ?@SLB?LR MPE?LGA KMJCASJCQ GL JGTGLE QWQRCKQ ?LB F?TC RFC KMQR BGTCPQC P?LEC MD DSLARGMLQ MD ?JJ K?APMKMJCASJCQ. 1PMRCGLQ K?W @C QRPSARSP?J, PCESJ?RMPW, AMLRP?ARGJC, MP NPMRCARGTC; RFCW K?W QCPTC GL RP?LQNMPR, QRMP?EC, MP KCK@P?LCQ; MP RFCW K?W @C RMVGLQ MP CLXWKCQ. &?AF ACJJ GL ? JGTGLE QWQRCK K?W AMLR?GL RFMSQ?LBQ MD NPMRCGLQ, C?AF UGRF ? SLGOSC DSLARGML. 5FCGP QRPSARSPCQ, JGIC RFCGP DSLARGMLQ, T?PW EPC?RJW. 5FCW ?PC ?JJ, FMUCTCP, NMJWKCPQ MD ?KGLM ?AGBQ, ?PP?LECB GL ? JGLC?P QCOSCLAC.
/PG<J 8E; !LE:K@FEJ F= +IFK<@EJ

Enzymes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ormones ?PC AFCKGA?J-QGEL?JGLE KMJCASJCQ, SQS?JJW QK?JJ NPMRCGLQ MP QRCPMGBQ, QCAPCRCB @W CLBMAPGLC ACJJQ RF?R ?AR RM AMLRPMJ MP PCESJ?RC QNCAGDGA NFWQGMJMEGA?J NPMACQQCQ, GLAJSBGLE EPMURF, BCTCJMNKCLR, KCR?@MJGQK, ?LB PCNPMBSARGML. 'MP CV?KNJC, GLQSJGL GQ ? NPMRCGL FMPKMLC RF?R FCJNQ RM PCESJ?RC RFC @JMMB EJSAMQC JCTCJ. 5FC NPGK?PW RWNCQ ?LB DSLARGMLQ MD NPMRCGLQ ?PC JGQRCB GL Table 3.1.
+IFK<@E /PG<J 8E; !LE:K@FEJ

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!JVI>PB, IFM>PB, MBMPFK, QOVMPFK (BJLDIL?FK, >I?RJFK !@QFK, QR?RIFK, HBO>QFK )KPRIFK, QEVOLUFKB )JJRKLDIL?RIFKP !@QFK, JVLPFK ,BDRJB PQLO>DB MOLQBFKP, BDD TEFQB (>I?RJFK)
!LE:K@FEJ
(BIM FK AFDBPQFLK LC CLLA ?V @>Q>?LIFWFKD KRQOFBKQP FKQL JLKLJBOF@ RKFQP #>OOV PR?PQ>K@BP FK QEB ?ILLA LO IVJME QEOLRDELRQ QEB ?LAV #LKPQOR@Q AFCCBOBKQ PQOR@QROBP, IFHB QEB @VQLPHBIBQLK #LLOAFK>QB QEB >@QFSFQV LC AFCCBOBKQ ?LAV PVPQBJP 0OLQB@Q QEB ?LAV COLJ CLOBFDK M>QELDBKP %CCB@Q JRP@IB @LKQO>@QFLK 0OLSFAB KLROFPEJBKQ FK B>OIV ABSBILMJBKQ LC QEB BJ?OVL >KA QEB PBBAIFKD
1PMRCGLQ F?TC BGDDCPCLR QF?NCQ ?LB KMJCASJ?P UCGEFRQ; QMKC NPMRCGLQ ?PC EJM@SJ?P GL QF?NC UFCPC?Q MRFCPQ ?PC DG@PMSQ GL L?RSPC. 'MP CV?KNJC, FCKMEJM@GL GQ ? EJM@SJ?P NPMRCGL, @SR AMJJ?ECL, DMSLB GL MSP QIGL, GQ ? DG@PMSQ NPMRCGL. 1PMRCGL QF?NC GQ APGRGA?J RM GRQ DSLARGML, ?LB RFGQ QF?NC GQ K?GLR?GLCB @W K?LW BGDDCPCLR RWNCQ MD AFCKGA?J @MLBQ. $F?LECQ GL RCKNCP?RSPC, N), ?LB CVNMQSPC RM AFCKGA?JQ K?W JC?B RM NCPK?LCLR AF?LECQ GL RFC QF?NC MD RFC NPMRCGL, JC?BGLE RM JMQQ MD DSLARGML, ILMUL ?Q denaturation. "JJ NPMRCGLQ ?PC K?BC SN MD BGDDCPCLR ?PP?LECKCLRQ MD RFC Q?KC 20 RWNCQ MD ?KGLM ?AGBQ.
AD@EF A:@;J
Amino acids ?PC RFC KMLMKCPQ RF?R K?IC SN NPMRCGLQ. &?AF ?KGLM ?AGB F?Q RFC Q?KC DSLB?KCLR?J QRPSARSPC, UFGAF AMLQGQRQ MD ? ACLRP?J A?P@ML ?RMK, ?JQM ILMUL ?Q RFC ?JNF? (C) A?P@ML, @MLBCB RM ?L ?KGLM EPMSN (/)2), ? A?P@MVWJ EPMSN ($00)), ?LB RM ? FWBPMECL ?RMK. &TCPW ?KGLM ?AGB ?JQM F?Q ?LMRFCP ?RMK MP EPMSN MD ?RMKQ @MLBCB RM RFC ACLRP?J ?RMK ILMUL ?Q RFC 3 EPMSN (Figure 3.22).
90
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!@>LI< 3.22 !JFKL >@FAP E>SB > @BKQO>I >PVJJBQOF@ @>O?LK QL TEF@E >K >JFKL DOLRM, > @>O?LUVI DOLRM, > EVAOLDBK >QLJ, >KA > PFAB @E>FK (2 DOLRM) >OB >QQ>@EBA.
5FC L?KC "?KGLM ?AGB" GQ BCPGTCB DPMK RFC D?AR RF?R RFCW AMLR?GL @MRF ?KGLM EPMSN ?LB A?P@MVWJ?AGB-EPMSN GL RFCGP @?QGA QRPSARSPC. "Q KCLRGMLCB, RFCPC ?PC 20 ?KGLM ?AGBQ NPCQCLR GL NPMRCGLQ. 5CL MD RFCQC ?PC AMLQGBCPCB CQQCLRG?J ?KGLM ?AGBQ GL FSK?LQ @CA?SQC RFC FSK?L @MBW A?LLMR NPMBSAC RFCK ?LB RFCW ?PC M@R?GLCB DPMK RFC BGCR. 'MP C?AF ?KGLM ?AGB, RFC 3 EPMSN (MP QGBC AF?GL) GQ BGDDCPCLR (Figure 3.23).
#(!04%2 3 X ")/,/')#!, -!#2/-/,%#5,%3
91
!@>LI< 3.23 4EBOB >OB 20 @LJJLK >JFKL >@FAP @LJJLKIV CLRKA FK MOLQBFKP, B>@E TFQE > AFCCBOBKQ 2 DOLRM (S>OF>KQ DOLRM) QE>Q ABQBOJFKBP FQP @EBJF@>I K>QROB.
7EF@E @>QBDLOFBP LC >JFKL >@FA TLRIA VLR BUMB@Q QL CFKA LK QEB PROC>@B LC > PLIR?IB MOLQBFK, >KA TEF@E TLRIA VLR BUMB@Q QL CFKA FK QEB FKQBOFLO 7E>Q AFPQOF?RQFLK LC >JFKL >@FAP TLRIA VLR BUMB@Q QL CFKA FK > MOLQBFK BJ?BAABA FK > IFMFA ?FI>VBO 5FC AFCKGA?J L?RSPC MD RFC QGBC AF?GL BCRCPKGLCQ RFC L?RSPC MD RFC ?KGLM ?AGB (RF?R GQ, UFCRFCP GR GQ ?AGBGA, @?QGA, NMJ?P, MP LMLNMJ?P). 'MP CV?KNJC, RFC ?KGLM ?AGB EJWAGLC F?Q ? FWBPMECL ?RMK ?Q RFC 3 EPMSN. "KGLM ?AGBQ QSAF ?Q T?JGLC, KCRFGMLGLC, ?LB ?J?LGLC ?PC LMLNMJ?P MP FWBPMNFM@GA GL L?RSPC, UFGJC ?KGLM ?AGBQ QSAF ?Q QCPGLC, RFPCMLGLC, ?LB AWQRCGLC ?PC NMJ?P ?LB F?TC FWBPMNFGJGA QGBC AF?GLQ. 5FC QGBC AF?GLQ MD JWQGLC ?LB ?PEGLGLC ?PC NMQGRGTCJW AF?PECB, ?LB RFCPCDMPC RFCQC ?KGLM ?AGBQ ?PC ?JQM ILMUL ?Q @?QGA ?KGLM ?AGBQ. 1PMJGLC F?Q ?L 3 EPMSN RF?R GQ JGLICB RM RFC ?KGLM EPMSN, DMPKGLE ? PGLE-JGIC QRPSARSPC. 1PMJGLC GQ ?L CVACNRGML RM RFC QR?LB?PB QRPSARSPC MD ?L ?LGKM ?AGB QGLAC GRQ ?KGLM EPMSN GQ LMR QCN?P?RC DPMK RFC QGBC AF?GL (Figure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peptide bond, UFGAF GQ DMPKCB @W ? BCFWBP?RGML PC?ARGML. 5FC A?P@MVWJ EPMSN MD MLC ?KGLM ?AGB ?LB RFC ?KGLM EPMSN MD RFC GLAMKGLE ?KGLM ?AGB AMK@GLC, PCJC?QGLE ? KMJCASJC MD U?RCP. 5FC PCQSJRGLE @MLB GQ RFC NCNRGBC @MLB (Figure 3.24).
92
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!@>LI< 3.24 0BMQFAB ?LKA CLOJ>QFLK FP > ABEVAO>QFLK PVKQEBPFP OB>@QFLK. 4EB @>O?LUVI DOLRM LC LKB >JFKL >@FA FP IFKHBA QL QEB >JFKL DOLRM LC QEB FK@LJFKD >JFKL >@FA. )K QEB MOL@BPP, > JLIB@RIB LC T>QBO FP OBIB>PBA.
5FC NPMBSARQ DMPKCB @W QSAF JGLI?ECQ ?PC A?JJCB NCNRGBCQ. "Q KMPC ?KGLM ?AGBQ HMGL RM RFGQ EPMUGLE AF?GL, RFC PCQSJRGLE AF?GL GQ ILMUL ?Q ? NMJWNCNRGBC. &?AF NMJWNCNRGBC F?Q ? DPCC ?KGLM EPMSN ?R MLC CLB. 5FGQ CLB GQ A?JJCB RFC / RCPKGL?J, MP RFC ?KGLM RCPKGL?J, ?LB RFC MRFCP CLB F?Q ? DPCC A?P@MVWJ EPMSN, ?JQM ILMUL ?Q RFC $ MP A?P@MVWJ RCPKGL?J. 8FGJC RFC RCPKQ NMJWNCNRGBC ?LB NPMRCGL ?PC QMKCRGKCQ SQCB GLRCPAF?LEC?@JW, ? NMJWNCNRGBC GQ RCAFLGA?JJW ? NMJWKCP MD ?KGLM ?AGBQ, UFCPC?Q RFC RCPK NPMRCGL GQ SQCB DMP ? NMJWNCNRGBC MP NMJWNCNRGBCQ RF?R F?TC AMK@GLCB RMECRFCP, MDRCL F?TC @MSLB LML-NCNRGBC NPMQRFCRGA EPMSNQ, F?TC ? BGQRGLAR QF?NC, ?LB F?TC ? SLGOSC DSLARGML. "DRCP NPMRCGL QWLRFCQGQ (RP?LQJ?RGML), KMQR NPMRCGLQ ?PC KMBGDGCB. 5FCQC ?PC ILMUL ?Q NMQR-RP?LQJ?RGML?J KMBGDGA?RGMLQ. 5FCW K?W SLBCPEM AJC?T?EC, NFMQNFMPWJ?RGML, MP K?W PCOSGPC RFC ?BBGRGML MD MRFCP AFCKGA?J EPMSNQ. 0LJW ?DRCP RFCQC KMBGDGA?RGMLQ GQ RFC NPMRCGL AMKNJCRCJW DSLARGML?J.
$JGAI RFPMSEF RFC QRCNQ MD NPMRCGL QWLRFCQGQ GL RFGQ interactive tutorial (http://openstaxcollege.org/l/ protein_synth) .
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93
/?< MFCLK@FE8IP .@>E@=@:8E:< F= CPKF:?IFD< :

#VQL@EOLJB @ FP >K FJMLOQ>KQ @LJMLKBKQ LC QEB BIB@QOLK QO>KPMLOQ @E>FK, > M>OQ LC @BIIRI>O OBPMFO>QFLK, >KA FQ FP KLOJ>IIV CLRKA FK QEB @BIIRI>O LOD>KBIIB, QEB JFQL@ELKAOFLK. 4EFP MOLQBFK E>P > EBJB MOLPQEBQF@ DOLRM, >KA QEB @BKQO>I FLK LC QEB EBJB DBQP >IQBOK>QBIV OBAR@BA >KA LUFAFWBA AROFKD BIB@QOLK QO>KPCBO. "B@>RPB QEFP BPPBKQF>I MOLQBFKwP OLIB FK MOLAR@FKD @BIIRI>O BKBODV FP @OR@F>I, FQ E>P @E>KDBA SBOV IFQQIB LSBO JFIIFLKP LC VB>OP. 0OLQBFK PBNRBK@FKD E>P PELTK QE>Q QEBOB FP > @LKPFABO>?IB >JLRKQ LC @VQL@EOLJB @ >JFKL >@FA PBNRBK@B ELJLILDV >JLKD AFCCBOBKQ PMB@FBP; FK LQEBO TLOAP, BSLIRQFLK>OV HFKPEFM @>K ?B >PPBPPBA ?V JB>PROFKD QEB PFJFI>OFQFBP LO AFCCBOBK@BP >JLKD S>OFLRP PMB@FBPw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
+IFK<@E .KIL:KLI<
"Q BGQASQQCB C?PJGCP, RFC QF?NC MD ? NPMRCGL GQ APGRGA?J RM GRQ DSLARGML. 'MP CV?KNJC, ?L CLXWKC A?L @GLB RM ? QNCAGDGA QS@QRP?RC ?R ? QGRC ILMUL ?Q RFC ?ARGTC QGRC. *D RFGQ ?ARGTC QGRC GQ ?JRCPCB @CA?SQC MD JMA?J AF?LECQ MP AF?LECQ GL MTCP?JJ NPMRCGL QRPSARSPC, RFC CLXWKC K?W @C SL?@JC RM @GLB RM RFC QS@QRP?RC. 5M SLBCPQR?LB FMU RFC NPMRCGL ECRQ GRQ DGL?J QF?NC MP AMLDMPK?RGML, UC LCCB RM SLBCPQR?LB RFC DMSP JCTCJQ MD NPMRCGL QRPSARSPC: NPGK?PW, QCAMLB?PW, RCPRG?PW, ?LB OS?RCPL?PW. P7/2'7> !97:)9:7+ 5FC SLGOSC QCOSCLAC MD ?KGLM ?AGBQ GL ? NMJWNCNRGBC AF?GL GQ GRQ primary structure. 'MP CV?KNJC, RFC N?LAPC?RGA FMPKMLC GLQSJGL F?Q RUM NMJWNCNRGBC AF?GLQ, " ?LB #, ?LB RFCW ?PC JGLICB RMECRFCP @W BGQSJDGBC @MLBQ. 5FC / RCPKGL?J ?KGLM ?AGB MD RFC " AF?GL GQ EJWAGLC, UFCPC?Q RFC $ RCPKGL?J ?KGLM ?AGB GQ ?QN?P?EGLC (Figure 3.25). 5FC QCOSCLACQ MD ?KGLM ?AGBQ GL RFC " ?LB # AF?GLQ ?PC SLGOSC RM GLQSJGL.
!@>LI< 3.25 "LSFKB PBORJ FKPRIFK FP > MOLQBFK ELOJLKB J>AB LC QTL MBMQFAB @E>FKP, ! (21 >JFKL >@FAP ILKD) >KA " (30 >JFKL >@FAP ILKD). )K B>@E @E>FK, MOFJ>OV PQOR@QROB FP FKAF@>QBA ?V QEOBB-IBQQBO >??OBSF>QFLKP QE>Q OBMOBPBKQ QEB K>JBP LC QEB >JFKL >@FAP FK QEB LOABO QEBV >OB MOBPBKQ. 4EB >JFKL >@FA @VPQBFKB (@VP) E>P > PRICEVAOVI (3() DOLRM >P > PFAB @E>FK. 4TL PRICEVAOVI DOLRMP @>K OB>@Q FK QEB MOBPBK@B LC LUVDBK QL CLOJ > AFPRICFAB (3-3) ?LKA. 4TL AFPRICFAB ?LKAP @LKKB@Q QEB ! >KA " @E>FKP QLDBQEBO, >KA > QEFOA EBIMP QEB ! @E>FK CLIA FKQL QEB @LOOB@Q PE>MB. .LQB QE>Q >II AFPRICFAB ?LKAP >OB QEB P>JB IBKDQE, ?RQ >OB AO>TK AFCCBOBKQ PFWBP CLO @I>OFQV.
5FC SLGOSC QCOSCLAC DMP CTCPW NPMRCGL GQ SJRGK?RCJW BCRCPKGLCB @W RFC ECLC CLAMBGLE RFC NPMRCGL. " AF?LEC GL LSAJCMRGBC QCOSCLAC MD RFC ECLCsQ AMBGLE PCEGML K?W JC?B RM ? BGDDCPCLR ?KGLM ?AGB @CGLE ?BBCB RM RFC EPMUGLE NMJWNCNRGBC AF?GL, A?SQGLE ? AF?LEC GL NPMRCGL QRPSARSPC ?LB DSLARGML. *L QGAIJC
94
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ACJJ ?LCKG?, RFC FCKMEJM@GL D AF?GL (? QK?JJ NMPRGML MD UFGAF GQ QFMUL GL Figure 3.26) F?Q ? QGLEJC ?KGLM ?AGB QS@QRGRSRGML, A?SQGLE ? AF?LEC GL NPMRCGL QRPSARSPC ?LB DSLARGML. 4NCAGDGA?JJW, RFC ?KGLM ?AGB EJSR?KGA ?AGB GQ QS@QRGRSRCB @W T?JGLC GL RFC D AF?GL. 8F?R GQ KMQR PCK?PI?@JC RM AMLQGBCP GQ RF?R ? FCKMEJM@GL KMJCASJC GQ K?BC SN MD RUM ?JNF? AF?GLQ ?LB RUM @CR? AF?GLQ RF?R C?AF AMLQGQR MD ?@MSR 150 ?KGLM ?AGBQ. 5FC KMJCASJC, RFCPCDMPC, F?Q ?@MSR 600 ?KGLM ?AGBQ. 5FC QRPSARSP?J BGDDCPCLAC @CRUCCL ? LMPK?J FCKMEJM@GL KMJCASJC ?LB ? QGAIJC ACJJ KMJCASJCqUFGAF BP?K?RGA?JJW BCAPC?QCQ JGDC CVNCAR?LAWqGQ ? QGLEJC ?KGLM ?AGB MD RFC 600. 8F?R GQ CTCL KMPC PCK?PI?@JC GQ RF?R RFMQC 600 ?KGLM ?AGBQ ?PC CLAMBCB @W RFPCC LSAJCMRGBCQ C?AF, ?LB RFC KSR?RGML GQ A?SQCB @W ? QGLEJC @?QC AF?LEC (NMGLR KSR?RGML), 1 GL 1800 @?QCQ.
!@>LI< 3.26 4EB ?BQ> @E>FK LC EBJLDIL?FK FP 147 OBPFARBP FK IBKDQE, VBQ > PFKDIB >JFKL >@FA PR?PQFQRQFLK IB>AP QL PF@HIB @BII >KBJF>. )K KLOJ>I EBJLDIL?FK, QEB >JFKL >@FA >Q MLPFQFLK PBSBK FP DIRQ>J>QB. )K PF@HIB @BII EBJLDIL?FK, QEFP DIRQ>J>QB FP OBMI>@BA ?V > S>IFKB.
#CA?SQC MD RFGQ AF?LEC MD MLC ?KGLM ?AGB GL RFC AF?GL, FCKMEJM@GL KMJCASJCQ DMPK JMLE DG@CPQ RF?R BGQRMPR RFC @GAMLA?TC, MP BGQA-QF?NCB, PCB @JMMB ACJJQ ?LB ?QQSKC ? APCQACLR MP tQGAIJCu QF?NC, UFGAF AJMEQ ?PRCPGCQ (Figure 3.27). 5FGQ A?L JC?B RM KWPG?B QCPGMSQ FC?JRF NPM@JCKQ QSAF ?Q @PC?RFJCQQLCQQ, BGXXGLCQQ, FC?B?AFCQ, ?LB ?@BMKGL?J N?GL DMP RFMQC ?DDCARCB @W RFGQ BGQC?QC.
!@>LI< 3.27 )K QEFP ?ILLA PJB>O, SFPR>IFWBA >Q 535U J>DKFCF@>QFLK RPFKD ?OFDEQ CFBIA JF@OLP@LMV, PF@HIB @BIIP >OB @OBP@BKQ PE>MBA, TEFIB KLOJ>I @BIIP >OB AFP@-PE>MBA. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V %A 5QEJ>K; P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
!+)43*'7> !97:)9:7+ 5FC JMA?J DMJBGLE MD RFC NMJWNCNRGBC GL QMKC PCEGMLQ EGTCQ PGQC RM RFC secondary structure MD RFC NPMRCGL. 5FC KMQR AMKKML ?PC RFC -helix ?LB -pleated sheet QRPSARSPCQ (Figure 3.28). #MRF QRPSARSPCQ ?PC RFC C-FCJGV QRPSARSPCqRFC FCJGV FCJB GL QF?NC @W FWBPMECL @MLBQ. 5FC FWBPMECL @MLBQ DMPK @CRUCCL RFC MVWECL ?RMK GL RFC A?P@MLWJ EPMSN GL MLC ?KGLM ?AGB ?LB ?LMRFCP ?KGLM ?AGB RF?R GQ DMSP ?KGLM ?AGBQ D?PRFCP ?JMLE RFC AF?GL.
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95
!@>LI< 3.28 4EB C-EBIFU >KA D-MIB>QBA PEBBQ >OB PB@LKA>OV PQOR@QROBP LC MOLQBFKP QE>Q CLOJ ?B@>RPB LC EVAOLDBK ?LKAFKD ?BQTBBK @>O?LKVI >KA >JFKL DOLRMP FK QEB MBMQFAB ?>@H?LKB. #BOQ>FK >JFKL >@FAP E>SB > MOLMBKPFQV QL CLOJ >K C-EBIFU, TEFIB LQEBOP E>SB > MOLMBKPFQV QL CLOJ > D-MIB>QBA PEBBQ.
&TCPW FCJGA?J RSPL GL ?L ?JNF? FCJGV F?Q 3.6 ?KGLM ?AGB PCQGBSCQ. 5FC 3 EPMSNQ (RFC T?PG?LR EPMSNQ) MD RFC NMJWNCNRGBC NPMRPSBC MSR DPMK RFC C-FCJGV AF?GL. *L RFC D-NJC?RCB QFCCR, RFC tNJC?RQu ?PC DMPKCB @W FWBPMECL @MLBGLE @CRUCCL ?RMKQ ML RFC @?AI@MLC MD RFC NMJWNCNRGBC AF?GL. 5FC 3 EPMSNQ ?PC ?RR?AFCB RM RFC A?P@MLQ ?LB CVRCLB ?@MTC ?LB @CJMU RFC DMJBQ MD RFC NJC?R. 5FC NJC?RCB QCEKCLRQ ?JGEL N?P?JJCJ MP ?LRGN?P?JJCJ RM C?AF MRFCP, ?LB FWBPMECL @MLBQ DMPK @CRUCCL RFC N?PRG?JJW NMQGRGTC LGRPMECL ?RMK GL RFC ?KGLM EPMSN ?LB RFC N?PRG?JJW LCE?RGTC MVWECL ?RMK GL RFC A?P@MLWJ EPMSN MD RFC NCNRGBC @?AI@MLC. 5FC C-FCJGV ?LB D-NJC?RCB QFCCR QRPSARSPCQ ?PC DMSLB GL KMQR EJM@SJ?P ?LB DG@PMSQ NPMRCGLQ ?LB RFCW NJ?W ?L GKNMPR?LR QRPSARSP?J PMJC. "+79/'7> !97:)9:7+ 5FC SLGOSC RFPCC-BGKCLQGML?J QRPSARSPC MD ? NMJWNCNRGBC GQ GRQ tertiary structure (Figure 3.29). 5FGQ QRPSARSPC GQ GL N?PR BSC RM AFCKGA?J GLRCP?ARGMLQ ?R UMPI ML RFC NMJWNCNRGBC AF?GL. 1PGK?PGJW, RFC GLRCP?ARGMLQ ?KMLE 3 EPMSNQ APC?RCQ RFC AMKNJCV RFPCC-BGKCLQGML?J RCPRG?PW QRPSARSPC MD ? NPMRCGL. 5FC L?RSPC MD RFC 3 EPMSNQ DMSLB GL RFC ?KGLM ?AGBQ GLTMJTCB A?L AMSLRCP?AR RFC DMPK?RGML MD RFC FWBPMECL @MLBQ BCQAPG@CB DMP QR?LB?PB QCAMLB?PW QRPSARSPCQ. 'MP CV?KNJC, 3 EPMSNQ UGRF JGIC AF?PECQ ?PC PCNCJJCB @W C?AF MRFCP ?LB RFMQC UGRF SLJGIC AF?PECQ ?PC ?RRP?ARCB RM C?AF MRFCP (GMLGA @MLBQ). 8FCL NPMRCGL DMJBGLE R?ICQ NJ?AC, RFC FWBPMNFM@GA 3 EPMSNQ MD LMLNMJ?P ?KGLM ?AGBQ J?W GL RFC GLRCPGMP MD RFC NPMRCGL, UFCPC?Q RFC FWBPMNFGJGA 3 EPMSNQ J?W ML RFC MSRQGBC. 5FC DMPKCP RWNCQ MD GLRCP?ARGMLQ ?PC ?JQM ILMUL ?Q FWBPMNFM@GA GLRCP?ARGMLQ. *LRCP?ARGML @CRUCCL AWQRCGLC QGBC AF?GLQ DMPKQ BGQSJDGBC JGLI?ECQ GL RFC NPCQCLAC MD MVWECL, RFC MLJW AMT?JCLR @MLB DMPKGLE BSPGLE NPMRCGL DMJBGLE.
96
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!@>LI< 3.29 4EB QBOQF>OV PQOR@QROB LC MOLQBFKP FP ABQBOJFKBA ?V > S>OFBQV LC @EBJF@>I FKQBO>@QFLKP. 4EBPB FK@IRAB EVAOLMEL?F@ FKQBO>@QFLKP, FLKF@ ?LKAFKD, EVAOLDBK ?LKAFKD >KA AFPRICFAB IFKH>DBP.
"JJ MD RFCQC GLRCP?ARGMLQ, UC?I ?LB QRPMLE, BCRCPKGLC RFC DGL?J RFPCC-BGKCLQGML?J QF?NC MD RFC NPMRCGL. 8FCL ? NPMRCGL JMQCQ GRQ RFPCC-BGKCLQGML?J QF?NC, GR K?W LM JMLECP @C DSLARGML?J. Q:'9+73'7> !97:)9:7+ *L L?RSPC, QMKC NPMRCGLQ ?PC DMPKCB DPMK QCTCP?J NMJWNCNRGBCQ, ?JQM ILMUL ?Q QS@SLGRQ, ?LB RFC GLRCP?ARGML MD RFCQC QS@SLGRQ DMPKQ RFC quaternary structure. 8C?I GLRCP?ARGMLQ @CRUCCL RFC QS@SLGRQ FCJN RM QR?@GJGXC RFC MTCP?JJ QRPSARSPC. 'MP CV?KNJC, GLQSJGL (? EJM@SJ?P NPMRCGL) F?Q ? AMK@GL?RGML MD FWBPMECL @MLBQ ?LB BGQSJDGBC @MLBQ RF?R A?SQC GR RM @C KMQRJW AJSKNCB GLRM ? @?JJ QF?NC. *LQSJGL QR?PRQ MSR ?Q ? QGLEJC NMJWNCNRGBC ?LB JMQCQ QMKC GLRCPL?J QCOSCLACQ GL RFC NPCQCLAC MD NMQR-RP?LQJ?RGML?J KMBGDGA?RGML ?DRCP RFC DMPK?RGML MD RFC BGQSJDGBC JGLI?ECQ RF?R FMJB RFC PCK?GLGLE AF?GLQ RMECRFCP. 4GJI (? DG@PMSQ NPMRCGL), FMUCTCP, F?Q ? D-NJC?RCB QFCCR QRPSARSPC RF?R GQ RFC PCQSJR MD FWBPMECL @MLBGLE @CRUCCL BGDDCPCLR AF?GLQ. 5FC DMSP JCTCJQ MD NPMRCGL QRPSARSPC (NPGK?PW, QCAMLB?PW, RCPRG?PW, ?LB OS?RCPL?PW) ?PC GJJSQRP?RCB GL Figure 3.30.
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97
!@>LI< 3.30 4EB CLRO IBSBIP LC MOLQBFK PQOR@QROB @>K ?B L?PBOSBA FK QEBPB FIIRPQO>QFLKP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .>QFLK>I (RJ>K 'BKLJB 2BPB>O@E )KPQFQRQB)
D<E8KLI8K@FE 8E; +IFK<@E !FC;@E>

&?AF NPMRCGL F?Q GRQ MUL SLGOSC QCOSCLAC ?LB QF?NC RF?R ?PC FCJB RMECRFCP @W AFCKGA?J GLRCP?ARGMLQ. *D RFC NPMRCGL GQ QS@HCAR RM AF?LECQ GL RCKNCP?RSPC, N), MP CVNMQSPC RM AFCKGA?JQ, RFC NPMRCGL QRPSARSPC K?W AF?LEC, JMQGLE GRQ QF?NC UGRFMSR JMQGLE GRQ NPGK?PW QCOSCLAC GL UF?R GQ ILMUL ?Q BCL?RSP?RGML. %CL?RSP?RGML GQ MDRCL PCTCPQG@JC @CA?SQC RFC NPGK?PW QRPSARSPC MD RFC NMJWNCNRGBC GQ AMLQCPTCB GL RFC NPMACQQ GD RFC BCL?RSPGLE ?ECLR GQ PCKMTCB, ?JJMUGLE RFC NPMRCGL RM PCQSKC GRQ DSLARGML. 4MKCRGKCQ BCL?RSP?RGML GQ GPPCTCPQG@JC, JC?BGLE RM JMQQ MD DSLARGML. 0LC CV?KNJC MD GPPCTCPQG@JC NPMRCGL BCL?RSP?RGML GQ UFCL ?L CEE GQ DPGCB. 5FC ?J@SKGL NPMRCGL GL RFC JGOSGB CEE UFGRC GQ BCL?RSPCB UFCL NJ?ACB GL ? FMR N?L. /MR ?JJ NPMRCGLQ ?PC BCL?RSPCB ?R FGEF RCKNCP?RSPCQ; DMP GLQR?LAC, @?ARCPG? RF?R QSPTGTC GL FMR QNPGLEQ F?TC NPMRCGLQ RF?R DSLARGML ?R RCKNCP?RSPCQ AJMQC RM @MGJGLE. 5FC QRMK?AF GQ ?JQM TCPW ?AGBGA, F?Q ? JMU N), ?LB BCL?RSPCQ NPMRCGLQ ?Q N?PR MD RFC BGECQRGML NPMACQQ; FMUCTCP, RFC BGECQRGTC CLXWKCQ MD RFC QRMK?AF PCR?GL RFCGP ?ARGTGRW SLBCP RFCQC AMLBGRGMLQ. 1PMRCGL DMJBGLE GQ APGRGA?J RM GRQ DSLARGML. *R U?Q MPGEGL?JJW RFMSEFR RF?R RFC NPMRCGLQ RFCKQCJTCQ UCPC PCQNMLQG@JC DMP RFC DMJBGLE NPMACQQ. 0LJW PCACLRJW U?Q GR DMSLB RF?R MDRCL RFCW PCACGTC ?QQGQR?LAC GL RFC DMJBGLE NPMACQQ DPMK NPMRCGL FCJNCPQ ILMUL ?Q chaperones (MP AF?NCPMLGLQ) RF?R ?QQMAG?RC UGRF RFC R?PECR NPMRCGL BSPGLE RFC DMJBGLE NPMACQQ. 5FCW ?AR @W NPCTCLRGLE ?EEPCE?RGML MD NMJWNCNRGBCQ RF?R K?IC SN RFC AMKNJCRC NPMRCGL QRPSARSPC, ?LB RFCW BGQ?QQMAG?RC DPMK RFC NPMRCGL MLAC RFC R?PECR NPMRCGL GQ DMJBCB.
98
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'MP ?L ?BBGRGML?J NCPQNCARGTC ML NPMRCGLQ, TGCU this animation (http://openstaxcollege.org/l/ proteins) A?JJCB t#GMKMJCASJCQ: 5FC 1PMRCGLQ.u
3.5 R )L:C<@: A:@;J

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC QRPSARSPC MD LSAJCGA ?AGBQ ?LB BCDGLC RFC RUM RWNCQ MD LSAJCGA ?AGBQ z &VNJ?GL RFC QRPSARSPC ?LB PMJC MD %/" z &VNJ?GL RFC QRPSARSPC ?LB PMJCQ MD 3/" Nucleic acids ?PC RFC KMQR GKNMPR?LR K?APMKMJCASJCQ DMP RFC AMLRGLSGRW MD JGDC. 5FCW A?PPW RFC ECLCRGA @JSCNPGLR MD ? ACJJ ?LB A?PPW GLQRPSARGMLQ DMP RFC DSLARGMLGLE MD RFC ACJJ.
D)A 8E; -)A

5FC RUM K?GL RWNCQ MD LSAJCGA ?AGBQ ?PC deoxyribonucleic acid (DNA) ?LB ribonucleic acid (RNA). %/" GQ RFC ECLCRGA K?RCPG?J DMSLB GL ?JJ JGTGLE MPE?LGQKQ, P?LEGLE DPMK QGLEJC-ACJJCB @?ARCPG? RM KSJRGACJJSJ?P K?KK?JQ. *R GQ DMSLB GL RFC LSAJCSQ MD CSI?PWMRCQ ?LB GL RFC MPE?LCJJCQ, AFJMPMNJ?QRQ, ?LB KGRMAFMLBPG?. *L NPMI?PWMRCQ, RFC %/" GQ LMR CLAJMQCB GL ? KCK@P?LMSQ CLTCJMNC. 5FC CLRGPC ECLCRGA AMLRCLR MD ? ACJJ GQ ILMUL ?Q GRQ ECLMKC, ?LB RFC QRSBW MD ECLMKCQ GQ ECLMKGAQ. *L CSI?PWMRGA ACJJQ @SR LMR GL NPMI?PWMRCQ, %/" DMPKQ ? AMKNJCV UGRF FGQRMLC NPMRCGLQ RM DMPK AFPMK?RGL, RFC QS@QR?LAC MD CSI?PWMRGA AFPMKMQMKCQ. " AFPMKMQMKC K?W AMLR?GL RCLQ MD RFMSQ?LBQ MD ECLCQ. .?LW ECLCQ AMLR?GL RFC GLDMPK?RGML RM K?IC NPMRCGL NPMBSARQ; MRFCP ECLCQ AMBC DMP 3/" NPMBSARQ. %/" AMLRPMJQ ?JJ MD RFC ACJJSJ?P ?ARGTGRGCQ @W RSPLGLE RFC ECLCQ tMLu MP tMDD.u 5FC MRFCP RWNC MD LSAJCGA ?AGB, 3/", GQ KMQRJW GLTMJTCB GL NPMRCGL QWLRFCQGQ. 5FC %/" KMJCASJCQ LCTCP JC?TC RFC LSAJCSQ @SR GLQRC?B SQC ?L GLRCPKCBG?PW RM AMKKSLGA?RC UGRF RFC PCQR MD RFC ACJJ. 5FGQ GLRCPKCBG?PW GQ RFC messenger RNA (mRNA). 0RFCP RWNCQ MD 3/"qJGIC P3/", R3/", ?LB KGAPM3/"q?PC GLTMJTCB GL NPMRCGL QWLRFCQGQ ?LB GRQ PCESJ?RGML. %/" ?LB 3/" ?PC K?BC SN MD KMLMKCPQ ILMUL ?Q nucleotides. 5FC LSAJCMRGBCQ AMK@GLC UGRF C?AF MRFCP RM DMPK ? polynucleotide, %/" MP 3/". &?AF LSAJCMRGBC GQ K?BC SN MD RFPCC AMKNMLCLRQ: ? LGRPMECLMSQ @?QC, ? NCLRMQC (DGTC-A?P@ML) QSE?P, ?LB ? NFMQNF?RC EPMSN (Figure 3.31). &?AF LGRPMECLMSQ @?QC GL ? LSAJCMRGBC GQ ?RR?AFCB RM ? QSE?P KMJCASJC, UFGAF GQ ?RR?AFCB RM MLC MP KMPC NFMQNF?RC EPMSNQ.
#(!04%2 3 X ")/,/')#!, -!#2/-/,%#5,%3
99
!@>LI< 3.31 ! KR@IBLQFAB FP J>AB RM LC QEOBB @LJMLKBKQP: > KFQOLDBKLRP ?>PB, > MBKQLPB PRD>O, >KA LKB LO JLOB MELPME>QB DOLRMP. #>O?LK OBPFARBP FK QEB MBKQLPB >OB KRJ?BOBA 1{ QEOLRDE 5{ (QEB MOFJB AFPQFKDRFPEBP QEBPB OBPFARBP COLJ QELPB FK QEB ?>PB, TEF@E >OB KRJ?BOBA TFQELRQ RPFKD > MOFJB KLQ>QFLK). 4EB ?>PB FP >QQ>@EBA QL QEB 1{ MLPFQFLK LC QEB OF?LPB, >KA QEB MELPME>QB FP >QQ>@EBA QL QEB 5{ MLPFQFLK. 7EBK > MLIVKR@IBLQFAB FP CLOJBA, QEB 5{ MELPME>QB LC QEB FK@LJFKD KR@IBLQFAB >QQ>@EBP QL QEB 3{ EVAOLUVI DOLRM >Q QEB BKA LC QEB DOLTFKD @E>FK. 4TL QVMBP LC MBKQLPB >OB CLRKA FK KR@IBLQFABP, ABLUVOF?LPB (CLRKA FK $.!) >KA OF?LPB (CLRKA FK 2.!). $BLUVOF?LPB FP PFJFI>O FK PQOR@QROB QL OF?LPB, ?RQ FQ E>P >K ( FKPQB>A LC >K /( >Q QEB 2{ MLPFQFLK. ">PBP @>K ?B AFSFABA FKQL QTL @>QBDLOFBP: MROFKBP >KA MVOFJFAFKBP. 0ROFKBP E>SB > ALR?IB OFKD PQOR@QROB, >KA MVOFJFAFKBP E>SB > PFKDIB OFKD.
5FC LGRPMECLMSQ @?QCQ, GKNMPR?LR AMKNMLCLRQ MD LSAJCMRGBCQ, ?PC MPE?LGA KMJCASJCQ ?LB ?PC QM L?KCB @CA?SQC RFCW AMLR?GL A?P@ML ?LB LGRPMECL. 5FCW ?PC @?QCQ @CA?SQC RFCW AMLR?GL ?L ?KGLM EPMSN RF?R F?Q RFC NMRCLRG?J MD @GLBGLE ?L CVRP? FWBPMECL, ?LB RFSQ, BCAPC?QCQ RFC FWBPMECL GML AMLACLRP?RGML GL GRQ CLTGPMLKCLR, K?IGLE GR KMPC @?QGA. &?AF LSAJCMRGBC GL %/" AMLR?GLQ MLC MD DMSP NMQQG@JC LGRPMECLMSQ @?QCQ: ?BCLGLC ("), ES?LGLC (() AWRMQGLC ($), ?LB RFWKGLC (5). "BCLGLC ?LB ES?LGLC ?PC AJ?QQGDGCB ?Q purines. 5FC NPGK?PW QRPSARSPC MD ? NSPGLC GQ RUM A?P@MLLGRPMECL PGLEQ. $WRMQGLC, RFWKGLC, ?LB SP?AGJ ?PC AJ?QQGDGCB ?Q pyrimidines UFGAF F?TC ? QGLEJC A?P@MLLGRPMECL PGLE ?Q RFCGP NPGK?PW QRPSARSPC (Figure 3.31). &?AF MD RFCQC @?QGA A?P@ML-LGRPMECL PGLEQ F?Q BGDDCPCLR DSLARGML?J EPMSNQ ?RR?AFCB RM GR. *L KMJCASJ?P @GMJMEW QFMPRF?LB, RFC LGRPMECLMSQ @?QCQ ?PC QGKNJW ILMUL @W RFCGP QWK@MJQ ", 5, (, $, ?LB 6. %/" AMLR?GLQ ", 5, (, ?LB $ UFCPC?Q 3/" AMLR?GLQ ", 6, (, ?LB $. 5FC NCLRMQC QSE?P GL %/" GQ BCMVWPG@MQC, ?LB GL 3/", RFC QSE?P GQ PG@MQC (Figure 3.31). 5FC BGDDCPCLAC @CRUCCL RFC QSE?PQ GQ RFC NPCQCLAC MD RFC FWBPMVWJ EPMSN ML RFC QCAMLB A?P@ML MD RFC PG@MQC ?LB FWBPMECL ML RFC QCAMLB A?P@ML MD RFC BCMVWPG@MQC. 5FC A?P@ML ?RMKQ MD RFC QSE?P KMJCASJC ?PC LSK@CPCB ?Q 1v, 2v, 3v, 4v, ?LB 5v (1v GQ PC?B ?Q tMLC NPGKCu). 5FC NFMQNF?RC PCQGBSC GQ ?RR?AFCB RM RFC FWBPMVWJ EPMSN MD RFC 5v A?P@ML MD MLC QSE?P ?LB RFC FWBPMVWJ EPMSN MD RFC 3v A?P@ML MD RFC QSE?P MD RFC LCVR LSAJCMRGBC, UFGAF DMPKQ ? 5vp3v phosphodiester JGLI?EC. 5FC NFMQNFMBGCQRCP JGLI?EC GQ LMR DMPKCB @W QGKNJC BCFWBP?RGML PC?ARGML JGIC RFC MRFCP JGLI?ECQ AMLLCARGLE KMLMKCPQ GL K?APMKMJCASJCQ:
100
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GRQ DMPK?RGML GLTMJTCQ RFC PCKMT?J MD RUM NFMQNF?RC EPMSNQ. " NMJWLSAJCMRGBC K?W F?TC RFMSQ?LBQ MD QSAF NFMQNFMBGCQRCP JGLI?ECQ.
D)A DFL9C<-#<C@O .KIL:KLI<

%/" F?Q ? BMS@JC-FCJGV QRPSARSPC (Figure 3.32). 5FC QSE?P ?LB NFMQNF?RC JGC ML RFC MSRQGBC MD RFC FCJGV, DMPKGLE RFC @?AI@MLC MD RFC %/". 5FC LGRPMECLMSQ @?QCQ ?PC QR?AICB GL RFC GLRCPGMP, JGIC RFC QRCNQ MD ? QR?GPA?QC, GL N?GPQ; RFC N?GPQ ?PC @MSLB RM C?AF MRFCP @W FWBPMECL @MLBQ. &TCPW @?QC N?GP GL RFC BMS@JC FCJGTV GQ QCN?P?RCB DPMK RFC LCVR @?QC N?GP @W 0.34 LK. 5FC RUM QRP?LBQ MD RFC FCJGV PSL GL MNNMQGRC BGPCARGMLQ, KC?LGLE RF?R RFC 5v A?P@ML CLB MD MLC QRP?LB UGJJ D?AC RFC 3v A?P@ML CLB MD GRQ K?RAFGLE QRP?LB. (5FGQ GQ PCDCPPCB RM ?Q ?LRGN?P?JJCJ MPGCLR?RGML ?LB GQ GKNMPR?LR RM %/" PCNJGA?RGML ?LB GL K?LW LSAJCGA ?AGB GLRCP?ARGMLQ.)
!@>LI< 3.32 .>QFSB $.! FP >K >KQFM>O>IIBI ALR?IB EBIFU. 4EB MELPME>QB ?>@H?LKB (FKAF@>QBA ?V QEB @ROSV IFKBP) FP LK QEB LRQPFAB, >KA QEB ?>PBP >OB LK QEB FKPFAB. %>@E ?>PB COLJ LKB PQO>KA FKQBO>@QP SF> EVAOLDBK ?LKAFKD TFQE > ?>PB COLJ QEB LMMLPFKD PQO>KA. (@OBAFQ: *BOLJB 7>IHBO/$BKKFP -VQP)
0LJW ACPR?GL RWNCQ MD @?QC N?GPGLE ?PC ?JJMUCB. 'MP CV?KNJC, ? ACPR?GL NSPGLC A?L MLJW N?GP UGRF ? ACPR?GL NWPGKGBGLC. 5FGQ KC?LQ " A?L N?GP UGRF 5, ?LB ( A?L N?GP UGRF $, ?Q QFMUL GL Figure 3.33. 5FGQ GQ ILMUL ?Q RFC @?QC AMKNJCKCLR?PW PSJC. *L MRFCP UMPBQ, RFC %/" QRP?LBQ ?PC AMKNJCKCLR?PW RM C?AF MRFCP. *D RFC QCOSCLAC MD MLC QRP?LB GQ ""55(($$, RFC AMKNJCKCLR?PW QRP?LB UMSJB F?TC RFC QCOSCLAC 55""$$((. %SPGLE %/" PCNJGA?RGML, C?AF QRP?LB GQ AMNGCB, PCQSJRGLE GL ? B?SEFRCP %/" BMS@JC FCJGV AMLR?GLGLE MLC N?PCLR?J %/" QRP?LB ?LB ? LCUJW QWLRFCQGXCB QRP?LB.
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101
!@>LI< 3.33 )K > ALR?IB PQO>KABA $.! JLIB@RIB, QEB QTL PQO>KAP ORK >KQFM>O>IIBI QL LKB >KLQEBO PL QE>Q LKB PQO>KA ORKP 5{ QL 3{ >KA QEB LQEBO 3{ QL 5{. 4EB MELPME>QB ?>@H?LKB FP IL@>QBA LK QEB LRQPFAB, >KA QEB ?>PBP >OB FK QEB JFAAIB. !ABKFKB CLOJP EVAOLDBK ?LKAP (LO ?>PB M>FOP) TFQE QEVJFKB, >KA DR>KFKB ?>PB M>FOP TFQE @VQLPFKB.
! JRQ>QFLK L@@ROP, >KA @VQLPFKB FP OBMI>@BA TFQE >ABKFKB. 7E>Q FJM>@Q AL VLR QEFKH QEFP TFII E>SB LK QEB $.! PQOR@QROB
-)A
3G@MLSAJCGA ?AGB, MP 3/", GQ K?GLJW GLTMJTCB GL RFC NPMACQQ MD NPMRCGL QWLRFCQGQ SLBCP RFC BGPCARGML MD %/". 3/" GQ SQS?JJW QGLEJC-QRP?LBCB ?LB GQ K?BC MD PG@MLSAJCMRGBCQ RF?R ?PC JGLICB @W NFMQNFMBGCQRCP @MLBQ. " PG@MLSAJCMRGBC GL RFC 3/" AF?GL AMLR?GLQ PG@MQC (RFC NCLRMQC QSE?P), MLC MD RFC DMSP LGRPMECLMSQ @?QCQ (", 6, (, ?LB $), ?LB RFC NFMQNF?RC EPMSN. 5FCPC ?PC DMSP K?HMP RWNCQ MD 3/": KCQQCLECP 3/" (K3/"), PG@MQMK?J 3/" (P3/"), RP?LQDCP 3/" (R3/"), ?LB KGAPM3/" (KG3/"). 5FC DGPQR, K3/", A?PPGCQ RFC KCQQ?EC DPMK %/", UFGAF AMLRPMJQ ?JJ MD RFC ACJJSJ?P ?ARGTGRGCQ GL ? ACJJ. *D ? ACJJ PCOSGPCQ ? ACPR?GL NPMRCGL RM @C QWLRFCQGXCB, RFC ECLC DMP RFGQ NPMBSAR GQ RSPLCB tMLu ?LB RFC KCQQCLECP 3/" GQ QWLRFCQGXCB GL RFC LSAJCSQ. 5FC 3/" @?QC QCOSCLAC GQ AMKNJCKCLR?PW RM RFC AMBGLE QCOSCLAC MD RFC %/" DPMK UFGAF GR F?Q @CCL AMNGCB. )MUCTCP, GL 3/", RFC @?QC 5 GQ ?@QCLR ?LB 6 GQ NPCQCLR GLQRC?B. *D RFC %/" QRP?LB F?Q ? QCOSCLAC ""55($($, RFC QCOSCLAC MD RFC AMKNJCKCLR?PW 3/" GQ 66""$($(. *L RFC AWRMNJ?QK, RFC K3/" GLRCP?ARQ UGRF PG@MQMKCQ ?LB MRFCP ACJJSJ?P K?AFGLCPW (Figure 3.34).
!@>LI< 3.34 ! OF?LPLJB E>P QTL M>OQP: > I>ODB PR?RKFQ >KA > PJ>II PR?RKFQ. 4EB J2.! PFQP FK ?BQTBBK QEB QTL PR?RKFQP. ! Q2.! JLIB@RIB OB@LDKFWBP > @LALK LK QEB J2.!, ?FKAP QL FQ ?V @LJMIBJBKQ>OV ?>PB M>FOFKD, >KA >AAP QEB @LOOB@Q >JFKL >@FA QL QEB DOLTFKD MBMQFAB @E>FK.
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104
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106
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5FC KGAPMQAMNCQ UC SQC RMB?W ?PC D?P KMPC AMKNJCV RF?L RFMQC SQCB GL RFC 1600Q @W "LRMLW T?L -CCSUCLFMCI, ? %SRAF QFMNICCNCP UFM F?B EPC?R QIGJJ GL AP?DRGLE JCLQCQ. %CQNGRC RFC JGKGR?RGMLQ MD FGQ LMU-?LAGCLR JCLQCQ, T?L -CCSUCLFMCI M@QCPTCB RFC KMTCKCLRQ MD NPMRGQR? (? RWNC MD QGLEJC-ACJJCB MPE?LGQK) ?LB QNCPK, UFGAF FC AMJJCARGTCJW RCPKCB t?LGK?JASJCQ.u *L ? 1665 NS@JGA?RGML A?JJCB M0*96.9(7/0(, CVNCPGKCLR?J QAGCLRGQR 3M@CPR )MMIC AMGLCB RFC RCPK tACJJu DMP RFC @MV-JGIC QRPSARSPCQ FC M@QCPTCB UFCL TGCUGLE AMPI RGQQSC RFPMSEF ? JCLQ. *L RFC 1670Q, T?L -CCSUCLFMCI BGQAMTCPCB @?ARCPG? ?LB NPMRMXM?. -?RCP ?BT?LACQ GL JCLQCQ, KGAPMQAMNC AMLQRPSARGML, ?LB QR?GLGLE RCAFLGOSCQ CL?@JCB MRFCP QAGCLRGQRQ RM QCC QMKC AMKNMLCLRQ GLQGBC ACJJQ. #W RFC J?RC 1830Q, @MR?LGQR .?RRFG?Q 4AFJCGBCL ?LB XMMJMEGQR 5FCMBMP 4AFU?LL UCPC QRSBWGLE RGQQSCQ ?LB NPMNMQCB RFC unified cell theory, UFGAF QR?RCQ RF?R ?JJ JGTGLE RFGLEQ ?PC AMKNMQCB MD MLC MP KMPC ACJJQ, RFC ACJJ GQ RFC @?QGA SLGR MD JGDC, ?LB LCU ACJJQ ?PGQC DPMK CVGQRGLE ACJJQ. 3SBMJD 7GPAFMU J?RCP K?BC GKNMPR?LR AMLRPG@SRGMLQ RM RFGQ RFCMPW.
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(>SB VLR BSBO EB>OA LC > JBAF@>I QBPQ @>IIBA > 0>M PJB>O (!@>LI< 4.4) )K QEFP QBPQ, > AL@QLO Q>HBP > PJ>II P>JMIB LC @BIIP COLJ QEB RQBOFKB @BOSFU LC > M>QFBKQ >KA PBKAP FQ QL > JBAF@>I I>? TEBOB > @VQLQB@EKLILDFPQ PQ>FKP QEB @BIIP >KA BU>JFKBP QEBJ CLO >KV @E>KDBP QE>Q @LRIA FKAF@>QB @BOSF@>I @>K@BO LO > JF@OL?F>I FKCB@QFLK. #VQLQB@EKLILDFPQP (@VQL- = x@BIIy) >OB MOLCBPPFLK>IP TEL PQRAV @BIIP SF> JF@OLP@LMF@ BU>JFK>QFLKP >KA LQEBO I>?LO>QLOV QBPQP. 4EBV >OB QO>FKBA QL ABQBOJFKB TEF@E @BIIRI>O @E>KDBP >OB TFQEFK KLOJ>I IFJFQP >KA TEF@E >OB >?KLOJ>I. 4EBFO CL@RP FP KLQ IFJFQBA QL @BOSF@>I @BIIP; QEBV PQRAV @BIIRI>O PMB@FJBKP QE>Q @LJB COLJ >II LOD>KP. 7EBK QEBV KLQF@B >?KLOJ>IFQFBP, QEBV @LKPRIQ > M>QELILDFPQ, TEL FP > JBAF@>I AL@QLO TEL @>K J>HB > @IFKF@>I AF>DKLPFP. #VQLQB@EKLILDFPQP MI>V > SFQ>I OLIB FK P>SFKD MBLMIBwP IFSBP. 7EBK >?KLOJ>IFQFBP >OB AFP@LSBOBA B>OIV, > M>QFBKQwP QOB>QJBKQ @>K ?BDFK PLLKBO, TEF@E RPR>IIV FK@OB>PBP QEB @E>K@BP LC > PR@@BPPCRI LRQ@LJB.
!@>LI< 4.4 4EBPB RQBOFKB @BOSFU @BIIP, SFBTBA QEOLRDE > IFDEQ JF@OLP@LMB, TBOB L?Q>FKBA COLJ > 0>M PJB>O. .LOJ>I @BIIP >OB LK QEB IBCQ. 4EB @BIIP LK QEB OFDEQ >OB FKCB@QBA TFQE ERJ>K M>MFIILJ>SFORP ((06). .LQF@B QE>Q QEB FKCB@QBA @BIIP >OB I>ODBO; >IPL, QTL LC QEBPB @BIIP B>@E E>SB QTL KR@IBF FKPQB>A LC LKB, QEB KLOJ>I KRJ?BO. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V %A 5QEJ>K, -$; P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
#(!04%2 4 X #%,, 3425#452%
113
4.2 R +IFB8IPFK@: C<CCJ

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z /?KC CV?KNJCQ MD NPMI?PWMRGA ?LB CSI?PWMRGA MPE?LGQKQ z $MKN?PC ?LB AMLRP?QR NPMI?PWMRGA ACJJQ ?LB CSI?PWMRGA ACJJQ z %CQAPG@C RFC PCJ?RGTC QGXCQ MD BGDDCPCLR IGLBQ MD ACJJQ z &VNJ?GL UFW ACJJQ KSQR @C QK?JJ $CJJQ D?JJ GLRM MLC MD RUM @PM?B A?RCEMPGCQ: NPMI?PWMRGA ?LB CSI?PWMRGA. 0LJW RFC NPCBMKGL?LRJW QGLEJCACJJCB MPE?LGQKQ MD RFC BMK?GLQ #?ARCPG? ?LB "PAF?C? ?PC AJ?QQGDGCB ?Q NPMI?PWMRCQ (NPM- = t@CDMPCu; -I?PW- = tLSAJCSQu). $CJJQ MD ?LGK?JQ, NJ?LRQ, DSLEG, ?LB NPMRGQRQ ?PC ?JJ CSI?PWMRCQ (ACS- = tRPSCu) ?LB ?PC K?BC SN MD CSI?PWMRGA ACJJQ.
CFDGFE<EKJ F= +IFB8IPFK@: C<CCJ

"JJ ACJJQ QF?PC DMSP AMKKML AMKNMLCLRQ: 1) ? NJ?QK? KCK@P?LC, ?L MSRCP AMTCPGLE RF?R QCN?P?RCQ RFC ACJJsQ GLRCPGMP DPMK GRQ QSPPMSLBGLE CLTGPMLKCLR; 2) AWRMNJ?QK, AMLQGQRGLE MD ? HCJJW-JGIC AWRMQMJ UGRFGL RFC ACJJ GL UFGAF MRFCP ACJJSJ?P AMKNMLCLRQ ?PC DMSLB; 3) %/", RFC ECLCRGA K?RCPG?J MD RFC ACJJ; ?LB 4) PG@MQMKCQ, UFGAF QWLRFCQGXC NPMRCGLQ. )MUCTCP, NPMI?PWMRCQ BGDDCP DPMK CSI?PWMRGA ACJJQ GL QCTCP?J U?WQ. " prokaryote GQ ? QGKNJC, KMQRJW QGLEJC-ACJJCB (SLGACJJSJ?P) MPE?LGQK RF?R J?AIQ ? LSAJCSQ, MP ?LW MRFCP KCK@P?LC-@MSLB MPE?LCJJC. 8C UGJJ QFMPRJW AMKC RM QCC RF?R RFGQ GQ QGELGDGA?LRJW BGDDCPCLR GL CSI?PWMRCQ. 1PMI?PWMRGA %/" GQ DMSLB GL ? ACLRP?J N?PR MD RFC ACJJ: RFC nucleoid (Figure 4.5).
!@>LI< 4.5 4EFP CFDROB PELTP QEB DBKBO>IFWBA PQOR@QROB LC > MOLH>OVLQF@ @BII. !II MOLH>OVLQBP E>SB @EOLJLPLJ>I $.! IL@>IFWBA FK > KR@IBLFA, OF?LPLJBP, > @BII JBJ?O>KB, >KA > @BII T>II. 4EB LQEBO PQOR@QROBP PELTK >OB MOBPBKQ FK PLJB, ?RQ KLQ >II, ?>@QBOF>.
.MQR NPMI?PWMRCQ F?TC ? NCNRGBMEJWA?L ACJJ U?JJ ?LB K?LW F?TC ? NMJWQ?AAF?PGBC A?NQSJC (Figure 4.5). 5FC ACJJ U?JJ ?ARQ ?Q ?L CVRP? J?WCP MD NPMRCARGML, FCJNQ RFC ACJJ K?GLR?GL GRQ QF?NC, ?LB NPCTCLRQ BCFWBP?RGML. 5FC A?NQSJC CL?@JCQ RFC ACJJ RM ?RR?AF RM QSPD?ACQ GL GRQ CLTGPMLKCLR. 4MKC NPMI?PWMRCQ F?TC DJ?ECJJ?, NGJG, MP DGK@PG?C. 'J?ECJJ? ?PC SQCB DMP JMAMKMRGML. 1GJG ?PC SQCB RM CVAF?LEC ECLCRGA K?RCPG?J BSPGLE ? RWNC MD PCNPMBSARGML A?JJCB AMLHSE?RGML. 'GK@PG?C ?PC SQCB @W @?ARCPG? RM ?RR?AF RM ? FMQR ACJJ.
114
#(!04%2 4 X #%,, 3425#452%
(@:IF9@FCF>@JK
4EB JLPQ BCCB@QFSB >@QFLK >KVLKB @>K Q>HB QL MOBSBKQ QEB PMOB>A LC @LKQ>DFLRP FIIKBPPBP FP QL T>PE EFP LO EBO E>KAP. 7EV "B@>RPB JF@OL?BP (LOD>KFPJP PL QFKV QE>Q QEBV @>K LKIV ?B PBBK TFQE JF@OLP@LMBP) >OB R?FNRFQLRP. 4EBV IFSB LK ALLOHKL?P, JLKBV, VLRO E>KAP, >KA J>KV LQEBO PROC>@BP. )C PLJBLKB PKBBWBP FKQL EFP E>KA >KA QLR@EBP > ALLOHKL?, >KA >CQBOT>OAP VLR QLR@E QE>Q P>JB ALLOHKL?, QEB JF@OL?BP COLJ QEB PKBBWBOw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mK GL BG?KCRCP, NPMI?PWMRGA ACJJQ ?PC QGELGDGA?LRJW QK?JJCP RF?L CSI?PWMRGA ACJJQ, UFGAF F?TC BG?KCRCPQ P?LEGLE DPMK 10 RM 100 mK (Figure 4.6). 5FC QK?JJ QGXC MD NPMI?PWMRCQ ?JJMUQ GMLQ ?LB MPE?LGA KMJCASJCQ RF?R CLRCP RFCK RM OSGAIJW BGDDSQC RM MRFCP N?PRQ MD RFC ACJJ. 4GKGJ?PJW, ?LW U?QRCQ NPMBSACB UGRFGL ? NPMI?PWMRGA ACJJ A?L OSGAIJW BGDDSQC MSR. 5FGQ GQ LMR RFC A?QC GL CSI?PWMRGA ACJJQ, UFGAF F?TC BCTCJMNCB BGDDCPCLR QRPSARSP?J ?B?NR?RGMLQ RM CLF?LAC GLRP?ACJJSJ?P RP?LQNMPR.
!@>LI< 4.6 4EFP CFDROB PELTP OBI>QFSB PFWBP LC JF@OL?BP LK > ILD>OFQEJF@ P@>IB (OB@>II QE>Q B>@E RKFQ LC FK@OB>PB FK > ILD>OFQEJF@ P@>IB OBMOBPBKQP > 10-CLIA FK@OB>PB FK QEB NR>KQFQV ?BFKD JB>PROBA).
4K?JJ QGXC, GL ECLCP?J, GQ LCACQQ?PW DMP ?JJ ACJJQ, UFCRFCP NPMI?PWMRGA MP CSI?PWMRGA. -CRsQ CV?KGLC UFW RF?R GQ QM. 'GPQR, UCsJJ AMLQGBCP RFC ?PC? ?LB TMJSKC MD ? RWNGA?J ACJJ. /MR ?JJ ACJJQ ?PC QNFCPGA?J GL QF?NC, @SR KMQR RCLB RM ?NNPMVGK?RC ? QNFCPC. :MS K?W PCKCK@CP DPMK WMSP FGEF QAFMMJ ECMKCRPW
#(!04%2 4 X #%,, 3425#452%
115
AMSPQC RF?R RFC DMPKSJ? DMP RFC QSPD?AC ?PC? MD ? QNFCPC GQ 4nP2, UFGJC RFC DMPKSJ? DMP GRQ TMJSKC GQ 4nP3/3. 5FSQ, ?Q RFC P?BGSQ MD ? ACJJ GLAPC?QCQ, GRQ QSPD?AC ?PC? GLAPC?QCQ ?Q RFC QOS?PC MD GRQ P?BGSQ, @SR GRQ TMJSKC GLAPC?QCQ ?Q RFC AS@C MD GRQ P?BGSQ (KSAF KMPC P?NGBJW). 5FCPCDMPC, ?Q ? ACJJ GLAPC?QCQ GL QGXC, GRQ QSPD?AC ?PC?-RM-TMJSKC P?RGM BCAPC?QCQ. 5FGQ Q?KC NPGLAGNJC UMSJB ?NNJW GD RFC ACJJ F?B RFC QF?NC MD ? AS@C (Figure 4.7). *D RFC ACJJ EPMUQ RMM J?PEC, RFC NJ?QK? KCK@P?LC UGJJ LMR F?TC QSDDGAGCLR QSPD?AC ?PC? RM QSNNMPR RFC P?RC MD BGDDSQGML PCOSGPCB DMP RFC GLAPC?QCB TMJSKC. *L MRFCP UMPBQ, ?Q ? ACJJ EPMUQ, GR @CAMKCQ JCQQ CDDGAGCLR. 0LC U?W RM @CAMKC KMPC CDDGAGCLR GQ RM BGTGBC; ?LMRFCP U?W GQ RM BCTCJMN MPE?LCJJCQ RF?R NCPDMPK QNCAGDGA R?QIQ. 5FCQC ?B?NR?RGMLQ JC?B RM RFC BCTCJMNKCLR MD KMPC QMNFGQRGA?RCB ACJJQ A?JJCB CSI?PWMRGA ACJJQ.
!@>LI< 4.7 .LQF@B QE>Q >P > @BII FK@OB>PBP FK PFWB, FQP PROC>@B >OB>-QL-SLIRJB O>QFL AB@OB>PBP. 7EBK QEBOB FP FKPRCCF@FBKQ PROC>@B >OB> QL PRMMLOQ > @BIIwP FK@OB>PFKD SLIRJB, > @BII TFII BFQEBO AFSFAB LO AFB. 4EB @BII LK QEB IBCQ E>P > SLIRJB LC 1 JJ3 >KA > PROC>@B >OB> LC 6 JJ2, TFQE > PROC>@B >OB>-QL-SLIRJB O>QFL LC 6 QL 1, TEBOB>P QEB @BII LK QEB OFDEQ E>P > SLIRJB LC 8 JJ3 >KA > PROC>@B >OB> LC 24 JJ2, TFQE > PROC>@B >OB>-QL-SLIRJB O>QFL LC 3 QL 1.
0OLH>OVLQF@ @BIIP >OB JR@E PJ>IIBO QE>K BRH>OVLQF@ @BIIP. 7E>Q >AS>KQ>DBP JFDEQ PJ>II @BII PFWB @LKCBO LK > @BII 7E>Q >AS>KQ>DBP JFDEQ I>ODB @BII PFWB E>SB
4.3 R LB8IPFK@: C<CCJ

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC QRPSARSPC MD CSI?PWMRGA ACJJQ z $MKN?PC ?LGK?J ACJJQ UGRF NJ?LR ACJJQ z 4R?RC RFC PMJC MD RFC NJ?QK? KCK@P?LC z 4SKK?PGXC RFC DSLARGMLQ MD RFC K?HMP ACJJ MPE?LCJJCQ )?TC WMS CTCP FC?PB RFC NFP?QC tDMPK DMJJMUQ DSLARGML u *RsQ ? NFGJMQMNFW NP?ARGACB GL K?LW GLBSQRPGCQ. *L ?PAFGRCARSPC, RFGQ KC?LQ RF?R @SGJBGLEQ QFMSJB @C AMLQRPSARCB RM QSNNMPR RFC ?ARGTGRGCQ RF?R UGJJ @C A?PPGCB MSR GLQGBC RFCK. 'MP CV?KNJC, ? QIWQAP?NCP QFMSJB @C @SGJR UGRF QCTCP?J CJCT?RMP @?LIQ; ? FMQNGR?J QFMSJB @C @SGJR QM RF?R GRQ CKCPECLAW PMMK GQ C?QGJW ?AACQQG@JC. 0SP L?RSP?J UMPJB ?JQM SRGJGXCQ RFC NPGLAGNJC MD DMPK DMJJMUGLE DSLARGML, CQNCAG?JJW GL ACJJ @GMJMEW, ?LB RFGQ UGJJ @CAMKC AJC?P ?Q UC CVNJMPC CSI?PWMRGA ACJJQ ( Figure 4.8). 6LJGIC NPMI?PWMRGA ACJJQ, eukaryotic cells F?TC: 1) ? KCK@P?LC-@MSLB LSAJCSQ; 2) LSKCPMSQ KCK@P?LC-@MSLB organelles QSAF ?Q RFC CLBMNJ?QKGA PCRGASJSK, (MJEG ?NN?P?RSQ, AFJMPMNJ?QRQ, KGRMAFMLBPG?, ?LB MRFCPQ; ?LB 3) QCTCP?J, PMB-QF?NCB AFPMKMQMKCQ. #CA?SQC ? CSI?PWMRGA ACJJsQ LSAJCSQ GQ QSPPMSLBCB @W ? KCK@P?LC, GR GQ MDRCL Q?GB RM F?TC ? tRPSC LSAJCSQ.u 5FC UMPB tMPE?LCJJCu KC?LQ tJGRRJC MPE?L,u ?LB, ?Q ?JPC?BW KCLRGMLCB, MPE?LCJJCQ F?TC QNCAG?JGXCB ACJJSJ?P DSLARGMLQ, HSQR ?Q RFC MPE?LQ MD WMSP @MBW F?TC QNCAG?JGXCB DSLARGMLQ. "R RFGQ NMGLR, GR QFMSJB @C AJC?P RM WMS RF?R CSI?PWMRGA ACJJQ F?TC ? KMPC AMKNJCV QRPSARSPC RF?L NPMI?PWMRGA ACJJQ. 0PE?LCJJCQ ?JJMU BGDDCPCLR DSLARGMLQ RM @C AMKN?PRKCLR?JGXCB GL BGDDCPCLR ?PC?Q MD RFC ACJJ. #CDMPC RSPLGLE RM MPE?LCJJCQ, JCRsQ DGPQR CV?KGLC RUM GKNMPR?LR AMKNMLCLRQ MD RFC ACJJ: RFC NJ?QK? KCK@P?LC ?LB RFC AWRMNJ?QK.
116
#(!04%2 4 X #%,, 3425#452%
(8)
(9) !@>LI< 4.8 4EBPB CFDROBP PELT QEB J>GLO LOD>KBIIBP >KA LQEBO @BII @LJMLKBKQP LC (>) > QVMF@>I >KFJ>I @BII >KA (?) > QVMF@>I BRH>OVLQF@ MI>KQ @BII. 4EB MI>KQ @BII E>P > @BII T>II, @EILOLMI>PQP, MI>PQFAP, >KA > @BKQO>I S>@RLIBuPQOR@QROBP KLQ CLRKA FK >KFJ>I @BIIP. 0I>KQ @BIIP AL KLQ E>SB IVPLPLJBP LO @BKQOLPLJBP.
)C QEB KR@IBLIRP TBOB KLQ >?IB QL @>OOV LRQ FQP CRK@QFLK, TE>Q LQEBO @BIIRI>O LOD>KBIIBP TLRIA ?B >CCB@QBA
/?< +C8JD8 (<D9I8E<

-GIC NPMI?PWMRCQ, CSI?PWMRGA ACJJQ F?TC ? plasma membrane ( Figure 4.9), ? NFMQNFMJGNGB @GJ?WCP UGRF CK@CBBCB NPMRCGLQ RF?R QCN?P?RCQ RFC GLRCPL?J AMLRCLRQ MD RFC ACJJ DPMK GRQ QSPPMSLBGLE CLTGPMLKCLR. " NFMQNFMJGNGB GQ ? JGNGB KMJCASJC UGRF RUM D?RRW ?AGB AF?GLQ ?LB ? NFMQNF?RC-AMLR?GLGLE EPMSN. 5FC NJ?QK? KCK@P?LC AMLRPMJQ RFC N?QQ?EC MD MPE?LGA KMJCASJCQ, GMLQ, U?RCP, ?LB MVWECL GLRM ?LB MSR MD RFC
#(!04%2 4 X #%,, 3425#452%
117
ACJJ. 8?QRCQ (QSAF ?Q A?P@ML BGMVGBC ?LB ?KKMLG?) ?JQM JC?TC RFC ACJJ @W N?QQGLE RFPMSEF RFC NJ?QK? KCK@P?LC.
!@>LI< 4.9 4EB BRH>OVLQF@ MI>PJ> JBJ?O>KB FP > MELPMELIFMFA ?FI>VBO TFQE MOLQBFKP >KA @ELIBPQBOLI BJ?BAABA FK FQ.
5FC NJ?QK? KCK@P?LCQ MD ACJJQ RF?R QNCAG?JGXC GL ?@QMPNRGML ?PC DMJBCB GLRM DGLECPJGIC NPMHCARGMLQ A?JJCB KGAPMTGJJG (QGLESJ?P = KGAPMTGJJSQ); ( Figure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
!@>LI< 4.10 -F@OLSFIIF, PELTK EBOB >P QEBV >MMB>O LK @BIIP IFKFKD QEB PJ>II FKQBPQFKB, FK@OB>PB QEB PROC>@B >OB> >S>FI>?IB CLO >?PLOMQFLK. 4EBPB JF@OLSFIIF >OB LKIV CLRKA LK QEB >OB> LC QEB MI>PJ> JBJ?O>KB QE>Q C>@BP QEB @>SFQV COLJ TEF@E PR?PQ>K@BP TFII ?B >?PLO?BA. (@OBAFQ "JF@OLDO>ME": JLAFCF@>QFLK LC TLOH ?V ,LRFP> (LT>OA)
/?< CPKFGC8JD
5FC cytoplasm GQ RFC CLRGPC PCEGML MD ? ACJJ @CRUCCL RFC NJ?QK? KCK@P?LC ?LB RFC LSAJC?P CLTCJMNC (? QRPSARSPC RM @C BGQASQQCB QFMPRJW). *R GQ K?BC SN MD MPE?LCJJCQ QSQNCLBCB GL RFC ECJ-JGIC cytosol, RFC AWRMQICJCRML, ?LB T?PGMSQ AFCKGA?JQ ( Figure 4.8). &TCL RFMSEF RFC AWRMNJ?QK AMLQGQRQ MD 70 RM 80 NCPACLR U?RCP, GR F?Q ? QCKG-QMJGB AMLQGQRCLAW, UFGAF AMKCQ DPMK RFC NPMRCGLQ UGRFGL GR. )MUCTCP, NPMRCGLQ ?PC LMR RFC MLJW MPE?LGA KMJCASJCQ DMSLB GL RFC AWRMNJ?QK. (JSAMQC ?LB MRFCP QGKNJC QSE?PQ, NMJWQ?AAF?PGBCQ, ?KGLM ?AGBQ, LSAJCGA ?AGBQ, D?RRW ?AGBQ, ?LB BCPGT?RGTCQ MD EJWACPMJ ?PC DMSLB RFCPC, RMM. *MLQ MD QMBGSK, NMR?QQGSK, A?JAGSK, ?LB K?LW MRFCP CJCKCLRQ ?PC ?JQM BGQQMJTCB GL RFC AWRMNJ?QK. .?LW KCR?@MJGA PC?ARGMLQ, GLAJSBGLE NPMRCGL QWLRFCQGQ, R?IC NJ?AC GL RFC AWRMNJ?QK.
/?< )L:C<LJ
5WNGA?JJW, RFC LSAJCSQ GQ RFC KMQR NPMKGLCLR MPE?LCJJC GL ? ACJJ ( Figure 4.8). 5FC nucleus (NJSP?J = LSAJCG) FMSQCQ RFC ACJJsQ %/" ?LB BGPCARQ RFC QWLRFCQGQ MD PG@MQMKCQ ?LB NPMRCGLQ. -CRsQ JMMI ?R GR GL KMPC BCR?GJ ( Figure 4.11).
118
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119
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120
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121
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122
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5FGQ RGEFR ?BFCPCLAC NPCTCLRQ K?RCPG?JQ DPMK JC?IGLE @CRUCCL RFC ACJJQ; RGEFR HSLARGMLQ ?PC RWNGA?JJW DMSLB GL CNGRFCJG?J RGQQSCQ RF?R JGLC GLRCPL?J MPE?LQ ?LB A?TGRGCQ, ?LB AMKNPGQC KMQR MD RFC QIGL. 'MP CV?KNJC, RFC RGEFR HSLARGMLQ MD RFC CNGRFCJG?J ACJJQ JGLGLE WMSP SPGL?PW @J?BBCP NPCTCLR SPGLC DPMK JC?IGLE MSR GLRM RFC CVRP?ACJJSJ?P QN?AC. D+824842+8 "JQM DMSLB MLJW GL ?LGK?J ACJJQ ?PC desmosomes, UFGAF ?AR JGIC QNMR UCJBQ @CRUCCL ?BH?ACLR CNGRFCJG?J ACJJQ (Figure 4.30). 4FMPR NPMRCGLQ A?JJCB A?BFCPGLQ GL RFC NJ?QK? KCK@P?LC AMLLCAR RM GLRCPKCBG?RC DGJ?KCLRQ RM APC?RC BCQKMQMKCQ. 5FC A?BFCPGLQ HMGL RUM ?BH?ACLR ACJJQ RMECRFCP ?LB K?GLR?GL RFC ACJJQ GL ? QFCCR-JGIC DMPK?RGML GL MPE?LQ ?LB RGQQSCQ RF?R QRPCRAF, JGIC RFC QIGL, FC?PR, ?LB KSQAJCQ.
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5M AMLBSAR ? TGPRS?J KGAPMQAMNW J?@ ?LB PCTGCU RFC N?PRQ MD ? ACJJ, UMPI RFPMSEF RFC QRCNQ MD RFGQ interactive assignment (http://openstaxcollege.org/l/microscopy_lab) .
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C@>?K D@:IFJ:FG< ?L GLQRPSKCLR RF?R K?ELGDGCQ ?L M@HCAR SQGLE ? @C?K TGQG@JC JGEFR N?QQCB ?LB @CLR RFPMSEF ? JCLQ QWQRCK RM TGQS?JGXC ? QNCAGKCL CPJFJFD< MPE?LCJJC GL ?L ?LGK?J ACJJ RF?R DSLARGMLQ ?Q RFC ACJJs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sQ K?GL CLCPEW-A?PPWGLE KMJCASJC EL:C<8I <EM<CFG< BMS@JC-KCK@P?LC QRPSARSPC RF?R AMLQRGRSRCQ RFC MSRCPKMQR NMPRGML MD RFC LSAJCSQ EL:C<F@; ACLRP?J N?PR MD ? NPMI?PWMRGA ACJJ GL UFGAF RFC AFPMKMQMKC GQ DMSLB EL:C<FCLJ B?PIJW QR?GLGLE @MBW UGRFGL RFC LSAJCSQ RF?R GQ PCQNMLQG@JC DMP ?QQCK@JGLE RFC QS@SLGRQ MD RFC PG@MQMKCQ EL:C<FGC8JD QCKG-QMJGB DJSGB GLQGBC RFC LSAJCSQ RF?R AMLR?GLQ RFC AFPMK?RGL ?LB LSAJCMJSQ EL:C<LJ ACJJ MPE?LCJJC RF?R FMSQCQ RFC ACJJs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
138
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" ACJJ GQ RFC QK?JJCQR SLGR MD JGDC. .MQR ACJJQ ?PC QM RGLW RF?R RFCW A?LLMR @C QCCL UGRF RFC L?ICB CWC. 5FCPCDMPC, QAGCLRGQRQ SQC KGAPMQAMNCQ RM QRSBW ACJJQ. &JCARPML KGAPMQAMNCQ NPMTGBC FGEFCP K?ELGDGA?RGML, FGEFCP PCQMJSRGML, ?LB KMPC BCR?GJ RF?L JGEFR KGAPMQAMNCQ. 5FC SLGDGCB ACJJ RFCMPW QR?RCQ RF?R ?JJ MPE?LGQKQ ?PC AMKNMQCB MD MLC MP KMPC ACJJQ, RFC ACJJ GQ RFC @?QGA SLGR MD JGDC, ?LB LCU ACJJQ ?PGQC DPMK CVGQRGLE ACJJQ.
4.2 +IFB8IPFK@: C<CCJ

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4.3 LB8IPFK@: C<CCJ

-GIC ? NPMI?PWMRGA ACJJ, ? CSI?PWMRGA ACJJ F?Q ? NJ?QK? KCK@P?LC, AWRMNJ?QK, ?LB PG@MQMKCQ, @SR ? CSI?PWMRGA ACJJ GQ RWNGA?JJW J?PECP RF?L ? NPMI?PWMRGA ACJJ, F?Q ? RPSC LSAJCSQ (KC?LGLE GRQ %/" GQ QSPPMSLBCB @W ? KCK@P?LC), ?LB F?Q MRFCP KCK@P?LC-@MSLB MPE?LCJJCQ RF?R ?JJMU DMP AMKN?PRKCLR?JGX?RGML MD DSLARGMLQ. 5FC NJ?QK? KCK@P?LC GQ ? NFMQNFMJGNGB @GJ?WCP CK@CBBCB UGRF NPMRCGLQ. 5FC LSAJCSQs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4.5 /?< CPKFJB<C<KFE

5FC AWRMQICJCRML F?Q RFPCC BGDDCPCLR RWNCQ MD NPMRCGL CJCKCLRQ. 'PMK L?PPMUCQR RM UGBCQR, RFCW ?PC RFC KGAPMDGJ?KCLRQ (?ARGL DGJ?KCLRQ), GLRCPKCBG?RC DGJ?KCLRQ, ?LB KGAPMRS@SJCQ. .GAPMDGJ?KCLRQ ?PC MDRCL ?QQMAG?RCB UGRF KWMQGL. 5FCW NPMTGBC PGEGBGRW ?LB QF?NC RM RFC ACJJ ?LB D?AGJGR?RC ACJJSJ?P KMTCKCLRQ. *LRCPKCBG?RC DGJ?KCLRQ @C?P RCLQGML ?LB ?LAFMP RFC LSAJCSQ ?LB MRFCP MPE?LCJJCQ GL NJ?AC. .GAPMRS@SJCQ FCJN RFC ACJJ PCQGQR AMKNPCQQGML, QCPTC ?Q RP?AIQ DMP KMRMP NPMRCGLQ RF?R KMTC TCQGAJCQ RFPMSEF RFC ACJJ, ?LB NSJJ PCNJGA?RCB AFPMKMQMKCQ RM MNNMQGRC CLBQ MD ? BGTGBGLE ACJJ. 5FCW ?PC ?JQM RFC QRPSARSP?J CJCKCLR MD ACLRPGMJCQ, DJ?ECJJ?, ?LB AGJG?.
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1. Figure 4.7 1PMI?PWMRGA ACJJQ ?PC KSAF QK?JJCP RF?L CSI?PWMRGA ACJJQ. 8F?R ?BT?LR?ECQ KGEFR QK?JJ ACJJ QGXC AMLDCP ML ? ACJJ 8F?R ?BT?LR?ECQ KGEFR J?PEC ACJJ QGXC F?TC 2. Figure 4.8 *D RFC LSAJCMJSQ UCPC LMR ?@JC RM A?PPW MSR GRQ DSLARGML, UF?R MRFCP ACJJSJ?P MPE?LCJJCQ UMSJB @C ?DDCARCB 3. Figure 4.18 *D ? NCPGNFCP?J KCK@P?LC NPMRCGL UCPC QWLRFCQGXCB GL RFC JSKCL (GLQGBC) MD RFC &3, UMSJB GR CLB SN ML RFC GLQGBC MP MSRQGBC MD RFC NJ?QK? KCK@P?LC
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4. 8FCL TGCUGLE ? QNCAGKCL RFPMSEF ? JGEFR KGAPMQAMNC, QAGCLRGQRQ SQC >>>>>>>> RM BGQRGLESGQF RFC GLBGTGBS?J AMKNMLCLRQ MD ACJJQ. >. ? @C?K MD CJCARPMLQ ?. P?BGM?ARGTC GQMRMNCQ @. QNCAG?J QR?GLQ A. FGEF RCKNCP?RSPCQ 5. 5FC >>>>>>>> GQ RFC @?QGA SLGR MD JGDC. >. ?. @. A. MPE?LGQK ACJJ RGQQSC MPE?L ?. NPMBSACB BSPGLE RFCGP MVGB?RGML PC?ARGMLQ @. GLAMPNMP?RCB GLRM RFCGP KCK@P?LCQ A. ? AMD?ARMP DMP RFC MPE?LCJJCQs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
140
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Diffusion GQ ? N?QQGTC NPMACQQ MD RP?LQNMPR. " QGLEJC QS@QR?LAC RCLBQ RM KMTC DPMK ?L ?PC? MD FGEF AMLACLRP?RGML RM ?L ?PC? MD JMU AMLACLRP?RGML SLRGJ RFC AMLACLRP?RGML GQ COS?J ?APMQQ ? QN?AC. :MS ?PC D?KGJG?P UGRF BGDDSQGML MD QS@QR?LACQ RFPMSEF RFC ?GP. 'MP CV?KNJC, RFGLI ?@MSR QMKCMLC MNCLGLE ? @MRRJC MD ?KKMLG? GL ? PMMK DGJJCB UGRF NCMNJC. 5FC ?KKMLG? E?Q GQ ?R GRQ FGEFCQR AMLACLRP?RGML GL RFC @MRRJC; GRQ JMUCQR AMLACLRP?RGML GQ ?R RFC CBECQ MD RFC PMMK. 5FC ?KKMLG? T?NMP UGJJ BGDDSQC, MP QNPC?B ?U?W, DPMK RFC @MRRJC, ?LB EP?BS?JJW, KMPC ?LB KMPC NCMNJC UGJJ QKCJJ RFC ?KKMLG? ?Q GR QNPC?BQ. .?RCPG?JQ KMTC UGRFGL RFC ACJJsQ AWRMQMJ @W BGDDSQGML, ?LB ACPR?GL K?RCPG?JQ KMTC RFPMSEF RFC NJ?QK? KCK@P?LC @W BGDDSQGML (Figure 5.8). %GDDSQGML CVNCLBQ LM CLCPEW. 0L RFC AMLRP?PW, AMLACLRP?RGML EP?BGCLRQ ?PC ? DMPK MD NMRCLRG?J CLCPEW, BGQQGN?RCB ?Q RFC EP?BGCLR GQ CJGKGL?RCB.
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Osmosis GQ RFC KMTCKCLR MD U?RCP RFPMSEF ? QCKGNCPKC?@JC KCK@P?LC ?AAMPBGLE RM RFC AMLACLRP?RGML EP?BGCLR MD U?RCP ?APMQQ RFC KCK@P?LC, UFGAF GQ GLTCPQCJW NPMNMPRGML?J RM RFC AMLACLRP?RGML MD QMJSRCQ. 8FGJC BGDDSQGML RP?LQNMPRQ K?RCPG?J ?APMQQ KCK@P?LCQ ?LB UGRFGL ACJJQ, MQKMQGQ RP?LQNMPRQ 653@ >(;,9 ?APMQQ ? KCK@P?LC ?LB RFC KCK@P?LC JGKGRQ RFC BGDDSQGML MD QMJSRCQ GL RFC U?RCP. /MR QSPNPGQGLEJW, RFC ?OS?NMPGLQ RF?R D?AGJGR?RC U?RCP KMTCKCLR NJ?W ? J?PEC PMJC GL MQKMQGQ, KMQR NPMKGLCLRJW GL PCB @JMMB ACJJQ ?LB RFC KCK@P?LCQ MD IGBLCW RS@SJCQ. M+).'3/82 0QKMQGQ GQ ? QNCAG?J A?QC MD BGDDSQGML. 8?RCP, JGIC MRFCP QS@QR?LACQ, KMTCQ DPMK ?L ?PC? MD FGEF AMLACLRP?RGML RM MLC MD JMU AMLACLRP?RGML. "L M@TGMSQ OSCQRGML GQ UF?R K?ICQ U?RCP KMTC ?R ?JJ *K?EGLC ? @C?ICP UGRF ? QCKGNCPKC?@JC KCK@P?LC QCN?P?RGLE RFC RUM QGBCQ MP F?JTCQ (Figure 5.11). 0L @MRF QGBCQ MD RFC KCK@P?LC RFC U?RCP JCTCJ GQ RFC Q?KC, @SR RFCPC ?PC BGDDCPCLR AMLACLRP?RGMLQ MD ? BGQQMJTCB QS@QR?LAC, MP solute, RF?R A?LLMR APMQQ RFC KCK@P?LC (MRFCPUGQC RFC AMLACLRP?RGMLQ ML C?AF QGBC UMSJB @C @?J?LACB @W RFC QMJSRC APMQQGLE RFC KCK@P?LC). *D RFC TMJSKC MD RFC QMJSRGML ML @MRF QGBCQ MD RFC KCK@P?LC GQ RFC Q?KC, @SR RFC AMLACLRP?RGMLQ MD QMJSRC ?PC BGDDCPCLR, RFCL RFCPC ?PC BGDDCPCLR ?KMSLRQ MD U?RCP, RFC QMJTCLR, ML CGRFCP QGBC MD RFC KCK@P?LC.
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5FC NPGK?PW ?ARGTC RP?LQNMPR RF?R DSLARGMLQ UGRF RFC ?ARGTC RP?LQNMPR MD QMBGSK ?LB NMR?QQGSK ?JJMUQ QCAMLB?PW ?ARGTC RP?LQNMPR RM MAASP. 5FC QCAMLB RP?LQNMPR KCRFMB GQ QRGJJ AMLQGBCPCB ?ARGTC @CA?SQC GR BCNCLBQ ML RFC SQC MD CLCPEW ?Q BMCQ NPGK?PW RP?LQNMPR (Figure 5.18).
!@>LI< 5.18 0OFJ>OV >@QFSB QO>KPMLOQ JLSBP FLKP >@OLPP > JBJ?O>KB, @OB>QFKD >K BIB@QOL@EBJF@>I DO>AFBKQ (BIB@QOLDBKF@ QO>KPMLOQ). (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
0LC MD RFC KMQR GKNMPR?LR NSKNQ GL ?LGK?JQ ACJJQ GQ RFC QMBGSK-NMR?QQGSK NSKN (/?+-,+ "51?QC), UFGAF K?GLR?GLQ RFC CJCARPMAFCKGA?J EP?BGCLR (?LB RFC AMPPCAR AMLACLRP?RGMLQ MD /?+ ?LB ,+) GL JGTGLE ACJJQ. 5FC QMBGSK-NMR?QQGSK NSKN KMTCQ ,+ GLRM RFC ACJJ UFGJC KMTGLE /?+ MSR ?R RFC Q?KC RGKC, ?R ? P?RGM MD RFPCC /?+ DMP CTCPW RUM ,+ GMLQ KMTCB GL. 5FC /?+-,+ "51?QC CVGQRQ GL RUM DMPKQ, BCNCLBGLE ML GRQ MPGCLR?RGML RM RFC GLRCPGMP MP CVRCPGMP MD RFC ACJJ ?LB GRQ ?DDGLGRW DMP CGRFCP QMBGSK MP NMR?QQGSK GMLQ. 5FC NPMACQQ AMLQGQRQ MD RFC DMJJMUGLE QGV QRCNQ. 1. 8GRF RFC CLXWKC MPGCLRCB RMU?PBQ RFC GLRCPGMP MD RFC ACJJ, RFC A?PPGCP F?Q ? FGEF ?DDGLGRW DMP QMBGSK GMLQ. 5FPCC GMLQ @GLB RM RFC NPMRCGL. 2. "51 GQ FWBPMJWXCB @W RFC NPMRCGL A?PPGCP ?LB ? JMU-CLCPEW NFMQNF?RC EPMSN ?RR?AFCQ RM GR. 3. "Q ? PCQSJR, RFC A?PPGCP AF?LECQ QF?NC ?LB PC-MPGCLRQ GRQCJD RMU?PBQ RFC CVRCPGMP MD RFC KCK@P?LC. 5FC NPMRCGLsQ ?DDGLGRW DMP QMBGSK BCAPC?QCQ ?LB RFC RFPCC QMBGSK GMLQ JC?TC RFC A?PPGCP. 4. 5FC QF?NC AF?LEC GLAPC?QCQ RFC A?PPGCPsQ ?DDGLGRW DMP NMR?QQGSK GMLQ, ?LB RUM QSAF GMLQ ?RR?AF RM RFC NPMRCGL. 4S@QCOSCLRJW, RFC JMU-CLCPEW NFMQNF?RC EPMSN BCR?AFCQ DPMK RFC A?PPGCP. 5. 8GRF RFC NFMQNF?RC EPMSN PCKMTCB ?LB NMR?QQGSK GMLQ ?RR?AFCB, RFC A?PPGCP NPMRCGL PCNMQGRGMLQ GRQCJD RMU?PBQ RFC GLRCPGMP MD RFC ACJJ. 6. 5FC A?PPGCP NPMRCGL, GL GRQ LCU AMLDGESP?RGML, F?Q ? BCAPC?QCB ?DDGLGRW DMP NMR?QQGSK, ?LB RFC RUM GMLQ ?PC PCJC?QCB GLRM RFC AWRMNJ?QK. 5FC NPMRCGL LMU F?Q ? FGEFCP ?DDGLGRW DMP QMBGSK GMLQ, ?LB RFC NPMACQQ QR?PRQ ?E?GL. 4CTCP?J RFGLEQ F?TC F?NNCLCB ?Q ? PCQSJR MD RFGQ NPMACQQ. "R RFGQ NMGLR, RFCPC ?PC KMPC QMBGSK GMLQ MSRQGBC MD RFC ACJJ RF?L GLQGBC ?LB KMPC NMR?QQGSK GMLQ GLQGBC RF?L MSR. 'MP CTCPW RFPCC GMLQ MD QMBGSK RF?R KMTC MSR, RUM GMLQ MD NMR?QQGSK KMTC GL. 5FGQ PCQSJRQ GL RFC GLRCPGMP @CGLE QJGEFRJW KMPC LCE?RGTC PCJ?RGTC RM RFC CVRCPGMP. 5FGQ BGDDCPCLAC GL AF?PEC GQ GKNMPR?LR GL APC?RGLE RFC AMLBGRGMLQ LCACQQ?PW DMP
160
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RFC QCAMLB?PW NPMACQQ. 5FC QMBGSK-NMR?QQGSK NSKN GQ, RFCPCDMPC, ?L electrogenic pump (? NSKN RF?R APC?RCQ ? AF?PEC GK@?J?LAC), APC?RGLE ?L CJCARPGA?J GK@?J?LAC ?APMQQ RFC KCK@P?LC ?LB AMLRPG@SRGLE RM RFC KCK@P?LC NMRCLRG?J.
7GQGR RFC site (http://openstaxcollege.org/l/Na_K_ATPase) RM QCC ? QGKSJ?RGML MD ?ARGTC RP?LQNMPR GL ? QMBGSK-NMR?QQGSK "51?QC.
.<:FE;8IP A:K@M< /I8EJGFIK (CF-KI8EJGFIK)

4CAMLB?PW ?ARGTC RP?LQNMPR @PGLEQ QMBGSK GMLQ, ?LB NMQQG@JW MRFCP AMKNMSLBQ, GLRM RFC ACJJ. "Q QMBGSK GML AMLACLRP?RGMLQ @SGJB MSRQGBC MD RFC NJ?QK? KCK@P?LC @CA?SQC MD RFC ?ARGML MD RFC NPGK?PW ?ARGTC RP?LQNMPR NPMACQQ, ?L CJCARPMAFCKGA?J EP?BGCLR GQ APC?RCB. *D ? AF?LLCJ NPMRCGL CVGQRQ ?LB GQ MNCL, RFC QMBGSK GMLQ UGJJ @C NSJJCB RFPMSEF RFC KCK@P?LC. 5FGQ KMTCKCLR GQ SQCB RM RP?LQNMPR MRFCP QS@QR?LACQ RF?R A?L ?RR?AF RFCKQCJTCQ RM RFC RP?LQNMPR NPMRCGL RFPMSEF RFC KCK@P?LC (Figure 5.19). .?LW ?KGLM ?AGBQ, ?Q UCJJ ?Q EJSAMQC, CLRCP ? ACJJ RFGQ U?W. 5FGQ QCAMLB?PW NPMACQQ GQ ?JQM SQCB RM QRMPC FGEF-CLCPEW FWBPMECL GMLQ GL RFC KGRMAFMLBPG? MD NJ?LR ?LB ?LGK?J ACJJQ DMP RFC NPMBSARGML MD "51. 5FC NMRCLRG?J CLCPEW RF?R ?AASKSJ?RCQ GL RFC QRMPCB FWBPMECL GMLQ GQ RP?LQJ?RCB GLRM IGLCRGA CLCPEW ?Q RFC GMLQ QSPEC RFPMSEF RFC AF?LLCJ NPMRCGL "51 QWLRF?QC, ?LB RF?R CLCPEW GQ SQCB RM AMLTCPR "%1 GLRM "51.
!@>LI< 5.19 !K BIB@QOL@EBJF@>I DO>AFBKQ, @OB>QBA ?V MOFJ>OV >@QFSB QO>KPMLOQ, @>K JLSB LQEBO PR?PQ>K@BP >D>FKPQ QEBFO @LK@BKQO>QFLK DO>AFBKQP, > MOL@BPP @>IIBA @L-QO>KPMLOQ LO PB@LKA>OV >@QFSB QO>KPMLOQ. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
)C QEB M( LRQPFAB QEB @BII AB@OB>PBP, TLRIA VLR BUMB@Q QEB >JLRKQ LC >JFKL >@FAP QO>KPMLOQBA FKQL QEB @BII QL FK@OB>PB LO AB@OB>PB
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161
5.4 R BLCB /I8EJGFIK

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C CLBMAWRMQGQ, GLAJSBGLE NF?EMAWRMQGQ, NGLMAWRMQGQ, ?LB PCACNRMP-KCBG?RCB CLBMAWRMQGQ z 6LBCPQR?LB RFC NPMACQQ MD CVMAWRMQGQ *L ?BBGRGML RM KMTGLE QK?JJ GMLQ ?LB KMJCASJCQ RFPMSEF RFC KCK@P?LC, ACJJQ ?JQM LCCB RM PCKMTC ?LB R?IC GL J?PECP KMJCASJCQ ?LB N?PRGAJCQ (QCC Table 5.2 DMP CV?KNJCQ). 4MKC ACJJQ ?PC CTCL A?N?@JC MD CLESJDGLE CLRGPC SLGACJJSJ?P KGAPMMPE?LGQKQ. :MS KGEFR F?TC AMPPCARJW FWNMRFCQGXCB RF?R RFC SNR?IC ?LB PCJC?QC MD J?PEC N?PRGAJCQ @W RFC ACJJ PCOSGPCQ CLCPEW. " J?PEC N?PRGAJC, FMUCTCP, A?LLMR N?QQ RFPMSEF RFC KCK@P?LC, CTCL UGRF CLCPEW QSNNJGCB @W RFC ACJJ.
E;F:PKFJ@J
Endocytosis GQ ? RWNC MD ?ARGTC RP?LQNMPR RF?R KMTCQ N?PRGAJCQ, QSAF ?Q J?PEC KMJCASJCQ, N?PRQ MD ACJJQ, ?LB CTCL UFMJC ACJJQ, GLRM ? ACJJ. 5FCPC ?PC BGDDCPCLR T?PG?RGMLQ MD CLBMAWRMQGQ, @SR ?JJ QF?PC ? AMKKML AF?P?ARCPGQRGA: 5FC NJ?QK? KCK@P?LC MD RFC ACJJ GLT?EGL?RCQ, DMPKGLE ? NMAICR ?PMSLB RFC R?PECR N?PRGAJC. 5FC NMAICR NGLAFCQ MDD, PCQSJRGLE GL RFC N?PRGAJC @CGLE AMLR?GLCB GL ? LCUJW APC?RCB GLRP?ACJJSJ?P TCQGAJC DMPKCB DPMK RFC NJ?QK? KCK@P?LC. P.'-4)>948/8 1F?EMAWRMQGQ (RFC AMLBGRGML MD tACJJ C?RGLEu) GQ RFC NPMACQQ @W UFGAF J?PEC N?PRGAJCQ, QSAF ?Q ACJJQ MP PCJ?RGTCJW J?PEC N?PRGAJCQ, ?PC R?ICL GL @W ? ACJJ. 'MP CV?KNJC, UFCL KGAPMMPE?LGQKQ GLT?BC RFC FSK?L @MBW, ? RWNC MD UFGRC @JMMB ACJJ A?JJCB ? LCSRPMNFGJ UGJJ PCKMTC RFC GLT?BCPQ RFPMSEF RFGQ NPMACQQ, QSPPMSLBGLE ?LB CLESJDGLE RFC KGAPMMPE?LGQK, UFGAF GQ RFCL BCQRPMWCB @W RFC LCSRPMNFGJ (Figure 5.20).
!@>LI< 5.20 )K ME>DL@VQLPFP, QEB @BII JBJ?O>KB PROOLRKAP QEB M>OQF@IB >KA BKDRICP FQ. (@OBAFQ: ->OF>K> 2RFW 6FII>OB>I)
*L NPCN?P?RGML DMP NF?EMAWRMQGQ, ? NMPRGML MD RFC GLU?PB-D?AGLE QSPD?AC MD RFC NJ?QK? KCK@P?LC @CAMKCQ AM?RCB UGRF ? NPMRCGL A?JJCB clathrin, UFGAF QR?@GJGXCQ RFGQ QCARGML MD RFC KCK@P?LC. 5FC AM?RCB NMPRGML MD RFC KCK@P?LC RFCL CVRCLBQ DPMK RFC @MBW MD RFC ACJJ ?LB QSPPMSLBQ RFC N?PRGAJC, CTCLRS?JJW CLAJMQGLE GR. 0LAC RFC TCQGAJC AMLR?GLGLE RFC N?PRGAJC GQ CLAJMQCB UGRFGL RFC ACJJ, RFC AJ?RFPGL BGQCLE?ECQ DPMK RFC KCK@P?LC ?LB RFC TCQGAJC KCPECQ UGRF ? JWQMQMKC DMP RFC @PC?IBMUL MD RFC K?RCPG?J GL RFC LCUJW DMPKCB AMKN?PRKCLR (CLBMQMKC). 8FCL ?AACQQG@JC LSRPGCLRQ DPMK RFC BCEP?B?RGML MD RFC TCQGASJ?P AMLRCLRQ F?TC @CCL CVRP?ARCB, RFC LCUJW DMPKCB CLBMQMKC KCPECQ UGRF RFC NJ?QK? KCK@P?LC ?LB PCJC?QCQ GRQ AMLRCLRQ GLRM RFC CVRP?ACJJSJ?P DJSGB. 5FC CLBMQMK?J KCK@P?LC ?E?GL @CAMKCQ N?PR MD RFC NJ?QK? KCK@P?LC.
162
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P/34)>948/8 " T?PG?RGML MD CLBMAWRMQGQ GQ A?JJCB pinocytosis. 5FGQ JGRCP?JJW KC?LQ tACJJ BPGLIGLEu ?LB U?Q L?KCB ?R ? RGKC UFCL RFC ?QQSKNRGML U?Q RF?R RFC ACJJ U?Q NSPNMQCDSJJW R?IGLE GL CVRP?ACJJSJ?P DJSGB. *L PC?JGRW, RFGQ GQ ? NPMACQQ RF?R R?ICQ GL KMJCASJCQ, GLAJSBGLE U?RCP, UFGAF RFC ACJJ LCCBQ DPMK RFC CVRP?ACJJSJ?P DJSGB. 1GLMAWRMQGQ PCQSJRQ GL ? KSAF QK?JJCP TCQGAJC RF?L BMCQ NF?EMAWRMQGQ, ?LB RFC TCQGAJC BMCQ LMR LCCB RM KCPEC UGRF ? JWQMQMKC (Figure 5.21).
!@>LI< 5.21 )K MFKL@VQLPFP, QEB @BII JBJ?O>KB FKS>DFK>QBP, PROOLRKAP > PJ>II SLIRJB LC CIRFA, >KA MFK@EBP LCC. (@OBAFQ: ->OF>K> 2RFW 6FII>OB>I)
" T?PG?RGML MD NGLMAWRMQGQ GQ A?JJCB potocytosis. 5FGQ NPMACQQ SQCQ ? AM?RGLE NPMRCGL, A?JJCB caveolin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igure 5.22).
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163
!@>LI< 5.22 )K OB@BMQLO-JBAF>QBA BKAL@VQLPFP, RMQ>HB LC PR?PQ>K@BP ?V QEB @BII FP Q>ODBQBA QL > PFKDIB QVMB LC PR?PQ>K@B QE>Q ?FKAP QL QEB OB@BMQLO LK QEB BUQBOK>I PROC>@B LC QEB @BII JBJ?O>KB. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
*L receptor-mediated endocytosis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t@?Bu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
4CC PCACNRMP-KCBG?RCB CLBMAWRMQGQ GL ?ARGML, ?LB AJGAI ML BGDDCPCLR parts (http://openstaxcollege.org/ l/endocytosis) DMP ? DMASQCB ?LGK?RGML.
OF:PKFJ@J
5FC PCTCPQC NPMACQQ MD KMTGLE K?RCPG?J GLRM ? ACJJ GQ RFC NPMACQQ MD CVMAWRMQGQ. Exocytosis GQ RFC MNNMQGRC MD RFC NPMACQQCQ BGQASQQCB ?@MTC GL RF?R GRQ NSPNMQC GQ RM CVNCJ K?RCPG?J DPMK RFC ACJJ GLRM RFC CVRP?ACJJSJ?P DJSGB. 8?QRC K?RCPG?J GQ CLTCJMNCB GL ? KCK@P?LC ?LB DSQCQ UGRF RFC GLRCPGMP MD RFC NJ?QK? KCK@P?LC. 5FGQ DSQGML MNCLQ RFC KCK@P?LMSQ CLTCJMNC ML RFC CVRCPGMP MD RFC ACJJ, ?LB RFC U?QRC K?RCPG?J GQ CVNCJJCB GLRM RFC CVRP?ACJJSJ?P QN?AC (Figure 5.23). 0RFCP CV?KNJCQ MD ACJJQ PCJC?QGLE KMJCASJCQ TG? CVMAWRMQGQ
164
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GLAJSBC RFC QCAPCRGML MD NPMRCGLQ MD RFC CVRP?ACJJSJ?P K?RPGV ?LB QCAPCRGML MD LCSPMRP?LQKGRRCPQ GLRM RFC QWL?NRGA AJCDR @W QWL?NRGA TCQGAJCQ.
!@>LI< 5.23 )K BUL@VQLPFP, SBPF@IBP @LKQ>FKFKD PR?PQ>K@BP CRPB TFQE QEB MI>PJ> JBJ?O>KB. 4EB @LKQBKQP >OB QEBK OBIB>PBA QL QEB BUQBOFLO LC QEB @BII. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
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5.4 BLCB /I8EJGFIK

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180
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188
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189
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190
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191
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198
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5FC ?BBGRGML MD ? NFMQNF?RC EPMSN RM ? KMJCASJC PCOSGPCQ CLCPEW. 1FMQNF?RC EPMSNQ ?PC LCE?RGTCJW AF?PECB ?LB RFSQ PCNCJ MLC ?LMRFCP UFCL RFCW ?PC ?PP?LECB GL QCPGCQ, ?Q RFCW ?PC GL "%1 ?LB "51. 5FGQ PCNSJQGML K?ICQ RFC "%1 ?LB "51 KMJCASJCQ GLFCPCLRJW SLQR?@JC. 5FC PCJC?QC MD MLC MP RUM NFMQNF?RC EPMSNQ DPMK "51, ? NPMACQQ A?JJCB dephosphorylation, PCJC?QCQ CLCPEW. E3+7-> ,742 A"P )WBPMJWQGQ GQ RFC NPMACQQ MD @PC?IGLE AMKNJCV K?APMKMJCASJCQ ?N?PR. %SPGLE FWBPMJWQGQ, U?RCP GQ QNJGR, MP JWQCB, ?LB RFC PCQSJRGLE FWBPMECL ?RMK ()+) ?LB ? FWBPMVWJ EPMSN (0)-) ?PC ?BBCB RM RFC J?PECP KMJCASJC. 5FC FWBPMJWQGQ MD "51 NPMBSACQ "%1, RMECRFCP UGRF ?L GLMPE?LGA NFMQNF?RC GML (1G), ?LB RFC PCJC?QC MD DPCC CLCPEW. 5M A?PPW MSR JGDC NPMACQQCQ, "51 GQ AMLRGLSMSQJW @PMICL BMUL GLRM "%1, ?LB JGIC ? PCAF?PEC?@JC @?RRCPW, "%1 GQ AMLRGLSMSQJW PCECLCP?RCB GLRM "51 @W RFC PC?RR?AFKCLR MD ? RFGPB NFMQNF?RC EPMSN. 8?RCP, UFGAF U?Q @PMICL BMUL GLRM GRQ FWBPMECL ?RMK ?LB FWBPMVWJ EPMSN BSPGLE "51 FWBPMJWQGQ, GQ PCECLCP?RCB UFCL ? RFGPB NFMQNF?RC GQ ?BBCB RM RFC "%1 KMJCASJC, PCDMPKGLE "51. 0@TGMSQJW, CLCPEW KSQR @C GLDSQCB GLRM RFC QWQRCK RM PCECLCP?RC "51. 8FCPC BMCQ RFGQ CLCPEW AMKC DPMK *L LC?PJW CTCPW JGTGLE RFGLE ML C?PRF, RFC CLCPEW AMKCQ DPMK RFC KCR?@MJGQK MD EJSAMQC. *L RFGQ U?W, "51 GQ ? BGPCAR JGLI @CRUCCL RFC JGKGRCB QCR MD CVCPEMLGA N?RFU?WQ MD EJSAMQC A?R?@MJGQK ?LB RFC KSJRGRSBC MD CLBCPEMLGA N?RFU?WQ RF?R NMUCP JGTGLE ACJJQ. P.485.47>1'9/43 3CA?JJ RF?R, GL QMKC AFCKGA?J PC?ARGMLQ, CLXWKCQ K?W @GLB RM QCTCP?J QS@QRP?RCQ RF?R PC?AR UGRF C?AF MRFCP ML RFC CLXWKC, DMPKGLE ?L GLRCPKCBG?RC AMKNJCV. "L GLRCPKCBG?RC AMKNJCV GQ ? RCKNMP?PW QRPSARSPC, ?LB GR ?JJMUQ MLC MD RFC QS@QRP?RCQ (QSAF ?Q "51) ?LB PC?AR?LRQ RM KMPC PC?BGJW PC?AR UGRF C?AF MRFCP; GL PC?ARGMLQ GLTMJTGLE "51, "51 GQ MLC MD RFC QS@QRP?RCQ ?LB "%1 GQ ? NPMBSAR. %SPGLE ?L CLBCPEMLGA AFCKGA?J PC?ARGML, "51 DMPKQ ?L GLRCPKCBG?RC AMKNJCV UGRF RFC QS@QRP?RC ?LB CLXWKC GL RFC PC?ARGML. 5FGQ GLRCPKCBG?RC AMKNJCV ?JJMUQ RFC "51 RM RP?LQDCP GRQ RFGPB NFMQNF?RC EPMSN, UGRF GRQ CLCPEW, RM RFC QS@QRP?RC, ? NPMACQQ A?JJCB NFMQNFMPWJ?RGML. Phosphorylation PCDCPQ RM RFC ?BBGRGML MD RFC NFMQNF?RC (Y1). 5FGQ GQ GJJSQRP?RCB @W RFC DMJJMUGLE ECLCPGA PC?ARGML:
A + enzyme + ATP P A enzyme B + enzyme + ADP + phosphate ion
8FCL RFC GLRCPKCBG?RC AMKNJCV @PC?IQ ?N?PR, RFC CLCPEW GQ SQCB RM KMBGDW RFC QS@QRP?RC ?LB AMLTCPR GR GLRM ? NPMBSAR MD RFC PC?ARGML. 5FC "%1 KMJCASJC ?LB ? DPCC NFMQNF?RC GML ?PC PCJC?QCB GLRM RFC KCBGSK ?LB ?PC ?T?GJ?@JC DMP PCAWAJGLE RFPMSEF ACJJ KCR?@MJGQK. !:(897'9+ P.485.47>1'9/43 "51 GQ ECLCP?RCB RFPMSEF RUM KCAF?LGQKQ BSPGLE RFC @PC?IBMUL MD EJSAMQC. " DCU "51 KMJCASJCQ ?PC ECLCP?RCB (RF?R GQ, PCECLCP?RCB DPMK "%1) ?Q ? BGPCAR PCQSJR MD RFC AFCKGA?J PC?ARGMLQ RF?R MAASP GL RFC A?R?@MJGA N?RFU?WQ. " NFMQNF?RC EPMSN GQ PCKMTCB DPMK ?L GLRCPKCBG?RC PC?AR?LR GL RFC N?RFU?W, ?LB RFC DPCC CLCPEW MD RFC PC?ARGML GQ SQCB RM ?BB RFC RFGPB NFMQNF?RC RM ?L ?T?GJ?@JC "%1 KMJCASJC, NPMBSAGLE "51 (Figure 7.4). 5FGQ TCPW BGPCAR KCRFMB MD NFMQNFMPWJ?RGML GQ A?JJCB substrate-level phosphorylation.
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199
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O=/*'9/;+ P.485.47>1'9/43 .MQR MD RFC "51 ECLCP?RCB BSPGLE EJSAMQC A?R?@MJGQK, FMUCTCP, GQ BCPGTCB DPMK ? KSAF KMPC AMKNJCV NPMACQQ, AFCKGMQKMQGQ, UFGAF R?ICQ NJ?AC GL KGRMAFMLBPG? (Figure 7.5) UGRFGL ? CSI?PWMRGA ACJJ MP RFC NJ?QK? KCK@P?LC MD ? NPMI?PWMRGA ACJJ. Chemiosmosis, ? NPMACQQ MD "51 NPMBSARGML GL ACJJSJ?P KCR?@MJGQK, GQ SQCB RM ECLCP?RC 90 NCPACLR MD RFC "51 K?BC BSPGLE EJSAMQC A?R?@MJGQK ?LB GQ ?JQM RFC KCRFMB SQCB GL RFC JGEFR PC?ARGMLQ MD NFMRMQWLRFCQGQ RM F?PLCQQ RFC CLCPEW MD QSLJGEFR. 5FC NPMBSARGML MD "51 SQGLE RFC NPMACQQ MD AFCKGMQKMQGQ GQ A?JJCB oxidative phosphorylation @CA?SQC MD RFC GLTMJTCKCLR MD MVWECL GL RFC NPMACQQ.
!@>LI< 7.5 )K BRH>OVLQBP, LUFA>QFSB MELPMELOVI>QFLK Q>HBP MI>@B FK JFQL@ELKAOF>. )K MOLH>OVLQBP, QEFP MOL@BPP Q>HBP MI>@B FK QEB MI>PJ> JBJ?O>KB. (#OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
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200
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC MTCP?JJ PCQSJR GL RCPKQ MD KMJCASJCQ NPMBSACB GL RFC @PC?IBMUL MD EJSAMQC @W EJWAMJWQGQ z $MKN?PC RFC MSRNSR MD EJWAMJWQGQ GL RCPKQ MD "51 KMJCASJCQ ?LB /"%) KMJCASJCQ NPMBSACB :MS F?TC PC?B RF?R LC?PJW ?JJ MD RFC CLCPEW SQCB @W JGTGLE ACJJQ AMKCQ RM RFCK GL RFC @MLBQ MD RFC QSE?P, EJSAMQC. Glycolysis GQ RFC DGPQR QRCN GL RFC @PC?IBMUL MD EJSAMQC RM CVRP?AR CLCPEW DMP ACJJSJ?P KCR?@MJGQK. /C?PJW ?JJ JGTGLE MPE?LGQKQ A?PPW MSR EJWAMJWQGQ ?Q N?PR MD RFCGP KCR?@MJGQK. 5FC NPMACQQ BMCQ LMR SQC MVWECL ?LB GQ RFCPCDMPC anaerobic. (JWAMJWQGQ R?ICQ NJ?AC GL RFC AWRMNJ?QK MD @MRF NPMI?PWMRGA ?LB CSI?PWMRGA ACJJQ. (JSAMQC CLRCPQ FCRCPMRPMNFGA ACJJQ GL RUM U?WQ. 0LC KCRFMB GQ RFPMSEF QCAMLB?PW ?ARGTC RP?LQNMPR GL UFGAF RFC RP?LQNMPR R?ICQ NJ?AC ?E?GLQR RFC EJSAMQC AMLACLRP?RGML EP?BGCLR. 5FC MRFCP KCAF?LGQK SQCQ ? EPMSN MD GLRCEP?J NPMRCGLQ A?JJCB (-65 NPMRCGLQ, ?JQM ILMUL ?Q EJSAMQC RP?LQNMPRCP NPMRCGLQ. 5FCQC RP?LQNMPRCPQ ?QQGQR GL RFC D?AGJGR?RCB BGDDSQGML MD EJSAMQC. (JWAMJWQGQ @CEGLQ UGRF RFC QGV A?P@ML PGLE-QF?NCB QRPSARSPC MD ? QGLEJC EJSAMQC KMJCASJC ?LB CLBQ UGRF RUM KMJCASJCQ MD ? RFPCC-A?P@ML QSE?P A?JJCB pyruvate. (JWAMJWQGQ AMLQGQRQ MD RUM BGQRGLAR NF?QCQ. 5FC DGPQR N?PR MD RFC EJWAMJWQGQ N?RFU?W RP?NQ RFC EJSAMQC KMJCASJC GL RFC ACJJ ?LB SQCQ CLCPEW RM KMBGDW GR QM RF?R RFC QGV-A?P@ML QSE?P KMJCASJC A?L @C QNJGR CTCLJW GLRM RFC RUM RFPCC-A?P@ML KMJCASJCQ. 5FC QCAMLB N?PR MD EJWAMJWQGQ CVRP?ARQ CLCPEW DPMK RFC KMJCASJCQ ?LB QRMPCQ GR GL RFC DMPK MD "51 ?LB /"%), RFC PCBSACB DMPK MD /"%.
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4RCN 1. 5FC DGPQR QRCN GL EJWAMJWQGQ (Figure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isomerase GQ ?L CLXWKC RF?R A?R?JWXCQ RFC AMLTCPQGML MD ? KMJCASJC GLRM MLC MD GRQ GQMKCPQ. (5FGQ AF?LEC DPMK NFMQNFMEJSAMQC RM NFMQNFMDPSARMQC ?JJMUQ RFC CTCLRS?J QNJGR MD RFC QSE?P GLRM RUM RFPCC-A?P@ML KMJCASJCQ.). 4RCN 3. 5FC RFGPB QRCN GQ RFC NFMQNFMPWJ?RGML MD DPSARMQC-6-NFMQNF?RC, A?R?JWXCB @W RFC CLXWKC NFMQNFMDPSARMIGL?QC. " QCAMLB "51 KMJCASJC BML?RCQ ? FGEF-CLCPEW NFMQNF?RC RM DPSARMQC-6-NFMQNF?RC, NPMBSAGLE DPSARMQC-1,6-@GQNFMQNF?RC. *L RFGQ N?RFU?W, NFMQNFMDPSARMIGL?QC GQ ? P?RC-JGKGRGLE CLXWKC. *R GQ ?ARGTC UFCL RFC AMLACLRP?RGML MD "%1 GQ FGEF; GR GQ JCQQ ?ARGTC UFCL "%1 JCTCJQ ?PC JMU ?LB RFC AMLACLRP?RGML MD "51 GQ FGEF. 5FSQ, GD RFCPC GQ tQSDDGAGCLRu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
!@>LI< 7.6 4EB CFOPQ E>IC LC DIV@LIVPFP RPBP QTL !40 JLIB@RIBP FK QEB MELPMELOVI>QFLK LC DIR@LPB, TEF@E FP QEBK PMIFQ FKQL QTL QEOBB-@>O?LK JLIB@RIBP.
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201
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4M D?P, EJWAMJWQGQ F?Q AMQR RFC ACJJ RUM "51 KMJCASJCQ ?LB NPMBSACB RUM QK?JJ, RFPCC-A?P@ML QSE?P KMJCASJCQ. #MRF MD RFCQC KMJCASJCQ UGJJ NPMACCB RFPMSEF RFC QCAMLB F?JD MD RFC N?RFU?W, ?LB QSDDGAGCLR CLCPEW UGJJ @C CVRP?ARCB RM N?W @?AI RFC RUM "51 KMJCASJCQ SQCB ?Q ?L GLGRG?J GLTCQRKCLR ?LB NPMBSAC ? NPMDGR DMP RFC ACJJ MD RUM ?BBGRGML?J "51 KMJCASJCQ ?LB RUM CTCL FGEFCP-CLCPEW /"%) KMJCASJCQ. 4RCN 6. 5FC QGVRF QRCN GL EJWAMJWQGQ (Figure 7.7) MVGBGXCQ RFC QSE?P (EJWACP?JBCFWBC-3-NFMQNF?RC), CVRP?ARGLE FGEF-CLCPEW CJCARPMLQ, UFGAF ?PC NGAICB SN @W RFC CJCARPML A?PPGCP /"%+, NPMBSAGLE /"%). 5FC QSE?P GQ RFCL NFMQNFMPWJ?RCB @W RFC ?BBGRGML MD ? QCAMLB NFMQNF?RC EPMSN, NPMBSAGLE 1,3-@GQNFMQNFMEJWACP?RC. /MRC RF?R RFC QCAMLB NFMQNF?RC EPMSN BMCQ LMR PCOSGPC ?LMRFCP "51 KMJCASJC.
!@>LI< 7.7 4EB PB@LKA E>IC LC DIV@LIVPFP FKSLISBP MELPMELOVI>QFLK TFQELRQ !40 FKSBPQJBKQ (PQBM 6) >KA MOLAR@BP QTL .!$( >KA CLRO !40 JLIB@RIBP MBO DIR@LPB.
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sQ AMLTCPQGML GLRM 1&1) ?LB PCQSJRQ GL RFC NPMBSARGML MD ? QCAMLB "51 KMJCASJC @W QS@QRP?RC-JCTCJ NFMQNFMPWJ?RGML ?LB RFC AMKNMSLB NWPSTGA ?AGB (MP GRQ Q?JR DMPK, NWPST?RC). .?LW CLXWKCQ GL CLXWK?RGA N?RFU?WQ ?PC L?KCB DMP RFC PCTCPQC PC?ARGMLQ, QGLAC RFC CLXWKC A?L A?R?JWXC @MRF DMPU?PB ?LB PCTCPQC PC?ARGMLQ (RFCQC K?W F?TC @CCL BCQAPG@CB GLGRG?JJW @W RFC PCTCPQC PC?ARGML RF?R R?ICQ NJ?AC GL TGRPM, SLBCP LML-NFWQGMJMEGA?J AMLBGRGMLQ).
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*L MPBCP DMP NWPST?RC, RFC NPMBSAR MD EJWAMJWQGQ, RM CLRCP RFC LCVR N?RFU?W, GR KSQR SLBCPEM QCTCP?J AF?LECQ. 5FC AMLTCPQGML GQ ? RFPCC-QRCN NPMACQQ (Figure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RFC QSAAGLWJ EPMSN RM DMPK QSAAGLWJ $M". 5FC CLXWKC RF?R A?R?JWXCQ QRCN DMSP GQ PCESJ?RCB @W DCCB@?AI GLFG@GRGML MD "51, QSAAGLWJ $M", ?LB /"%). 4RCN 5. *L QRCN DGTC, ? NFMQNF?RC EPMSN GQ QS@QRGRSRCB DMP AMCLXWKC ", ?LB ? FGEF-CLCPEW @MLB GQ DMPKCB. 5FGQ CLCPEW GQ SQCB GL QS@QRP?RC-JCTCJ NFMQNFMPWJ?RGML (BSPGLE RFC AMLTCPQGML MD RFC QSAAGLWJ EPMSN RM QSAAGL?RC) RM DMPK CGRFCP ES?LGLC RPGNFMQNF?RC ((51) MP "51. 5FCPC ?PC RUM DMPKQ MD RFC CLXWKC, A?JJCB GQMCLXWKCQ, DMP RFGQ QRCN, BCNCLBGLE SNML RFC RWNC MD ?LGK?J RGQQSC GL UFGAF RFCW ?PC DMSLB. 0LC DMPK GQ DMSLB GL RGQQSCQ RF?R SQC J?PEC ?KMSLRQ MD "51, QSAF ?Q FC?PR ?LB QICJCR?J KSQAJC. 5FGQ DMPK NPMBSACQ "51. 5FC QCAMLB DMPK MD RFC CLXWKC GQ DMSLB GL RGQQSCQ RF?R F?TC ? FGEF LSK@CP MD ?L?@MJGA N?RFU?WQ, QSAF ?Q JGTCP. 5FGQ DMPK NPMBSACQ (51. (51 GQ CLCPECRGA?JJW COSGT?JCLR RM "51; FMUCTCP, GRQ SQC GQ KMPC PCQRPGARCB. *L N?PRGASJ?P, NPMRCGL QWLRFCQGQ NPGK?PGJW SQCQ (51. 4RCN 6. 4RCN QGV GQ ? BCFWBP?RGML NPMACQQ RF?R AMLTCPRQ QSAAGL?RC GLRM DSK?P?RC. 5UM FWBPMECL ?RMKQ ?PC RP?LQDCPPCB RM '"%, NPMBSAGLE '"%)2. 5FC CLCPEW AMLR?GLCB GL RFC CJCARPMLQ MD RFCQC ?RMKQ GQ GLQSDDGAGCLR RM PCBSAC /"%+ @SR ?BCOS?RC RM PCBSAC '"%. 6LJGIC /"%), RFGQ A?PPGCP PCK?GLQ ?RR?AFCB RM RFC CLXWKC ?LB RP?LQDCPQ RFC CJCARPMLQ RM RFC CJCARPML RP?LQNMPR AF?GL BGPCARJW. 5FGQ NPMACQQ GQ K?BC NMQQG@JC @W RFC JMA?JGX?RGML MD RFC CLXWKC A?R?JWXGLE RFGQ QRCN GLQGBC RFC GLLCP KCK@P?LC MD RFC KGRMAFMLBPGML. 4RCN 7. 8?RCP GQ ?BBCB RM DSK?P?RC BSPGLE QRCN QCTCL, ?LB K?J?RC GQ NPMBSACB. 5FC J?QR QRCN GL RFC AGRPGA ?AGB AWAJC PCECLCP?RCQ MV?JM?ACR?RC @W MVGBGXGLE K?J?RC. "LMRFCP KMJCASJC MD /"%) GQ NPMBSACB GL RFC NPMACQQ.
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5FC CJCARPML RP?LQNMPR AF?GL (Figure 7.10) GQ RFC J?QR AMKNMLCLR MD ?CPM@GA PCQNGP?RGML ?LB GQ RFC MLJW N?PR MD EJSAMQC KCR?@MJGQK RF?R SQCQ ?RKMQNFCPGA MVWECL. 0VWECL AMLRGLSMSQJW BGDDSQCQ GLRM NJ?LRQ; GL ?LGK?JQ, GR CLRCPQ RFC @MBW RFPMSEF RFC PCQNGP?RMPW QWQRCK. &JCARPML RP?LQNMPR GQ ? QCPGCQ MD PCBMV PC?ARGMLQ RF?R PCQCK@JC ? PCJ?W P?AC MP @SAICR @PGE?BC GL RF?R CJCARPMLQ ?PC N?QQCB P?NGBJW DPMK MLC AMKNMLCLR RM RFC LCVR, RM RFC CLBNMGLR MD RFC AF?GL UFCPC RFC CJCARPMLQ PCBSAC KMJCASJ?P MVWECL, NPMBSAGLE U?RCP. 5FCPC ?PC DMSP AMKNJCVCQ AMKNMQCB MD NPMRCGLQ, J?@CJCB * RFPMSEF *7 GL Figure 7.10, ?LB RFC ?EEPCE?RGML MD RFCQC DMSP AMKNJCVCQ, RMECRFCP UGRF ?QQMAG?RCB KM@GJC, ?AACQQMPW CJCARPML A?PPGCPQ, GQ A?JJCB RFC CJCARPML RP?LQNMPR AF?GL. 5FC CJCARPML RP?LQNMPR AF?GL GQ NPCQCLR GL KSJRGNJC AMNGCQ GL RFC GLLCP KGRMAFMLBPG?J KCK@P?LC MD CSI?PWMRCQ ?LB RFC NJ?QK? KCK@P?LC MD NPMI?PWMRCQ.
!@>LI< 7.10 4EB BIB@QOLK QO>KPMLOQ @E>FK FP > PBOFBP LC BIB@QOLK QO>KPMLOQBOP BJ?BAABA FK QEB FKKBO JFQL@ELKAOF>I JBJ?O>KB QE>Q PERQQIBP BIB@QOLKP COLJ .!$( >KA &!$(2 QL JLIB@RI>O LUVDBK. )K QEB MOL@BPP, MOLQLKP >OB MRJMBA COLJ QEB JFQL@ELKAOF>I J>QOFU QL QEB FKQBOJBJ?O>KB PM>@B, >KA LUVDBK FP OBAR@BA QL CLOJ T>QBO.
C4251+= I 5M QR?PR, RUM CJCARPMLQ ?PC A?PPGCB RM RFC DGPQR AMKNJCV ?@M?PB /"%). 5FGQ AMKNJCV, J?@CJCB *, GQ AMKNMQCB MD DJ?TGL KMLMLSAJCMRGBC ('./) ?LB ?L GPML-QSJDSP ('C-4)-AMLR?GLGLE NPMRCGL. './, UFGAF GQ BCPGTCB DPMK TGR?KGL #2, ?JQM A?JJCB PG@MDJ?TGL, GQ MLC MD QCTCP?J NPMQRFCRGA EPMSNQ MP AM-D?ARMPQ GL RFC CJCARPML RP?LQNMPR AF?GL. " prosthetic group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ubiquinone (2). 5FC 2 KMJCASJC GQ JGNGB QMJS@JC ?LB DPCCJW KMTCQ RFPMSEF RFC FWBPMNFM@GA AMPC MD RFC KCK@P?LC. 0LAC GR GQ PCBSACB, (2)2), S@GOSGLMLC BCJGTCPQ GRQ CJCARPMLQ RM RFC LCVR AMKNJCV GL RFC CJCARPML RP?LQNMPR AF?GL. 2 PCACGTCQ RFC CJCARPMLQ BCPGTCB DPMK /"%) DPMK AMKNJCV * ?LB RFC CJCARPMLQ BCPGTCB DPMK '"%)2 DPMK AMKNJCV **, GLAJSBGLE QSAAGL?RC BCFWBPMECL?QC. 5FGQ CLXWKC ?LB '"%)2 DMPK ? QK?JJ AMKNJCV RF?R BCJGTCPQ CJCARPMLQ BGPCARJW RM RFC CJCARPML RP?LQNMPR AF?GL, @WN?QQGLE RFC DGPQR AMKNJCV. 4GLAC RFCQC CJCARPMLQ @WN?QQ ?LB RFSQ BM LMR CLCPEGXC RFC NPMRML NSKN GL RFC DGPQR AMKNJCV, DCUCP "51 KMJCASJCQ ?PC K?BC DPMK RFC '"%)2 CJCARPMLQ. 5FC LSK@CP MD "51 KMJCASJCQ SJRGK?RCJW M@R?GLCB GQ BGPCARJW NPMNMPRGML?J RM RFC LSK@CP MD NPMRMLQ NSKNCB ?APMQQ RFC GLLCP KGRMAFMLBPG?J KCK@P?LC. C4251+= III 5FC RFGPB AMKNJCV GQ AMKNMQCB MD AWRMAFPMKC @, ?LMRFCP 'C-4 NPMRCGL, 3GCQIC ACLRCP (2'C-24 ACLRCP), ?LB AWRMAFPMKC A NPMRCGLQ; RFGQ AMKNJCV GQ ?JQM A?JJCB AWRMAFPMKC MVGBMPCBSAR?QC. $WRMAFPMKC NPMRCGLQ F?TC ? NPMQRFCRGA EPMSN MD FCKC. 5FC FCKC KMJCASJC GQ QGKGJ?P RM RFC FCKC GL FCKMEJM@GL, @SR GR A?PPGCQ
#(!04%2 7 X #%,,5,!2 2%30)2!4)/.
207
CJCARPMLQ, LMR MVWECL. "Q ? PCQSJR, RFC GPML GML ?R GRQ AMPC GQ PCBSACB ?LB MVGBGXCB ?Q GR N?QQCQ RFC CJCARPMLQ, DJSARS?RGLE @CRUCCL BGDDCPCLR MVGB?RGML QR?RCQ: 'C++ (PCBSACB) ?LB 'C+++ (MVGBGXCB). 5FC FCKC KMJCASJCQ GL RFC AWRMAFPMKCQ F?TC QJGEFRJW BGDDCPCLR AF?P?ARCPGQRGAQ BSC RM RFC CDDCARQ MD RFC BGDDCPCLR NPMRCGLQ @GLBGLE RFCK, EGTGLE QJGEFRJW BGDDCPCLR AF?P?ARCPGQRGAQ RM C?AF AMKNJCV. $MKNJCV *** NSKNQ NPMRMLQ RFPMSEF RFC KCK@P?LC ?LB N?QQCQ GRQ CJCARPMLQ RM AWRMAFPMKC A DMP RP?LQNMPR RM RFC DMSPRF AMKNJCV MD NPMRCGLQ ?LB CLXWKCQ (AWRMAFPMKC A GQ RFC ?AACNRMP MD CJCARPMLQ DPMK 2; FMUCTCP, UFCPC?Q 2 A?PPGCQ N?GPQ MD CJCARPMLQ, AWRMAFPMKC A A?L ?AACNR MLJW MLC ?R ? RGKC). C4251+= I$ 5FC DMSPRF AMKNJCV GQ AMKNMQCB MD AWRMAFPMKC NPMRCGLQ A, ?, ?LB ?3. 5FGQ AMKNJCV AMLR?GLQ RUM FCKC EPMSNQ (MLC GL C?AF MD RFC RUM AWRMAFPMKCQ, ?, ?LB ?3) ?LB RFPCC AMNNCP GMLQ (? N?GP MD $S" ?LB MLC $S# GL AWRMAFPMKC ?3). 5FC AWRMAFPMKCQ FMJB ?L MVWECL KMJCASJC TCPW RGEFRJW @CRUCCL RFC GPML ?LB AMNNCP GMLQ SLRGJ RFC MVWECL GQ AMKNJCRCJW PCBSACB. 5FC PCBSACB MVWECL RFCL NGAIQ SN RUM FWBPMECL GMLQ DPMK RFC QSPPMSLBGLE KCBGSK RM K?IC U?RCP ()20). 5FC PCKMT?J MD RFC FWBPMECL GMLQ DPMK RFC QWQRCK AMLRPG@SRCQ RM RFC GML EP?BGCLR SQCB GL RFC NPMACQQ MD AFCKGMQKMQGQ.
C?<D@FJDFJ@J
*L AFCKGMQKMQGQ, RFC DPCC CLCPEW DPMK RFC QCPGCQ MD PCBMV PC?ARGMLQ HSQR BCQAPG@CB GQ SQCB RM NSKN FWBPMECL GMLQ (NPMRMLQ) ?APMQQ RFC KCK@P?LC. 5FC SLCTCL BGQRPG@SRGML MD )+ GMLQ ?APMQQ RFC KCK@P?LC CQR?@JGQFCQ @MRF AMLACLRP?RGML ?LB CJCARPGA?J EP?BGCLRQ (RFSQ, ?L CJCARPMAFCKGA?J EP?BGCLR), MUGLE RM RFC FWBPMECL GMLQs NMQGRGTC AF?PEC ?LB RFCGP ?EEPCE?RGML ML MLC QGBC MD RFC KCK@P?LC. *D RFC KCK@P?LC UCPC MNCL RM BGDDSQGML @W RFC FWBPMECL GMLQ, RFC GMLQ UMSJB RCLB RM BGDDSQC @?AI ?APMQQ GLRM RFC K?RPGV, BPGTCL @W RFCGP CJCARPMAFCKGA?J EP?BGCLR. 3CA?JJ RF?R K?LW GMLQ A?LLMR BGDDSQC RFPMSEF RFC LMLNMJ?P PCEGMLQ MD NFMQNFMJGNGB KCK@P?LCQ UGRFMSR RFC ?GB MD GML AF?LLCJQ. 4GKGJ?PJW, FWBPMECL GMLQ GL RFC K?RPGV QN?AC A?L MLJW N?QQ RFPMSEF RFC GLLCP KGRMAFMLBPG?J KCK@P?LC RFPMSEF ?L GLRCEP?J KCK@P?LC NPMRCGL A?JJCB "51 QWLRF?QC (Figure 7.11). 5FGQ AMKNJCV NPMRCGL ?ARQ ?Q ? RGLW ECLCP?RMP, RSPLCB @W RFC DMPAC MD RFC FWBPMECL GMLQ BGDDSQGLE RFPMSEF GR, BMUL RFCGP CJCARPMAFCKGA?J EP?BGCLR. 5FC RSPLGLE MD N?PRQ MD RFGQ KMJCASJ?P K?AFGLC D?AGJGR?RCQ RFC ?BBGRGML MD ? NFMQNF?RC RM "%1, DMPKGLE "51, SQGLE RFC NMRCLRG?J CLCPEW MD RFC FWBPMECL GML EP?BGCLR.
208
#(!04%2 7 X #%,,5,!2 2%30)2!4)/.
!@>LI< 7.11 !40 PVKQE>PB FP > @LJMIBU, JLIB@RI>O J>@EFKB QE>Q RPBP > MOLQLK ((+) DO>AFBKQ QL CLOJ !40 COLJ !$0 >KA FKLOD>KF@ MELPME>QB (0F). (#OBAFQ: JLAFCF@>QFLK LC TLOH ?V +I>RP (LCCJBFBO)
$FKFQOLMEBKLI ($.0) FP >K RK@LRMIBO QE>Q J>HBP QEB FKKBO JFQL@ELKAOF>I JBJ?O>KB IB>HV QL MOLQLKP. )Q T>P RPBA RKQFI 1938 >P > TBFDEQ-ILPP AORD. 7E>Q BCCB@Q TLRIA VLR BUMB@Q $.0 QL E>SB LK QEB @E>KDB FK M( >@OLPP QEB FKKBO JFQL@ELKAOF>I JBJ?O>KB 7EV AL VLR QEFKH QEFP JFDEQ ?B >K BCCB@QFSB TBFDEQ-ILPP AORD $FCKGMQKMQGQ (Figure 7.12) GQ SQCB RM ECLCP?RC 90 NCPACLR MD RFC "51 K?BC BSPGLE ?CPM@GA EJSAMQC A?R?@MJGQK; GR GQ ?JQM RFC KCRFMB SQCB GL RFC JGEFR PC?ARGMLQ MD NFMRMQWLRFCQGQ RM F?PLCQQ RFC CLCPEW MD QSLJGEFR GL RFC NPMACQQ MD NFMRMNFMQNFMPWJ?RGML. 3CA?JJ RF?R RFC NPMBSARGML MD "51 SQGLE RFC NPMACQQ MD AFCKGMQKMQGQ GL KGRMAFMLBPG? GQ A?JJCB MVGB?RGTC NFMQNFMPWJ?RGML. 5FC MTCP?JJ PCQSJR MD RFCQC PC?ARGMLQ GQ RFC NPMBSARGML MD "51 DPMK RFC CLCPEW MD RFC CJCARPMLQ PCKMTCB DPMK FWBPMECL ?RMKQ. 5FCQC ?RMKQ UCPC MPGEGL?JJW N?PR MD ? EJSAMQC KMJCASJC. "R RFC CLB MD RFC N?RFU?W, RFC CJCARPMLQ ?PC SQCB RM PCBSAC ?L MVWECL KMJCASJC RM MVWECL GMLQ. 5FC CVRP? CJCARPMLQ ML RFC MVWECL ?RRP?AR FWBPMECL GMLQ (NPMRMLQ) DPMK RFC QSPPMSLBGLE KCBGSK, ?LB U?RCP GQ DMPKCB.
#(!04%2 7 X #%,,5,!2 2%30)2!4)/.
209
!@>LI< 7.12 )K LUFA>QFSB MELPMELOVI>QFLK, QEB M( DO>AFBKQ CLOJBA ?V QEB BIB@QOLK QO>KPMLOQ @E>FK FP RPBA ?V !40 PVKQE>PB QL CLOJ !40.
#V>KFAB FKEF?FQP @VQL@EOLJB @ LUFA>PB, > @LJMLKBKQ LC QEB BIB@QOLK QO>KPMLOQ @E>FK. )C @V>KFAB MLFPLKFKD L@@ROP, TLRIA VLR BUMB@Q QEB M( LC QEB FKQBOJBJ?O>KB PM>@B QL FK@OB>PB LO AB@OB>PB 7E>Q BCCB@Q TLRIA @V>KFAB E>SB LK !40 PVKQEBPFP
A/+ 4@<C;
5FC LSK@CP MD "51 KMJCASJCQ ECLCP?RCB DPMK RFC A?R?@MJGQK MD EJSAMQC T?PGCQ. 'MP CV?KNJC, RFC LSK@CP MD FWBPMECL GMLQ RF?R RFC CJCARPML RP?LQNMPR AF?GL AMKNJCVCQ A?L NSKN RFPMSEF RFC KCK@P?LC T?PGCQ @CRUCCL QNCAGCQ. "LMRFCP QMSPAC MD T?PG?LAC QRCKQ DPMK RFC QFSRRJC MD CJCARPMLQ ?APMQQ RFC KCK@P?LCQ MD RFC KGRMAFMLBPG?. (5FC /"%) ECLCP?RCB DPMK EJWAMJWQGQ A?LLMR C?QGJW CLRCP KGRMAFMLBPG?.) 5FSQ, CJCARPMLQ ?PC NGAICB SN ML RFC GLQGBC MD KGRMAFMLBPG? @W CGRFCP /"%+ MP '"%+. "Q WMS F?TC JC?PLCB C?PJGCP, RFCQC '"%+ KMJCASJCQ A?L RP?LQNMPR DCUCP GMLQ; AMLQCOSCLRJW, DCUCP "51 KMJCASJCQ ?PC ECLCP?RCB UFCL '"%+ ?ARQ ?Q ? A?PPGCP. /"%+ GQ SQCB ?Q RFC CJCARPML RP?LQNMPRCP GL RFC JGTCP ?LB '"%+ ?ARQ GL RFC @P?GL. "LMRFCP D?ARMP RF?R ?DDCARQ RFC WGCJB MD "51 KMJCASJCQ ECLCP?RCB DPMK EJSAMQC GQ RFC D?AR RF?R GLRCPKCBG?RC AMKNMSLBQ GL RFCQC N?RFU?WQ ?PC SQCB DMP MRFCP NSPNMQCQ. (JSAMQC A?R?@MJGQK AMLLCARQ UGRF RFC N?RFU?WQ RF?R @SGJB MP @PC?I BMUL ?JJ MRFCP @GMAFCKGA?J AMKNMSLBQ GL ACJJQ, ?LB RFC PCQSJR GQ QMKCUF?R KCQQGCP RF?L RFC GBC?J QGRS?RGMLQ BCQAPG@CB RFSQ D?P. 'MP CV?KNJC, QSE?PQ MRFCP RF?L EJSAMQC ?PC DCB GLRM RFC EJWAMJWRGA N?RFU?W DMP CLCPEW CVRP?ARGML. .MPCMTCP, RFC DGTC-A?P@ML QSE?PQ RF?R DMPK LSAJCGA ?AGBQ ?PC K?BC DPMK GLRCPKCBG?RCQ GL EJWAMJWQGQ. $CPR?GL LMLCQQCLRG?J ?KGLM ?AGBQ A?L @C K?BC DPMK GLRCPKCBG?RCQ MD @MRF EJWAMJWQGQ ?LB RFC AGRPGA ?AGB AWAJC. -GNGBQ, QSAF ?Q AFMJCQRCPMJ ?LB RPGEJWACPGBCQ, ?PC ?JQM K?BC DPMK GLRCPKCBG?RCQ GL RFCQC N?RFU?WQ, ?LB @MRF ?KGLM ?AGBQ ?LB RPGEJWACPGBCQ ?PC @PMICL BMUL DMP CLCPEW RFPMSEF RFCQC N?RFU?WQ. 0TCP?JJ, GL JGTGLE QWQRCKQ, RFCQC N?RFU?WQ MD EJSAMQC A?R?@MJGQK CVRP?AR ?@MSR 34 NCPACLR MD RFC CLCPEW AMLR?GLCB GL EJSAMQC.
7.5 R (<K89FC@JD N@K?FLK *OP><E

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %GQASQQ RFC DSLB?KCLR?J BGDDCPCLAC @CRUCCL ?L?CPM@GA ACJJSJ?P PCQNGP?RGML ?LB DCPKCLR?RGML z %CQAPG@C RFC RWNC MD DCPKCLR?RGML RF?R PC?BGJW MAASPQ GL ?LGK?J ACJJQ ?LB RFC AMLBGRGMLQ RF?R GLGRG?RC RF?R DCPKCLR?RGML *L ?CPM@GA PCQNGP?RGML, RFC DGL?J CJCARPML ?AACNRMP GQ ?L MVWECL KMJCASJC, 02. *D ?CPM@GA PCQNGP?RGML MAASPQ, RFCL "51 UGJJ @C NPMBSACB SQGLE RFC CLCPEW MD FGEF-CLCPEW CJCARPMLQ A?PPGCB @W /"%) MP
210
#(!04%2 7 X #%,,5,!2 2%30)2!4)/.
'"%)2 RM RFC CJCARPML RP?LQNMPR AF?GL. *D ?CPM@GA PCQNGP?RGML BMCQ LMR MAASP, /"%) KSQR @C PCMVGBGXCB RM /"%+ DMP PCSQC ?Q ?L CJCARPML A?PPGCP DMP RFC EJWAMJWRGA N?RFU?W RM AMLRGLSC. )MU GQ RFGQ BMLC 4MKC JGTGLE QWQRCKQ SQC ?L MPE?LGA KMJCASJC ?Q RFC DGL?J CJCARPML ?AACNRMP. 1PMACQQCQ RF?R SQC ?L MPE?LGA KMJCASJC RM PCECLCP?RC /"%+ DPMK /"%) ?PC AMJJCARGTCJW PCDCPPCB RM ?Q fermentation. *L AMLRP?QR, QMKC JGTGLE QWQRCKQ SQC ?L GLMPE?LGA KMJCASJC ?Q ? DGL?J CJCARPML ?AACNRMP. #MRF KCRFMBQ ?PC A?JJCB anaerobic cellular respiration GL UFGAF MPE?LGQKQ AMLTCPR CLCPEW DMP RFCGP SQC GL RFC ?@QCLAC MD MVWECL.
AE8<IF9@: C<CCLC8I -<JG@I8K@FE

$CPR?GL NPMI?PWMRCQ, GLAJSBGLE QMKC QNCAGCQ MD @?ARCPG? ?LB "PAF?C?, SQC ?L?CPM@GA PCQNGP?RGML. 'MP CV?KNJC, RFC EPMSN MD "PAF?C? A?JJCB KCRF?LMECLQ PCBSACQ A?P@ML BGMVGBC RM KCRF?LC RM MVGBGXC /"%). 5FCQC KGAPMMPE?LGQKQ ?PC DMSLB GL QMGJ ?LB GL RFC BGECQRGTC RP?ARQ MD PSKGL?LRQ, QSAF ?Q AMUQ ?LB QFCCN. 4GKGJ?PJW, QSJD?RC-PCBSAGLE @?ARCPG? ?LB "PAF?C?, KMQR MD UFGAF ?PC ?L?CPM@GA ( Figure 7.13), PCBSAC QSJD?RC RM FWBPMECL QSJDGBC RM PCECLCP?RC /"%+ DPMK /"%).
!@>LI< 7.13 4EB DOBBK @LILO PBBK FK QEBPB @L>PQ>I T>QBOP FP COLJ >K BORMQFLK LC EVAOLDBK PRICFABMOLAR@FKD ?>@QBOF>. 4EBPB >K>BOL?F@, PRIC>QB-OBAR@FKD ?>@QBOF> OBIB>PB EVAOLDBK PRICFAB D>P >P QEBV AB@LJMLPB >ID>B FK QEB T>QBO. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .!3!/*BCC 3@EJ>IQW, -/$)3 ,>KA 2>MFA 2BPMLKPB 4B>J >Q .!3! '3&#, 6FPF?IB %>OQE #>Q>ILD LC .!3! FJ>DBP)
7GQGR RFGQ site (http://openstaxcollege.org/l/fermentation) RM QCC ?L?CPM@GA ACJJSJ?P PCQNGP?RGML GL ?ARGML. L')9/) A)/* F+72+39'9/43 5FC DCPKCLR?RGML KCRFMB SQCB @W ?LGK?JQ ?LB ACPR?GL @?ARCPG?, JGIC RFMQC GL WMESPR, GQ J?ARGA ?AGB DCPKCLR?RGML ( Figure 7.14). 5FGQ RWNC MD DCPKCLR?RGML GQ SQCB PMSRGLCJW GL K?KK?JG?L PCB @JMMB ACJJQ ?LB GL QICJCR?J KSQAJC RF?R F?Q ?L GLQSDDGAGCLR MVWECL QSNNJW RM ?JJMU ?CPM@GA PCQNGP?RGML RM AMLRGLSC (RF?R GQ, GL KSQAJCQ SQCB RM RFC NMGLR MD D?RGESC). *L KSQAJCQ, J?ARGA ?AGB ?AASKSJ?RGML KSQR @C PCKMTCB @W RFC @JMMB AGPASJ?RGML ?LB RFC J?AR?RC @PMSEFR RM RFC JGTCP DMP DSPRFCP KCR?@MJGQK. 5FC AFCKGA?J PC?ARGMLQ MD J?ARGA ?AGB DCPKCLR?RGML ?PC RFC DMJJMUGLE:
#(!04%2 7 X #%,,5,!2 2%30)2!4)/.
211
Pyruvic acid + NADH lactic acid + NAD+

5FC CLXWKC SQCB GL RFGQ PC?ARGML GQ J?AR?RC BCFWBPMECL?QC (-%)). 5FC PC?ARGML A?L NPMACCB GL CGRFCP BGPCARGML, @SR RFC PC?ARGML DPMK JCDR RM PGEFR GQ GLFG@GRCB @W ?AGBGA AMLBGRGMLQ. 4SAF J?ARGA ?AGB ?AASKSJ?RGML U?Q MLAC @CJGCTCB RM A?SQC KSQAJC QRGDDLCQQ, D?RGESC, ?LB QMPCLCQQ, ?JRFMSEF KMPC PCACLR PCQC?PAF BGQNSRCQ RFGQ FWNMRFCQGQ. 0LAC RFC J?ARGA ?AGB F?Q @CCL PCKMTCB DPMK RFC KSQAJC ?LB AGPASJ?RCB RM RFC JGTCP, GR A?L @C PCAMLTCPRCB GLRM NWPSTGA ?AGB ?LB DSPRFCP A?R?@MJGXCB DMP CLCPEW.
!@>LI< 7.14 ,>@QF@ >@FA CBOJBKQ>QFLK FP @LJJLK FK JRP@IB @BIIP QE>Q E>SB ORK LRQ LC LUVDBK.
4OBJBQLI, > JBQ>?LIF@ MLFPLK CLRKA FK QEB TEFQB PK>HB OLLQ MI>KQ, MOBSBKQP QEB JBQ>?LIFPJ LC I>@Q>QB. 7EBK @LTP B>Q QEFP MI>KQ, FQ FP @LK@BKQO>QBA FK QEB JFIH QEBV MOLAR@B. (RJ>KP TEL @LKPRJB QEB JFIH ?B@LJB FII. 3VJMQLJP LC QEFP AFPB>PB, TEF@E FK@IRAB SLJFQFKD, >?ALJFK>I M>FK, >KA QOBJLOP, ?B@LJB TLOPB >CQBO BUBO@FPB. 7EV AL VLR QEFKH QEFP FP QEB @>PB A1)4.41 F+72+39'9/43 "LMRFCP D?KGJG?P DCPKCLR?RGML NPMACQQ GQ ?JAMFMJ DCPKCLR?RGML ( Figure 7.15) RF?R NPMBSACQ CRF?LMJ, ?L ?JAMFMJ. 5FC DGPQR AFCKGA?J PC?ARGML MD ?JAMFMJ DCPKCLR?RGML GQ RFC DMJJMUGLE ($02 BMCQ LMR N?PRGAGN?RC GL RFC QCAMLB PC?ARGML):
Pyruvic acid CO 2 + acetaldehyde + NADH ethanol + NAD+
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
212
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214
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216
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GL ? J?RCP QRCN. 5FC NPCQCLAC MD RFC LCE?RGTCJW AF?PECB NFMQNF?RC GL RFC KMJCASJC ?JQM NPCTCLRQ RFC QSE?P DPMK JC?TGLE RFC ACJJ. 8FCL FCVMIGL?QC GQ GLFG@GRCB, EJSAMQC BGDDSQCQ MSR MD RFC ACJJ ?LB BMCQ LMR @CAMKC ? QS@QRP?RC DMP RFC PCQNGP?RGML N?RFU?WQ GL RF?R RGQQSC. 5FC NPMBSAR MD RFC FCVMIGL?QC PC?ARGML GQ EJSAMQC-6-NFMQNF?RC, UFGAF ?AASKSJ?RCQ UFCL ? J?RCP CLXWKC, NFMQNFMDPSARMIGL?QC, GQ GLFG@GRCB.
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1FMQNFMDPSARMIGL?QC GQ RFC K?GL CLXWKC AMLRPMJJCB GL EJWAMJWQGQ. )GEF JCTCJQ MD "51, AGRP?RC, MP ? JMUCP, KMPC ?AGBGA N) BCAPC?QC RFC CLXWKCsQ ?ARGTGRW. "L GLAPC?QC GL AGRP?RC AMLACLRP?RGML A?L MAASP @CA?SQC MD ? @JMAI?EC GL RFC AGRPGA ?AGB AWAJC. 'CPKCLR?RGML, UGRF GRQ NPMBSARGML MD MPE?LGA ?AGBQ JGIC J?ARGA ?AGB, DPCOSCLRJW ?AAMSLRQ DMP RFC GLAPC?QCB ?AGBGRW GL ? ACJJ; FMUCTCP, RFC NPMBSARQ MD DCPKCLR?RGML BM LMR RWNGA?JJW ?AASKSJ?RC GL ACJJQ. 5FC J?QR QRCN GL EJWAMJWQGQ GQ A?R?JWXCB @W NWPST?RC IGL?QC. 5FC NWPST?RC NPMBSACB A?L NPMACCB RM @C A?R?@MJGXCB MP AMLTCPRCB GLRM RFC ?KGLM ?AGB ?J?LGLC. *D LM KMPC CLCPEW GQ LCCBCB ?LB ?J?LGLC GQ GL ?BCOS?RC QSNNJW, RFC CLXWKC GQ GLFG@GRCB. 5FC CLXWKCs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igure 7.9). 5FCQC CLXWKCQ ?PC GQMAGRP?RC BCFWBPMECL?QC ?LB C-ICRMEJSR?P?RC BCFWBPMECL?QC. 8FCL ?BCOS?RC "51 ?LB /"%) JCTCJQ ?PC ?T?GJ?@JC, RFC P?RCQ MD RFCQC PC?ARGMLQ BCAPC?QC. 8FCL KMPC "51 GQ LCCBCB, ?Q PCDJCARCB GL PGQGLE "%1 JCTCJQ, RFC P?RC GLAPC?QCQ. C,CRMEJSR?P?RC BCFWBPMECL?QC UGJJ ?JQM @C ?DDCARCB @W RFC JCTCJQ MD QSAAGLWJ $M"q? QS@QCOSCLR GLRCPKCBG?RC GL RFC AWAJCqA?SQGLE ? BCAPC?QC GL ?ARGTGRW. " BCAPC?QC GL RFC P?RC MD MNCP?RGML MD RFC N?RFU?W ?R RFGQ NMGLR GQ LMR LCACQQ?PGJW LCE?RGTC, ?Q RFC GLAPC?QCB JCTCJQ MD RFC C-ICRMEJSR?P?RC LMR SQCB @W RFC AGRPGA ?AGB AWAJC A?L @C SQCB @W RFC ACJJ DMP ?KGLM ?AGB (EJSR?K?RC) QWLRFCQGQ. E1+)9743 "7'385479 C.'/3 4NCAGDGA CLXWKCQ MD RFC CJCARPML RP?LQNMPR AF?GL ?PC SL?DDCARCB @W DCCB@?AI GLFG@GRGML, @SR RFC P?RC MD CJCARPML RP?LQNMPR RFPMSEF RFC N?RFU?W GQ ?DDCARCB @W RFC JCTCJQ MD "%1 ?LB "51. (PC?RCP "51 AMLQSKNRGML @W ? ACJJ GQ GLBGA?RCB @W ? @SGJBSN MD "%1. "Q "51 SQ?EC BCAPC?QCQ, RFC AMLACLRP?RGML MD
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217
"%1 BCAPC?QCQ, ?LB LMU, "51 @CEGLQ RM @SGJB SN GL RFC ACJJ. 5FGQ AF?LEC GQ RFC PCJ?RGTC AMLACLRP?RGML MD "%1 RM "51 RPGEECPQ RFC ACJJ RM QJMU BMUL RFC CJCARPML RP?LQNMPR AF?GL.
7GQGR RFGQ site (http://openstaxcollege.org/l/electron_transp) RM QCC ?L ?LGK?RGML MD RFC CJCARPML RP?LQNMPR AF?GL ?LB "51 QWLRFCQGQ. 'MP ? QSKK?PW MD DCCB@?AI AMLRPMJQ GL ACJJSJ?P PCQNGP?RGML, QCC Table 7.1.
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218
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A/+ JPEK?8J< (?JQM, '1'0 "51 QWLRF?QC) KCK@P?LC-CK@CBBCB NPMRCGL AMKNJCV RF?R ?BBQ ? NFMQNF?RC RM "%1 UGRF CLCPEW DPMK NPMRMLQ BGDDSQGLE RFPMSEF GR 8:<KPC CFA AMK@GL?RGML MD ?L ?ACRWJ EPMSN BCPGTCB DPMK NWPSTGA ?AGB ?LB AMCLXWKC ", UFGAF GQ K?BC DPMK N?LRMRFCLGA ?AGB (? #-EPMSN TGR?KGL) 8<IF9@: I<JG@I8K@FE NPMACQQ GL UFGAF MPE?LGQKQ AMLTCPR CLCPEW GL RFC NPCQCLAC MD MVWECL 8E8<IF9@: :<CCLC8I I<JG@I8K@FE NPMACQQ GL UFGAF MPE?LGQKQ AMLTCPR CLCPEW DMP RFCGP SQC GL RFC ?@QCLAC MD MVWECL 8E8<IF9@: NPMACQQ RF?R BMCQ LMR SQC MVWECL :?<D@FJDFJ@J NPMACQQ GL UFGAF RFCPC GQ ? NPMBSARGML MD ?BCLMQGLC RPGNFMQNF?RC ("51) GL ACJJSJ?P KCR?@MJGQK @W RFC GLTMJTCKCLR MD ? NPMRML EP?BGCLR ?APMQQ ? KCK@P?LC :@KI@: 8:@; :P:C< (?JQM, ,PC@Q AWAJC) QCPGCQ MD CLXWKC-A?R?JWXCB AFCKGA?J PC?ARGMLQ MD ACLRP?J GKNMPR?LAC GL ?JJ JGTGLE ACJJQ ;<G?FJG?FIPC8K@FE PCKMT?J MD ? NFMQNF?RC EPMSN DPMK ? KMJCASJC =<ID<EK8K@FE NPMACQQ MD PCECLCP?RGLE /"%+ UGRF CGRFCP ?L GLMPE?LGA MP MPE?LGA AMKNMSLB QCPTGLE ?Q RFC DGL?J CJCARPML ?AAACNRMP, MAASPQ GL RFC ?@QCLAC; MAASPQ GL RFC ?@QCLAC MD MVWECL "'0/ GIFK<@E GLRCEP?J KCK@P?LC NPMRCGL RF?R RP?LQNMPRQ EJSAMQC >CP:FCPJ@J NPMACQQ MD @PC?IGLE EJSAMQC GLRM RUM RFPCC-A?P@ML KMJCASJCQ UGRF RFC NPMBSARGML MD "51 ?LB /"%) @JFD<I8J< CLXWKC RF?R AMLTCPRQ ? KMJCASJC GLRM GRQ GQMKCP &I<9J :P:C< (?JQM, AGRPGA ?AGB AWAJC) ?JRCPL?RC L?KC DMP RFC AGRPGA ?AGB AWAJC, L?KCB ?DRCP )?LQ ,PC@Q UFM DGPQR GBCLRGDGCB RFC QRCNQ GL RFC N?RFU?W GL RFC 1930Q GL NGECML DJGEFR KSQAJCQ; QCC AGRPGA ?AGB AWAJC FO@;8K@M< G?FJG?FIPC8K@FE NPMBSARGML MD "51 SQGLE RFC NPMACQQ MD AFCKGMQKMQGQ ?LB MVWECL G?FJG?FIPC8K@FE ?BBGRGML MD ? FGEF-CLCPEW NFMQNF?RC RM ? AMKNMSLB, SQS?JJW ? KCR?@MJGA GLRCPKCBG?RC, ? NPMRCGL, MP "%1 GIFJK?<K@: >IFLG (?JQM, NPMQRFCRGA AMD?ARMP) KMJCASJC @MSLB RM ? NPMRCGL RF?R D?AGJGR?RCQ RFC DSLARGML MD RFC NPMRCGL GPILM8K< RFPCC-A?P@ML QSE?P RF?R A?L @C BCA?P@MVWJ?RCB ?LB MVGBGXCB RM K?IC ?ACRWJ $M", UFGAF CLRCPQ RFC AGRPGA ?AGB AWAJC SLBCP ?CPM@GA AMLBGRGMLQ; RFC CLB NPMBSAR MD EJWAMJWQGQ I<;FO I<8:K@FE AFCKGA?J PC?ARGML RF?R AMLQGQRQ MD RFC AMSNJGLE MD ?L MVGB?RGML PC?ARGML ?LB ? PCBSARGML PC?ARGML JL9JKI8K<-C<M<C G?FJG?FIPC8K@FE NPMBSARGML MD "51 DPMK "%1 SQGLE RFC CVACQQ CLCPEW DPMK ? AFCKGA?J PC?ARGML ?LB ? NFMQNF?RC EPMSN DPMK ? PC?AR?LR /CA :P:C< (?JQM, AGRPGA ?AGB AWAJC) ?JRCPL?RC L?KC DMP RFC AGRPGA ?AGB AWAJC, L?KCB ?DRCP RFC EPMSN L?KC DMP AGRPGA ?AGB, RPGA?P@MVWJGA ?AGB (5$"); QCC AGRPGA ?AGB AWAJC L9@HL@EFE< QMJS@JC CJCARPML RP?LQNMPRCP GL RFC CJCARPML RP?LQNMPR AF?GL RF?R AMLLCARQ RFC DGPQR MP QCAMLB AMKNJCV RM RFC RFGPB
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219
7.1 EP @E '@M@E> .PJK<DJ

"51 DSLARGMLQ ?Q RFC CLCPEW ASPPCLAW DMP ACJJQ. *R ?JJMUQ RFC ACJJ RM QRMPC CLCPEW @PGCDJW ?LB RP?LQNMPR GR UGRFGL RFC ACJJ RM QSNNMPR CLBCPEMLGA AFCKGA?J PC?ARGMLQ. 5FC QRPSARSPC MD "51 GQ RF?R MD ?L 3/" LSAJCMRGBC UGRF RFPCC NFMQNF?RCQ ?RR?AFCB. "Q "51 GQ SQCB DMP CLCPEW, ? NFMQNF?RC EPMSN MP RUM ?PC BCR?AFCB, ?LB CGRFCP "%1 MP ".1 GQ NPMBSACB. &LCPEW BCPGTCB DPMK EJSAMQC A?R?@MJGQK GQ SQCB RM AMLTCPR "%1 GLRM "51. 8FCL "51 GQ SQCB GL ? PC?ARGML, RFC RFGPB NFMQNF?RC GQ RCKNMP?PGJW ?RR?AFCB RM ? QS@QRP?RC GL ? NPMACQQ A?JJCB NFMQNFMPWJ?RGML. 5FC RUM NPMACQQCQ MD "51 PCECLCP?RGML RF?R ?PC SQCB GL AMLHSLARGML UGRF EJSAMQC A?R?@MJGQK ?PC QS@QRP?RC-JCTCJ NFMQNFMPWJ?RGML ?LB MVGB?RGTC NFMQNFMPWJ?RGML RFPMSEF RFC NPMACQQ MD AFCKGMQKMQGQ.
7.2 "CP:FCPJ@J
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
7.3 *O@;8K@FE F= +PILM8K< 8E; K?< C@KI@: A:@; CP:C<

*L RFC NPCQCLAC MD MVWECL, NWPST?RC GQ RP?LQDMPKCB GLRM ?L ?ACRWJ EPMSN ?RR?AFCB RM ? A?PPGCP KMJCASJC MD AMCLXWKC ". 5FC PCQSJRGLE ?ACRWJ $M" A?L CLRCP QCTCP?J N?RFU?WQ, @SR KMQR MDRCL, RFC ?ACRWJ EPMSN GQ BCJGTCPCB RM RFC AGRPGA ?AGB AWAJC DMP DSPRFCP A?R?@MJGQK. %SPGLE RFC AMLTCPQGML MD NWPST?RC GLRM RFC ?ACRWJ EPMSN, ? KMJCASJC MD A?P@ML BGMVGBC ?LB RUM FGEF-CLCPEW CJCARPMLQ ?PC PCKMTCB. 5FC A?P@ML BGMVGBC ?AAMSLRQ DMP RUM (AMLTCPQGML MD RUM NWPST?RC KMJCASJCQ) MD RFC QGV A?P@MLQ MD RFC MPGEGL?J EJSAMQC KMJCASJC. 5FC CJCARPMLQ ?PC NGAICB SN @W /"%+, ?LB RFC /"%) A?PPGCQ RFC CJCARPMLQ RM ? J?RCP N?RFU?W DMP "51 NPMBSARGML. "R RFGQ NMGLR, RFC EJSAMQC KMJCASJC RF?R MPGEGL?JJW CLRCPCB ACJJSJ?P PCQNGP?RGML F?Q @CCL AMKNJCRCJW MVGBGXCB. $FCKGA?J NMRCLRG?J CLCPEW QRMPCB UGRFGL RFC EJSAMQC KMJCASJC F?Q @CCL RP?LQDCPPCB RM CJCARPML A?PPGCPQ MP F?Q @CCL SQCB RM QWLRFCQGXC ? DCU "51Q. 5FC AGRPGA ?AGB AWAJC GQ ? QCPGCQ MD PCBMV ?LB BCA?P@MVWJ?RGML PC?ARGMLQ RF?R PCKMTC FGEF-CLCPEW CJCARPMLQ ?LB A?P@ML BGMVGBC. 5FC CJCARPMLQ RCKNMP?PGJW QRMPCB GL KMJCASJCQ MD /"%) ?LB '"%)2 ?PC SQCB RM ECLCP?RC "51 GL ? QS@QCOSCLR N?RFU?W. 0LC KMJCASJC MD CGRFCP (51 MP "51 GQ NPMBSACB @W QS@QRP?RC-JCTCJ NFMQNFMPWJ?RGML ML C?AF RSPL MD RFC AWAJC. 5FCPC GQ LM AMKN?PGQML MD RFC AWAJGA N?RFU?W UGRF ? JGLC?P MLC.
7.4 *O@;8K@M< +?FJG?FIPC8K@FE

5FC CJCARPML RP?LQNMPR AF?GL GQ RFC NMPRGML MD ?CPM@GA PCQNGP?RGML RF?R SQCQ DPCC MVWECL ?Q RFC DGL?J CJCARPML ?AACNRMP MD RFC CJCARPMLQ PCKMTCB DPMK RFC GLRCPKCBG?RC AMKNMSLBQ GL EJSAMQC A?R?@MJGQK. 5FC CJCARPML RP?LQNMPR AF?GL GQ AMKNMQCB MD DMSP J?PEC, KSJRGNPMRCGL AMKNJCVCQ CK@CBBCB GL RFC GLLCP KGRMAFMLBPG?J KCK@P?LC ?LB RUM QK?JJ BGDDSQG@JC CJCARPML A?PPGCPQ QFSRRJGLE CJCARPMLQ @CRUCCL RFCK. 5FC CJCARPMLQ ?PC N?QQCB RFPMSEF ? QCPGCQ MD PCBMV PC?ARGMLQ, UGRF ? QK?JJ ?KMSLR MD DPCC CLCPEW SQCB ?R RFPCC NMGLRQ RM RP?LQNMPR FWBPMECL GMLQ ?APMQQ ? KCK@P?LC. 5FGQ NPMACQQ AMLRPG@SRCQ RM RFC EP?BGCLR SQCB GL AFCKGMQKMQGQ. 5FC CJCARPMLQ N?QQGLE RFPMSEF RFC CJCARPML RP?LQNMPR AF?GL EP?BS?JJW JMQC CLCPEW, )GEF-CLCPEW CJCARPMLQ BML?RCB RM RFC AF?GL @W CGRFCP /"%) MP '"%)2 AMKNJCRC RFC AF?GL, ?Q JMUCLCPEW CJCARPMLQ PCBSAC MVWECL KMJCASJCQ ?LB DMPK U?RCP. 5FC JCTCJ MD DPCC CLCPEW MD RFC CJCARPMLQ BPMNQ DPMK ?@MSR 60 IA?J/KMJ GL /"%) MP 45 IA?J/KMJ GL '"%)2 RM ?@MSR 0 IA?J/KMJ GL U?RCP. 5FC CLB NPMBSARQ MD RFC CJCARPML RP?LQNMPR AF?GL ?PC U?RCP ?LB "51. " LSK@CP MD GLRCPKCBG?RC AMKNMSLBQ MD RFC AGRPGA ?AGB AWAJC A?L @C BGTCPRCB GLRM RFC ?L?@MJGQK MD MRFCP @GMAFCKGA?J KMJCASJCQ, QSAF ?Q LMLCQQCLRG?J ?KGLM ?AGBQ, QSE?PQ, ?LB JGNGBQ. 5FCQC Q?KC KMJCASJCQ A?L QCPTC ?Q CLCPEW QMSPACQ DMP RFC EJSAMQC N?RFU?WQ.
7.5 (<K89FC@JD N@K?FLK *OP><E

*D /"%) A?LLMR @C MVGBGXCB RFPMSEF ?CPM@GA PCQNGP?RGML, ?LMRFCP CJCARPML ?AACNRMP GQ SQCB. .MQR MPE?LGQKQ UGJJ SQC QMKC DMPK MD DCPKCLR?RGML RM ?AAMKNJGQF RFC PCECLCP?RGML MD /"%+, CLQSPGLE RFC AMLRGLS?RGML MD EJWAMJWQGQ. 5FC PCECLCP?RGML MD /"%+ GL DCPKCLR?RGML GQ LMR ?AAMKN?LGCB @W "51
220
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NPMBSARGML; RFCPCDMPC, RFC NMRCLRG?J MD /"%) RM NPMBSAC "51 SQGLE ?L CJCARPML RP?LQNMPR AF?GL GQ LMR SRGJGXCB.
7.6 CFEE<:K@FEJ F= C8I9F?P;I8K<, +IFK<@E, 8E; '@G@; (<K89FC@: +8K?N8PJ

5FC @PC?IBMUL ?LB QWLRFCQGQ MD A?P@MFWBP?RCQ, NPMRCGLQ, ?LB JGNGBQ AMLLCAR UGRF RFC N?RFU?WQ MD EJSAMQC A?R?@MJGQK. 5FC QGKNJC QSE?PQ ?PC E?J?ARMQC, DPSARMQC, EJWAMECL, ?LB NCLRMQC. 5FCQC ?PC A?R?@MJGXCB BSPGLE EJWAMJWQGQ. 5FC ?KGLM ?AGBQ DPMK NPMRCGLQ AMLLCAR UGRF EJSAMQC A?R?@MJGQK RFPMSEF NWPST?RC, ?ACRWJ $M", ?LB AMKNMLCLRQ MD RFC AGRPGA ?AGB AWAJC. $FMJCQRCPMJ QWLRFCQGQ QR?PRQ UGRF ?ACRWJ EPMSNQ, ?LB RFC AMKNMLCLRQ MD RPGEJWACPGBCQ AMKC DPMK EJWACPMJ-3-NFMQNF?RC DPMK EJWAMJWQGQ ?LB ?ACRWJ EPMSNQ NPMBSACB GL RFC KGRMAFMLBPG? DPMK NWPST?RC.
7.7 -<>LC8K@FE F= C<CCLC8I -<JG@I8K@FE

$CJJSJ?P PCQNGP?RGML GQ AMLRPMJJCB @W ? T?PGCRW MD KC?LQ. 5FC CLRPW MD EJSAMQC GLRM ? ACJJ GQ AMLRPMJJCB @W RFC RP?LQNMPR NPMRCGLQ RF?R ?GB EJSAMQC N?QQ?EC RFPMSEF RFC ACJJ KCK@P?LC. .MQR MD RFC AMLRPMJ MD RFC PCQNGP?RGML NPMACQQCQ GQ ?AAMKNJGQFCB RFPMSEF RFC AMLRPMJ MD QNCAGDGA CLXWKCQ GL RFC N?RFU?WQ. 5FGQ GQ ? RWNC MD LCE?RGTC DCCB@?AI, RSPLGLE RFC CLXWKCQ MDD. 5FC CLXWKCQ PCQNMLB KMQR MDRCL RM RFC JCTCJQ MD RFC ?T?GJ?@JC LSAJCMQGBCQ "51, "%1, ".1, /"%+, ?LB '"%. 0RFCP GLRCPKCBG?RCQ MD RFC N?RFU?W ?JQM ?DDCAR ACPR?GL CLXWKCQ GL RFC QWQRCKQ.
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C DMSP RWNCQ MD QGEL?JGLE DMSLB GL KSJRGACJJSJ?P MPE?LGQKQ z $MKN?PC GLRCPL?J PCACNRMPQ UGRF ACJJ-QSPD?AC PCACNRMPQ z 3CAMELGXC RFC PCJ?RGMLQFGN @CRUCCL ? JGE?LBsQ QRPSARSPC ?LB GRQ KCAF?LGQK MD ?ARGML 5FCPC ?PC RUM IGLBQ MD AMKKSLGA?RGML GL RFC UMPJB MD JGTGLE ACJJQ. $MKKSLGA?RGML @CRUCCL ACJJQ GQ A?JJCB intercellular signaling, ?LB AMKKSLGA?RGML UGRFGL ? ACJJ GQ A?JJCB intracellular signaling. "L C?QW U?W RM PCKCK@CP RFC BGQRGLARGML GQ @W SLBCPQR?LBGLE RFC -?RGL MPGEGL MD RFC NPCDGVCQ: GLRCP- KC?LQ "@CRUCCL" (DMP CV?KNJC, GLRCPQCARGLE JGLCQ ?PC RFMQC RF?R APMQQ C?AF MRFCP) ?LB GLRP?- KC?LQ "GLQGBC" (JGIC GLRP?TCLMSQ). $FCKGA?J QGEL?JQ ?PC PCJC?QCB @W signaling cells GL RFC DMPK MD QK?JJ, SQS?JJW TMJ?RGJC MP QMJS@JC KMJCASJCQ A?JJCB JGE?LBQ. " ligand GQ ? KMJCASJC RF?R @GLBQ ?LMRFCP QNCAGDGA KMJCASJC, GL QMKC A?QCQ, BCJGTCPGLE ? QGEL?J GL RFC NPMACQQ. -GE?LBQ A?L RFSQ @C RFMSEFR MD ?Q QGEL?JGLE KMJCASJCQ. -GE?LBQ GLRCP?AR UGRF NPMRCGLQ GL target cells, UFGAF ?PC ACJJQ RF?R ?PC ?DDCARCB @W AFCKGA?J QGEL?JQ; RFCQC NPMRCGLQ ?PC ?JQM A?JJCB receptors. -GE?LBQ ?LB PCACNRMPQ CVGQR GL QCTCP?J T?PGCRGCQ; FMUCTCP, ? QNCAGDGA JGE?LB UGJJ F?TC ? QNCAGDGA PCACNRMP RF?R RWNGA?JJW @GLBQ MLJW RF?R JGE?LB.
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5FCPC ?PC DMSP A?RCEMPGCQ MD AFCKGA?J QGEL?JGLE DMSLB GL KSJRGACJJSJ?P MPE?LGQKQ: N?P?APGLC QGEL?JGLE, CLBMAPGLC QGEL?JGLE, ?SRMAPGLC QGEL?JGLE, ?LB BGPCAR QGEL?JGLE ?APMQQ E?N HSLARGMLQ (Figure 9.2). 5FC K?GL BGDDCPCLAC @CRUCCL RFC BGDDCPCLR A?RCEMPGCQ MD QGEL?JGLE GQ RFC BGQR?LAC RF?R RFC QGEL?J RP?TCJQ RFPMSEF RFC MPE?LGQK RM PC?AF RFC R?PECR ACJJ. /MR ?JJ ACJJQ ?PC ?DDCARCB @W RFC Q?KC QGEL?JQ.
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P'7')7/3+ !/-3'1/34GEL?JQ RF?R ?AR JMA?JJW @CRUCCL ACJJQ RF?R ?PC AJMQC RMECRFCP ?PC A?JJCB paracrine signals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synaptic signal GQ ? AFCKGA?J QGEL?J RF?R RP?TCJQ @CRUCCL LCPTC ACJJQ. 4GEL?JQ UGRFGL RFC LCPTC ACJJQ ?PC NPMN?E?RCB @W D?QRKMTGLE CJCARPGA?J GKNSJQCQ. 8FCL RFCQC GKNSJQCQ PC?AF RFC CLB MD RFC ?VML, RFC QGEL?J AMLRGLSCQ ML RM ? BCLBPGRC MD RFC LCVR ACJJ @W RFC PCJC?QC MD AFCKGA?J JGE?LBQ A?JJCB neurotransmitters @W RFC NPCQWL?NRGA ACJJ (RFC ACJJ CKGRRGLE RFC QGEL?J). 5FC LCSPMRP?LQKGRRCPQ ?PC RP?LQNMPRCB ?APMQQ RFC TCPW QK?JJ BGQR?LACQ @CRUCCL LCPTC ACJJQ, UFGAF ?PC A?JJCB chemical synapses (Figure 9.3). 5FC QK?JJ BGQR?LAC @CRUCCL LCPTC ACJJQ ?JJMUQ RFC QGEL?J RM RP?TCJ OSGAIJW; RFGQ CL?@JCQ ?L GKKCBG?RC PCQNMLQC, QSAF ?Q, 5?IC WMSP F?LB MDD RFC QRMTC! 8FCL RFC LCSPMRP?LQKGRRCP @GLBQ RFC PCACNRMP ML RFC QSPD?AC MD RFC NMQRQWL?NRGA ACJJ, RFC CJCARPMAFCKGA?J NMRCLRG?J MD RFC R?PECR ACJJ AF?LECQ, ?LB RFC LCVR CJCARPGA?J GKNSJQC GQ J?SLAFCB. 5FC LCSPMRP?LQKGRRCPQ RF?R ?PC PCJC?QCB GLRM RFC AFCKGA?J QWL?NQC ?PC BCEP?BCB OSGAIJW MP ECR PC?@QMP@CB @W RFC NPCQWL?NRGA ACJJ QM RF?R RFC PCAGNGCLR LCPTC ACJJ A?L PCAMTCP OSGAIJW ?LB @C NPCN?PCB RM PCQNMLB P?NGBJW RM RFC LCVR QWL?NRGA QGEL?J.
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E3*4)7/3+ !/-3'1/34GEL?JQ DPMK BGQR?LR ACJJQ ?PC A?JJCB endocrine signals, ?LB RFCW MPGEGL?RC DPMK endocrine cells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utocrine signals ?PC NPMBSACB @W QGEL?JGLE ACJJQ RF?R A?L ?JQM @GLB RM RFC JGE?LB RF?R GQ PCJC?QCB. 5FGQ KC?LQ RFC QGEL?JGLE ACJJ ?LB RFC R?PECR ACJJ A?L @C RFC Q?KC MP ? QGKGJ?P ACJJ (RFC NPCDGV (<;6- KC?LQ QCJD, ? PCKGLBCP RF?R RFC QGEL?JGLE ACJJ QCLBQ ? QGEL?J RM GRQCJD). 5FGQ RWNC MD QGEL?JGLE MDRCL MAASPQ BSPGLE RFC C?PJW BCTCJMNKCLR MD ?L MPE?LGQK RM CLQSPC RF?R ACJJQ BCTCJMN GLRM RFC AMPPCAR RGQQSCQ ?LB R?IC ML RFC NPMNCP DSLARGML. "SRMAPGLC QGEL?JGLE ?JQM PCESJ?RCQ N?GL QCLQ?RGML ?LB GLDJ?KK?RMPW PCQNMLQCQ. 'SPRFCP, GD ? ACJJ GQ GLDCARCB UGRF ? TGPSQ, RFC ACJJ A?L QGEL?J GRQCJD RM SLBCPEM NPMEP?KKCB ACJJ BC?RF, IGJJGLE RFC TGPSQ GL RFC NPMACQQ. *L QMKC A?QCQ, LCGEF@MPGLE ACJJQ MD RFC Q?KC RWNC ?PC ?JQM GLDJSCLACB @W RFC PCJC?QCB JGE?LB. *L CK@PWMJMEGA?J BCTCJMNKCLR, RFGQ NPMACQQ MD QRGKSJ?RGLE ? EPMSN MD LCGEF@MPGLE ACJJQ K?W FCJN RM BGPCAR RFC BGDDCPCLRG?RGML MD GBCLRGA?J ACJJQ GLRM RFC Q?KC ACJJ RWNC, RFSQ CLQSPGLE RFC NPMNCP BCTCJMNKCLR?J MSRAMKC. D/7+)9 !/-3'1/3- A)7488 G'5 J:3)9/438 (?N HSLARGMLQ GL ?LGK?JQ ?LB NJ?QKMBCQK?R? GL NJ?LRQ ?PC AMLLCARGMLQ @CRUCCL RFC NJ?QK? KCK@P?LCQ MD LCGEF@MPGLE ACJJQ. 5FCQC U?RCP-DGJJCB AF?LLCJQ ?JJMU QK?JJ QGEL?JGLE KMJCASJCQ, A?JJCB intracellular mediators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3CACNRMPQ ?PC NPMRCGL KMJCASJCQ GL RFC R?PECR ACJJ MP ML GRQ QSPD?AC RF?R @GLB JGE?LB. 5FCPC ?PC RUM RWNCQ MD PCACNRMPQ, GLRCPL?J PCACNRMPQ ?LB ACJJ-QSPD?AC PCACNRMPQ. I39+73'1 7+)+59478 Internal receptors, ?JQM ILMUL ?Q GLRP?ACJJSJ?P MP AWRMNJ?QKGA PCACNRMPQ, ?PC DMSLB GL RFC AWRMNJ?QK MD RFC ACJJ ?LB PCQNMLB RM FWBPMNFM@GA JGE?LB KMJCASJCQ RF?R ?PC ?@JC RM RP?TCJ ?APMQQ RFC NJ?QK? KCK@P?LC. 0LAC GLQGBC RFC ACJJ, K?LW MD RFCQC KMJCASJCQ @GLB RM NPMRCGLQ RF?R ?AR ?Q PCESJ?RMPQ MD K3/" QWLRFCQGQ (RP?LQAPGNRGML) RM KCBG?RC ECLC CVNPCQQGML. (CLC CVNPCQQGML GQ RFC ACJJSJ?P NPMACQQ MD RP?LQDMPKGLE RFC GLDMPK?RGML GL ? ACJJ'Q %/" GLRM ? QCOSCLAC MD ?KGLM ?AGBQ, UFGAF SJRGK?RCJW DMPKQ ? NPMRCGL. 8FCL RFC JGE?LB @GLBQ RM RFC GLRCPL?J PCACNRMP, ? AMLDMPK?RGML?J AF?LEC GQ RPGEECPCB RF?R CVNMQCQ ? %/"@GLBGLE QGRC ML RFC NPMRCGL. 5FC JGE?LB-PCACNRMP AMKNJCV KMTCQ GLRM RFC LSAJCSQ, RFCL @GLBQ RM QNCAGDGA PCESJ?RMPW PCEGMLQ MD RFC AFPMKMQMK?J %/" ?LB NPMKMRCQ RFC GLGRG?RGML MD RP?LQAPGNRGML (Figure 9.4). 5P?LQAPGNRGML GQ RFC NPMACQQ MD AMNWGLE RFC GLDMPK?RGML GL ? ACJJQ %/" GLRM ? QNCAG?J DMPK MD 3/" A?JJCB KCQQCLECP 3/" (K3/"); RFC ACJJ SQCQ GLDMPK?RGML GL RFC K3/" (UFGAF KMTCQ MSR GLRM RFC AWRMNJ?QK ?LB ?QQMAG?RCQ UGRF PG@MQMKCQ) RM JGLI QNCAGDGA ?KGLM ?AGBQ GL RFC AMPPCAR MPBCP, NPMBSAGLE ? NPMRCGL. *LRCPL?J PCACNRMPQ A?L BGPCARJW GLDJSCLAC ECLC CVNPCQQGML UGRFMSR F?TGLE RM N?QQ RFC QGEL?J ML RM MRFCP PCACNRMPQ MP KCQQCLECPQ.
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C+11-!:7,')+ +)+59478 Cell-surface receptors, ?JQM ILMUL ?Q RP?LQKCK@P?LC PCACNRMPQ, ?PC ACJJ QSPD?AC, KCK@P?LC-?LAFMPCB (GLRCEP?J) NPMRCGLQ RF?R @GLB RM CVRCPL?J JGE?LB KMJCASJCQ. 5FGQ RWNC MD PCACNRMP QN?LQ RFC NJ?QK? KCK@P?LC ?LB NCPDMPKQ QGEL?J RP?LQBSARGML, GL UFGAF ?L CVRP?ACJJSJ?P QGEL?J GQ AMLTCPRCB GLRM ?L GLRCPACJJSJ?P QGEL?J. -GE?LBQ RF?R GLRCP?AR UGRF ACJJ-QSPD?AC PCACNRMPQ BM LMR F?TC RM CLRCP RFC ACJJ RF?R RFCW ?DDCAR. $CJJ-QSPD?AC PCACNRMPQ ?PC ?JQM A?JJCB ACJJ-QNCAGDGA NPMRCGLQ MP K?PICPQ @CA?SQC RFCW ?PC QNCAGDGA RM GLBGTGBS?J ACJJ RWNCQ. #CA?SQC ACJJ-QSPD?AC PCACNRMP NPMRCGLQ ?PC DSLB?KCLR?J RM LMPK?J ACJJ DSLARGMLGLE, GR QFMSJB AMKC ?Q LM QSPNPGQC RF?R ? K?JDSLARGML GL ?LW MLC MD RFCQC NPMRCGLQ AMSJB F?TC QCTCPC AMLQCOSCLACQ. &PPMPQ GL RFC NPMRCGL QRPSARSPCQ MD ACPR?GL PCACNRMP KMJCASJCQ F?TC @CCL QFMUL RM NJ?W ? PMJC GL FWNCPRCLQGML (FGEF @JMMB NPCQQSPC), ?QRFK?, FC?PR BGQC?QC, ?LB A?LACP. &?AF ACJJ-QSPD?AC PCACNRMP F?Q RFPCC K?GL AMKNMLCLRQ: ?L CVRCPL?J JGE?LB-@GLBGLE BMK?GL, ? FWBPMNFM@GA KCK@P?LC-QN?LLGLE PCEGML, ?LB ?L GLRP?ACJJSJ?P BMK?GL GLQGBC RFC ACJJ. 5FC JGE?LB-@GLBGLE BMK?GL GQ ?JQM A?JJCB RFC extracellular domain. 5FC QGXC ?LB CVRCLR MD C?AF MD RFCQC BMK?GLQ T?PW UGBCJW, BCNCLBGLE ML RFC RWNC MD PCACNRMP.
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G-protein-linked receptors @GLB ? JGE?LB ?LB ?ARGT?RC ? KCK@P?LC NPMRCGL A?JJCB ? (-NPMRCGL. 5FC ?ARGT?RCB (-NPMRCGL RFCL GLRCP?ARQ UGRF CGRFCP ?L GML AF?LLCJ MP ?L CLXWKC GL RFC KCK@P?LC (Figure 9.6). "JJ (-NPMRCGL-JGLICB PCACNRMPQ F?TC QCTCL RP?LQKCK@P?LC BMK?GLQ, @SR C?AF PCACNRMP F?Q GRQ MUL QNCAGDGA CVRP?ACJJSJ?P BMK?GL ?LB (-NPMRCGL-@GLBGLE QGRC.
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$CJJ QGEL?JGLE SQGLE (-NPMRCGL-JGLICB PCACNRMPQ MAASPQ ?Q ? AWAJGA QCPGCQ MD CTCLRQ. #CDMPC RFC JGE?LB @GLBQ, RFC GL?ARGTC (-NPMRCGL A?L @GLB RM ? LCUJW PCTC?JCB QGRC ML RFC PCACNRMP QNCAGDGA DMP GRQ @GLBGLE. 0LAC RFC (-NPMRCGL @GLBQ RM RFC PCACNRMP, RFC PCQSJR?LR QF?NC AF?LEC ?ARGT?RCQ RFC (-NPMRCGL, UFGAF PCJC?QCQ (%1 ?LB NGAIQ SN (51. 5FC QS@SLGRQ MD RFC (-NPMRCGL RFCL QNJGR GLRM RFC C QS@SLGR ?LB RFC DE QS@SLGR. 0LC MP @MRF MD RFCQC (-NPMRCGL DP?EKCLRQ K?W @C ?@JC RM ?ARGT?RC MRFCP NPMRCGLQ ?Q ? PCQSJR. "DRCP ?UFGJC, RFC (51 ML RFC ?ARGTC C QS@SLGR MD RFC (-NPMRCGL GQ FWBPMJWXCB RM (%1 ?LB RFC DE QS@SLGR GQ BC?ARGT?RCB. 5FC QS@SLGRQ PC?QQMAG?RC RM DMPK RFC GL?ARGTC (-NPMRCGL ?LB RFC AWAJC @CEGLQ ?LCU.
!@>LI< 9.6 (BQBOLQOFJBOF@ ' MOLQBFKP E>SB QEOBB PR?RKFQP: C, D, >KA E. 7EBK > PFDK>IFKD JLIB@RIB ?FKAP QL > '-MOLQBFK-@LRMIBA OB@BMQLO FK QEB MI>PJ> JBJ?O>KB, > '$0 JLIB@RIB >PPL@F>QBA TFQE QEB C PR?RKFQ FP BU@E>KDBA CLO '40. 4EB D >KA E PR?RKFQP AFPPL@F>QB COLJ QEB C PR?RKFQ, >KA > @BIIRI>O OBPMLKPB FP QOFDDBOBA BFQEBO ?V QEB C PR?RKFQ LO QEB AFPPL@F>QBA DE M>FO. (VAOLIVPFP LC '40 QL '$0 QBOJFK>QBP QEB PFDK>I.
(-NPMRCGL-JGLICB PCACNRMPQ F?TC @CCL CVRCLQGTCJW QRSBGCB ?LB KSAF F?Q @CCL JC?PLCB ?@MSR RFCGP PMJCQ GL K?GLR?GLGLE FC?JRF. #?ARCPG? RF?R ?PC N?RFMECLGA RM FSK?LQ A?L PCJC?QC NMGQMLQ RF?R GLRCPPSNR QNCAGDGA (-NPMRCGL-JGLICB PCACNRMP DSLARGML, JC?BGLE RM GJJLCQQCQ QSAF ?Q NCPRSQQGQ, @MRSJGQK, ?LB AFMJCP?. *L AFMJCP? (Figure 9.7), DMP CV?KNJC, RFC U?RCP-@MPLC @?ARCPGSK #0)906 */63,9(, NPMBSACQ ? RMVGL, AFMJCP?ECL, RF?R @GLBQ RM ACJJQ JGLGLE RFC QK?JJ GLRCQRGLC. 5FC RMVGL RFCL CLRCPQ RFCQC GLRCQRGL?J ACJJQ, UFCPC GR KMBGDGCQ ? (-NPMRCGL RF?R AMLRPMJQ RFC MNCLGLE MD ? AFJMPGBC AF?LLCJ ?LB A?SQCQ GR RM PCK?GL AMLRGLSMSQJW ?ARGTC, PCQSJRGLE GL J?PEC JMQQCQ MD DJSGBQ DPMK RFC @MBW ?LB NMRCLRG?JJW D?R?J BCFWBP?RGML ?Q ? PCQSJR.
252
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
Enzyme-linked receptors ?PC ACJJ-QSPD?AC PCACNRMPQ UGRF GLRP?ACJJSJ?P BMK?GLQ RF?R ?PC ?QQMAG?RCB UGRF ?L CLXWKC. *L QMKC A?QCQ, RFC GLRP?ACJJSJ?P BMK?GL MD RFC PCACNRMP GRQCJD GQ ?L CLXWKC. 0RFCP CLXWKC-JGLICB PCACNRMPQ F?TC ? QK?JJ GLRP?ACJJSJ?P BMK?GL RF?R GLRCP?ARQ BGPCARJW UGRF ?L CLXWKC. 5FC CLXWKC-JGLICB PCACNRMPQ LMPK?JJW F?TC J?PEC CVRP?ACJJSJ?P ?LB GLRP?ACJJSJ?P BMK?GLQ, @SR RFC KCK@P?LCQN?LLGLE PCEGML AMLQGQRQ MD ? QGLEJC ?JNF?-FCJGA?J PCEGML MD RFC NCNRGBC QRP?LB. 8FCL ? JGE?LB @GLBQ RM RFC CVRP?ACJJSJ?P BMK?GL, ? QGEL?J GQ RP?LQDCPPCB RFPMSEF RFC KCK@P?LC, ?ARGT?RGLE RFC CLXWKC. "ARGT?RGML MD RFC CLXWKC QCRQ MDD ? AF?GL MD CTCLRQ UGRFGL RFC ACJJ RF?R CTCLRS?JJW JC?BQ RM ? PCQNMLQC. 0LC CV?KNJC MD RFGQ RWNC MD CLXWKC-JGLICB PCACNRMP GQ RFC RWPMQGLC IGL?QC PCACNRMP (Figure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!@>LI< 9.8 ! OB@BMQLO QVOLPFKB HFK>PB FP >K BKWVJB-IFKHBA OB@BMQLO TFQE > PFKDIB QO>KPJBJ?O>KB OBDFLK, >KA BUQO>@BIIRI>O >KA FKQO>@BIIRI>O ALJ>FKP. "FKAFKD LC > PFDK>IFKD JLIB@RIB QL QEB BUQO>@BIIRI>O ALJ>FK @>RPBP QEB OB@BMQLO QL AFJBOFWB. 4VOLPFKB OBPFARBP LK QEB FKQO>@BIIRI>O ALJ>FK >OB QEBK >RQLMELPMELOVI>QBA, QOFDDBOFKD > ALTKPQOB>J @BIIRI>O OBPMLKPB. 4EB PFDK>I FP QBOJFK>QBA ?V > MELPME>Q>PB QE>Q OBJLSBP QEB MELPME>QBP COLJ QEB MELPMELQVOLPFKB OBPFARBP.
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
.@>E8C@E> (FC<:LC<J
1PMBSACB @W QGEL?JGLE ACJJQ ?LB RFC QS@QCOSCLR @GLBGLE RM PCACNRMPQ GL R?PECR ACJJQ, JGE?LBQ ?AR ?Q AFCKGA?J QGEL?JQ RF?R RP?TCJ RM RFC R?PECR ACJJQ RM AMMPBGL?RC PCQNMLQCQ. 5FC RWNCQ MD KMJCASJCQ RF?R QCPTC ?Q JGE?LBQ ?PC GLAPCBG@JW T?PGCB ?LB P?LEC DPMK QK?JJ NPMRCGLQ RM QK?JJ GMLQ JGIC A?JAGSK ($? 2+).
254
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igure 9.9). 0RFCP FWBPMNFM@GA FMPKMLCQ GLAJSBC RFWPMGB FMPKMLCQ ?LB TGR?KGL %. *L MPBCP RM @C QMJS@JC GL @JMMB, FWBPMNFM@GA JGE?LBQ KSQR @GLB RM A?PPGCP NPMRCGLQ UFGJC RFCW ?PC @CGLE RP?LQNMPRCB RFPMSEF RFC @JMMBQRPC?K.
!@>LI< 9.9 3QBOLFA ELOJLKBP E>SB PFJFI>O @EBJF@>I PQOR@QROBP QL QEBFO MOB@ROPLO, @ELIBPQBOLI. "B@>RPB QEBPB JLIB@RIBP >OB PJ>II >KA EVAOLMEL?F@, QEBV @>K AFCCRPB AFOB@QIV >@OLPP QEB MI>PJ> JBJ?O>KB FKQL QEB @BII, TEBOB QEBV FKQBO>@Q TFQE FKQBOK>I OB@BMQLOP.
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
9.2 R +IFG8>8K@FE F= K?< .@>E8C

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL FMU RFC @GLBGLE MD ? JGE?LB GLGRG?RCQ QGEL?J RP?LQBSARGML RFPMSEFMSR ? ACJJ z 3CAMELGXC RFC PMJC MD NFMQNFMPWJ?RGML GL RFC RP?LQKGQQGML MD GLRP?ACJJSJ?P QGEL?JQ z &T?JS?RC RFC PMJC MD QCAMLB KCQQCLECPQ GL QGEL?J RP?LQKGQQGML 0LAC ? JGE?LB @GLBQ RM ? PCACNRMP, RFC QGEL?J GQ RP?LQKGRRCB RFPMSEF RFC KCK@P?LC ?LB GLRM RFC AWRMNJ?QK. $MLRGLS?RGML MD ? QGEL?J GL RFGQ K?LLCP GQ A?JJCB signal transduction. 4GEL?J RP?LQBSARGML MLJW MAASPQ
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UGRF ACJJ-QSPD?AC PCACNRMPQ @CA?SQC GLRCPL?J PCACNRMPQ ?PC ?@JC RM GLRCP?AR BGPCARJW UGRF %/" GL RFC LSAJCSQ RM GLGRG?RC NPMRCGL QWLRFCQGQ. 8FCL ? JGE?LB @GLBQ RM GRQ PCACNRMP, AMLDMPK?RGML?J AF?LECQ MAASP RF?R ?DDCAR RFC PCACNRMPsQ GLRP?ACJJSJ?P BMK?GL. $MLDMPK?RGML?J AF?LECQ MD RFC CVRP?ACJJSJ?P BMK?GL SNML JGE?LB @GLBGLE A?L NPMN?E?RC RFPMSEF RFC KCK@P?LC PCEGML MD RFC PCACNRMP ?LB JC?B RM ?ARGT?RGML MD RFC GLRP?ACJJSJ?P BMK?GL MP GRQ ?QQMAG?RCB NPMRCGLQ. *L QMKC A?QCQ, @GLBGLE MD RFC JGE?LB A?SQCQ dimerization MD RFC PCACNRMP, UFGAF KC?LQ RF?R RUM PCACNRMPQ @GLB RM C?AF MRFCP RM DMPK ? QR?@JC AMKNJCV A?JJCB ? BGKCP. " dimer GQ ? AFCKGA?J AMKNMSLB DMPKCB UFCL RUM KMJCASJCQ (MDRCL GBCLRGA?J) HMGL RMECRFCP. 5FC @GLBGLE MD RFC PCACNRMPQ GL RFGQ K?LLCP CL?@JCQ RFCGP GLRP?ACJJSJ?P BMK?GLQ RM AMKC GLRM AJMQC AMLR?AR ?LB ?ARGT?RC C?AF MRFCP.
B@E;@E> $E@K@8K<J 8 .@>E8C@E> +8K?N8P

"DRCP RFC JGE?LB @GLBQ RM RFC ACJJ-QSPD?AC PCACNRMP, RFC ?ARGT?RGML MD RFC PCACNRMPsQ GLRP?ACJJSJ?P AMKNMLCLRQ QCRQ MDD ? AF?GL MD CTCLRQ RF?R GQ A?JJCB ? signaling pathway MP ? QGEL?JGLE A?QA?BC. *L ? QGEL?JGLE N?RFU?W, QCAMLB KCQQCLECPQ, CLXWKCQ, ?LB ?ARGT?RCB NPMRCGLQ GLRCP?AR UGRF QNCAGDGA NPMRCGLQ, UFGAF ?PC GL RSPL ?ARGT?RCB GL ? AF?GL PC?ARGML RF?R CTCLRS?JJW JC?BQ RM ? AF?LEC GL RFC ACJJsQ CLTGPMLKCLR (Figure 9.10). 5FC CTCLRQ GL RFC A?QA?BC MAASP GL ? QCPGCQ, KSAF JGIC ? ASPPCLR DJMUQ GL ? PGTCP. *LRCP?ARGMLQ RF?R MAASP @CDMPC ? ACPR?GL NMGLR ?PC BCDGLCB ?Q SNQRPC?K CTCLRQ, ?LB CTCLRQ ?DRCP RF?R NMGLR ?PC A?JJCB BMULQRPC?K CTCLRQ.
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!@>LI< 9.10 4EB BMFABOJ>I DOLTQE C>@QLO (%'&) OB@BMQLO (%'&2) FP > OB@BMQLO QVOLPFKB HFK>PB FKSLISBA FK QEB OBDRI>QFLK LC @BII DOLTQE, TLRKA EB>IFKD, >KA QFPPRB OBM>FO. 7EBK %'& ?FKAP QL QEB %'&2, > @>P@>AB LC ALTKPQOB>J BSBKQP @>RPBP QEB @BII QL DOLT >KA AFSFAB. )C %'&2 FP >@QFS>QBA >Q FK>MMOLMOF>QB QFJBP, RK@LKQOLIIBA @BII DOLTQE (@>K@BO) J>V L@@RO.
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signal integration MD RFC N?RFU?WQ, GL UFGAF QGEL?JQ DPMK RUM MP KMPC BGDDCPCLR ACJJ-QSPD?AC PCACNRMPQ KCPEC RM ?ARGT?RC RFC Q?KC PCQNMLQC GL RFC ACJJ. 5FGQ NPMACQQ A?L CLQSPC RF?R KSJRGNJC CVRCPL?J PCOSGPCKCLRQ ?PC KCR @CDMPC ? ACJJ AMKKGRQ RM ? QNCAGDGA PCQNMLQC. 5FC CDDCARQ MD CVRP?ACJJSJ?P QGEL?JQ A?L ?JQM @C ?KNJGDGCB @W CLXWK?RGA A?QA?BCQ. "R RFC GLGRG?RGML MD RFC QGEL?J, ? QGLEJC JGE?LB @GLBQ RM ? QGLEJC PCACNRMP. )MUCTCP, ?ARGT?RGML MD ? PCACNRMP-JGLICB CLXWKC A?L ?ARGT?RC K?LW AMNGCQ MD ? AMKNMLCLR MD RFC QGEL?JGLE A?QA?BC, UFGAF ?KNJGDGCQ RFC QGEL?J.
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(<K?F;J F= $EKI8:<CCLC8I .@>E8C@E>

5FC GLBSARGML MD ? QGEL?JGLE N?RFU?W BCNCLBQ ML RFC KMBGDGA?RGML MD ? ACJJSJ?P AMKNMLCLR @W ?L CLXWKC. 5FCPC ?PC LSKCPMSQ CLXWK?RGA KMBGDGA?RGMLQ RF?R A?L MAASP, ?LB RFCW ?PC PCAMELGXCB GL RSPL @W RFC LCVR AMKNMLCLR BMULQRPC?K. 5FC DMJJMUGLE ?PC QMKC MD RFC KMPC AMKKML CTCLRQ GL GLRP?ACJJSJ?P QGEL?JGLE.
0@QCPTC ?L ?LGK?RGML MD ACJJ QGEL?JGLE ?R RFGQ site (http://openstaxcollege.org/l/cell_signals) . P.485.47>1'9/43 0LC MD RFC KMQR AMKKML AFCKGA?J KMBGDGA?RGMLQ RF?R MAASPQ GL QGEL?JGLE N?RFU?WQ GQ RFC ?BBGRGML MD ? NFMQNF?RC EPMSN (104p3) RM ? KMJCASJC QSAF ?Q ? NPMRCGL GL ? NPMACQQ A?JJCB NFMQNFMPWJ?RGML. 5FC NFMQNF?RC A?L @C ?BBCB RM ? LSAJCMRGBC QSAF ?Q (.1 RM DMPK (%1 MP (51. 1FMQNF?RCQ ?PC ?JQM MDRCL ?BBCB RM QCPGLC, RFPCMLGLC, ?LB RWPMQGLC PCQGBSCQ MD NPMRCGLQ, UFCPC RFCW PCNJ?AC RFC FWBPMVWJ EPMSN MD RFC ?KGLM ?AGB (Figure 9.11). 5FC RP?LQDCP MD RFC NFMQNF?RC GQ A?R?JWXCB @W ?L CLXWKC A?JJCB ? kinase. 7?PGMSQ IGL?QCQ ?PC L?KCB DMP RFC QS@QRP?RC RFCW NFMQNFMPWJ?RC. 1FMQNFMPWJ?RGML MD QCPGLC ?LB RFPCMLGLC PCQGBSCQ MDRCL ?ARGT?RCQ CLXWKCQ. 1FMQNFMPWJ?RGML MD RWPMQGLC PCQGBSCQ A?L CGRFCP ?DDCAR RFC ?ARGTGRW MD ?L CLXWKC MP APC?RC ? @GLBGLE QGRC RF?R GLRCP?ARQ UGRF BMULQRPC?K AMKNMLCLRQ GL RFC QGEL?JGLE A?QA?BC. 1FMQNFMPWJ?RGML K?W ?ARGT?RC MP GL?ARGT?RC CLXWKCQ, ?LB RFC PCTCPQ?J MD NFMQNFMPWJ?RGML, BCNFMQNFMPWJ?RGML @W ? NFMQNF?R?QC, UGJJ PCTCPQC RFC CDDCAR.
!@>LI< 9.11 )K MOLQBFK MELPMELOVI>QFLK, > MELPME>QB DOLRM (0/4-3 ) FP >AABA QL OBPFARBP LC QEB >JFKL >@FAP PBOFKB, QEOBLKFKB, >KA QVOLPFKB.
!+)43* M+88+3-+78 Second messengers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
258
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A?JAGSK GML AF?LLCJQ ?JJMU RFC FGEFCP JCTCJQ MD $?2+ RF?R ?PC NPCQCLR MSRQGBC RFC ACJJ (MP GL GLRP?ACJJSJ?P QRMP?EC AMKN?PRKCLRQ) RM DJMU GLRM RFC AWRMNJ?QK, UFGAF P?GQCQ RFC AMLACLRP?RGML MD AWRMNJ?QKGA $?2+. 5FC PCQNMLQC RM RFC GLAPC?QC GL $?2+ T?PGCQ, BCNCLBGLE ML RFC ACJJ RWNC GLTMJTCB. 'MP CV?KNJC, GL RFC j-ACJJQ MD RFC N?LAPC?Q, $?2+ QGEL?JGLE JC?BQ RM RFC PCJC?QC MD GLQSJGL, ?LB GL KSQAJC ACJJQ, ?L GLAPC?QC GL $?2+ JC?BQ RM KSQAJC AMLRP?ARGMLQ. "LMRFCP QCAMLB KCQQCLECP SRGJGXCB GL K?LW BGDDCPCLR ACJJ RWNCQ GQ cyclic AMP (cAMP). $WAJGA ".1 GQ QWLRFCQGXCB @W RFC CLXWKC ?BCLWJWJ AWAJ?QC DPMK "51 (Figure 9.12). 5FC K?GL PMJC MD A".1 GL ACJJQ GQ RM @GLB RM ?LB ?ARGT?RC ?L CLXWKC A?JJCB cAMP-dependent kinase (A-kinase). "-IGL?QC PCESJ?RCQ K?LW TGR?J KCR?@MJGA N?RFU?WQ: *R NFMQNFMPWJ?RCQ QCPGLC ?LB RFPCMLGLC PCQGBSCQ MD GRQ R?PECR NPMRCGLQ, ?ARGT?RGLE RFCK GL RFC NPMACQQ. "-IGL?QC GQ DMSLB GL K?LW BGDDCPCLR RWNCQ MD ACJJQ, ?LB RFC R?PECR NPMRCGLQ GL C?AF IGLB MD ACJJ ?PC BGDDCPCLR. %GDDCPCLACQ EGTC PGQC RM RFC T?PG?RGML MD RFC PCQNMLQCQ RM A".1 GL BGDDCPCLR ACJJQ.
!@>LI< 9.12 4EFP AF>DO>J PELTP QEB JB@E>KFPJ CLO QEB CLOJ>QFLK LC @V@IF@ !-0 (@!-0). @!-0 PBOSBP >P > PB@LKA JBPPBKDBO QL >@QFS>QB LO FK>@QFS>QB MOLQBFKP TFQEFK QEB @BII. 4BOJFK>QFLK LC QEB PFDK>I L@@ROP TEBK >K BKWVJB @>IIBA MELPMELAFBPQBO>PB @LKSBOQP @!-0 FKQL !-0.
1PCQCLR GL QK?JJ AMLACLRP?RGMLQ GL RFC NJ?QK? KCK@P?LC, inositol phospholipids ?PC JGNGBQ RF?R A?L ?JQM @C AMLTCPRCB GLRM QCAMLB KCQQCLECPQ. #CA?SQC RFCQC KMJCASJCQ ?PC KCK@P?LC AMKNMLCLRQ, RFCW ?PC JMA?RCB LC?P KCK@P?LC-@MSLB PCACNRMPQ ?LB A?L C?QGJW GLRCP?AR UGRF RFCK. 1FMQNF?RGBWJGLMQGRMJ (1*) GQ RFC K?GL NFMQNFMJGNGB RF?R NJ?WQ ? PMJC GL ACJJSJ?P QGEL?JGLE. &LXWKCQ ILMUL ?Q IGL?QCQ NFMQNFMPWJ?RC 1* RM DMPK 1*-NFMQNF?RC (1*1) ?LB 1*-@GQNFMQNF?RC (1*12). 5FC CLXWKC NFMQNFMJGN?QC $ AJC?TCQ 1*12 RM DMPK diacylglycerol (DAG) ?LB inositol triphosphate (IP3) (Figure 9.13). 5FCQC NPMBSARQ MD RFC AJC?T?EC MD 1*12 QCPTC ?Q QCAMLB KCQQCLECPQ. %G?AWJEJWACPMJ (%"() PCK?GLQ GL RFC NJ?QK? KCK@P?LC ?LB ?ARGT?RCQ NPMRCGL IGL?QC $ (1,$), UFGAF RFCL NFMQNFMPWJ?RCQ QCPGLC ?LB RFPCMLGLC PCQGBSCQ GL GRQ R?PECR NPMRCGLQ. *13 BGDDSQCQ GLRM RFC AWRMNJ?QK ?LB @GLBQ RM JGE?LB-E?RCB A?JAGSK AF?LLCJQ GL RFC CLBMNJ?QKGA PCRGASJSK RM PCJC?QC $?2+ RF?R AMLRGLSCQ RFC QGEL?J A?QA?BC.
#(!04%2 9 X #%,, #/--5.)#!4)/.
259
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9.3 R -<JGFEJ< KF K?< .@>E8C

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C FMU QGEL?JGLE N?RFU?WQ BGPCAR NPMRCGL CVNPCQQGML, ACJJSJ?P KCR?@MJGQK, ?LB ACJJ EPMURF z *BCLRGDW RFC DSLARGML MD 1,$ GL QGEL?J RP?LQBSARGML N?RFU?WQ z 3CAMELGXC RFC PMJC MD ?NMNRMQGQ GL RFC BCTCJMNKCLR ?LB K?GLRCL?LAC MD ? FC?JRFW MPE?LGQK *LQGBC RFC ACJJ, JGE?LBQ @GLB RM RFCGP GLRCPL?J PCACNRMPQ, ?JJMUGLE RFCK RM BGPCARJW ?DDCAR RFC ACJJsQ %/" ?LB NPMRCGL-NPMBSAGLE K?AFGLCPW. 6QGLE QGEL?J RP?LQBSARGML N?RFU?WQ, PCACNRMPQ GL RFC NJ?QK? KCK@P?LC NPMBSAC ? T?PGCRW MD CDDCARQ ML RFC ACJJ. 5FC PCQSJRQ MD QGEL?JGLE N?RFU?WQ ?PC CVRPCKCJW T?PGCB ?LB BCNCLB ML RFC RWNC MD ACJJ GLTMJTCB ?Q UCJJ ?Q RFC CVRCPL?J ?LB GLRCPL?J AMLBGRGMLQ. " QK?JJ Q?KNJGLE MD PCQNMLQCQ GQ BCQAPG@CB @CJMU.
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4MKC QGEL?J RP?LQBSARGML N?RFU?WQ PCESJ?RC RFC RP?LQAPGNRGML MD 3/". 0RFCPQ PCESJ?RC RFC RP?LQJ?RGML MD NPMRCGLQ DPMK K3/". "L CV?KNJC MD ? NPMRCGL RF?R PCESJ?RCQ RP?LQJ?RGML GL RFC LSAJCSQ GQ RFC ."1 IGL?QC &3,. &3, GQ ?ARGT?RCB GL ? NFMQNFMPWJ?RGML A?QA?BC UFCL CNGBCPK?J EPMURF D?ARMP (&(') @GLBQ RFC &(' PCACNRMP (QCC Figure 9.10). 6NML NFMQNFMPWJ?RGML, &3, CLRCPQ RFC LSAJCSQ ?LB ?ARGT?RCQ ? NPMRCGL IGL?QC RF?R, GL RSPL, PCESJ?RCQ NPMRCGL RP?LQJ?RGML (Figure 9.14).
260
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5FC QCAMLB IGLB MD NPMRCGL UGRF UFGAF 1,$ A?L GLRCP?AR GQ ? NPMRCGL RF?R ?ARQ ?Q ?L GLFG@GRMP. "L inhibitor GQ ? KMJCASJC RF?R @GLBQ RM ? NPMRCGL ?LB NPCTCLRQ GR DPMK DSLARGMLGLE MP PCBSACQ GRQ DSLARGML. *L RFGQ A?QC, RFC GLFG@GRMP GQ ? NPMRCGL A?JJCB *k-#, UFGAF @GLBQ RM RFC PCESJ?RMPW NPMRCGL /'-k#. (5FC QWK@MJ k PCNPCQCLRQ RFC (PCCI JCRRCP I?NN?.) 8FCL *k-# GQ @MSLB RM /'-k#, RFC AMKNJCV A?LLMR CLRCP RFC LSAJCSQ MD RFC ACJJ, @SR UFCL *k-# GQ NFMQNFMPWJ?RCB @W 1,$, GR A?L LM JMLECP @GLB /'-k#, ?LB /'k# (? RP?LQAPGNRGML D?ARMP) A?L CLRCP RFC LSAJCSQ ?LB GLGRG?RC 3/" RP?LQAPGNRGML. *L RFGQ A?QC, RFC CDDCAR MD NFMQNFMPWJ?RGML GQ RM GL?ARGT?RC ?L GLFG@GRMP ?LB RFCPC@W ?ARGT?RC RFC NPMACQQ MD RP?LQAPGNRGML.
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5FC PCQSJR MD ?LMRFCP QGEL?JGLE N?RFU?W ?DDCARQ KSQAJC ACJJQ. 5FC ?ARGT?RGML MD j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qRFC tDGEFR MP DJGEFRu PCDJCV.
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$CJJ QGEL?JGLE N?RFU?WQ ?JQM NJ?W ? K?HMP PMJC GL ACJJ BGTGQGML. $CJJQ BM LMR LMPK?JJW BGTGBC SLJCQQ RFCW ?PC QRGKSJ?RCB @W QGEL?JQ DPMK MRFCP ACJJQ. 5FC JGE?LBQ RF?R NPMKMRC ACJJ EPMURF ?PC A?JJCB growth factors. .MQR EPMURF D?ARMPQ @GLB RM ACJJ-QSPD?AC PCACNRMPQ RF?R ?PC JGLICB RM RWPMQGLC IGL?QCQ. 5FCQC ACJJQSPD?AC PCACNRMPQ ?PC A?JJCB PCACNRMP RWPMQGLC IGL?QCQ (35,Q). "ARGT?RGML MD 35,Q GLGRG?RCQ ? QGEL?JGLE N?RFU?W RF?R GLAJSBCQ ? (-NPMRCGL A?JJCB 3"4, UFGAF ?ARGT?RCQ RFC ."1 IGL?QC N?RFU?W BCQAPG@CB C?PJGCP. 5FC CLXWKC ."1 IGL?QC RFCL QRGKSJ?RCQ RFC CVNPCQQGML MD NPMRCGLQ RF?R GLRCP?AR UGRF MRFCP ACJJSJ?P AMKNMLCLRQ RM GLGRG?RC ACJJ BGTGQGML.
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261
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8FCL ? ACJJ GQ B?K?ECB, QSNCPDJSMSQ, MP NMRCLRG?JJW B?LECPMSQ RM ?L MPE?LGQK, ? ACJJ A?L GLGRG?RC ? KCAF?LGQK RM RPGEECP NPMEP?KKCB ACJJ BC?RF, MP apoptosis. "NMNRMQGQ ?JJMUQ ? ACJJ RM BGC GL ? AMLRPMJJCB K?LLCP RF?R NPCTCLRQ RFC PCJC?QC MD NMRCLRG?JJW B?K?EGLE KMJCASJCQ DPMK GLQGBC RFC ACJJ. 5FCPC ?PC K?LW GLRCPL?J AFCAINMGLRQ RF?R KMLGRMP ? ACJJsQ FC?JRF; GD ?@LMPK?JGRGCQ ?PC M@QCPTCB, ? ACJJ A?L QNMLR?LCMSQJW GLGRG?RC RFC NPMACQQ MD ?NMNRMQGQ. )MUCTCP, GL QMKC A?QCQ, QSAF ?Q ? TGP?J GLDCARGML MP SLAMLRPMJJCB ACJJ BGTGQGML BSC RM A?LACP, RFC ACJJsQ LMPK?J AFCAIQ ?LB @?J?LACQ D?GJ. &VRCPL?J QGEL?JGLE A?L ?JQM GLGRG?RC ?NMNRMQGQ. 'MP CV?KNJC, KMQR LMPK?J ?LGK?J ACJJQ F?TC PCACNRMPQ RF?R GLRCP?AR UGRF RFC CVRP?ACJJSJ?P K?RPGV, ? LCRUMPI MD EJWAMNPMRCGLQ RF?R NPMTGBCQ QRPSARSP?J QSNNMPR DMP ACJJQ GL ?L MPE?LGQK. 5FC @GLBGLE MD ACJJSJ?P PCACNRMPQ RM RFC CVRP?ACJJSJ?P K?RPGV GLGRG?RCQ ? QGEL?JGLE A?QA?BC UGRFGL RFC ACJJ. )MUCTCP, GD RFC ACJJ KMTCQ ?U?W DPMK RFC CVRP?ACJJSJ?P K?RPGV, RFC QGEL?JGLE AC?QCQ, ?LB RFC ACJJ SLBCPEMCQ ?NMNRMQGQ. 5FGQ QWQRCK ICCNQ ACJJQ DPMK RP?TCJGLE RFPMSEF RFC @MBW ?LB NPMJGDCP?RGLE MSR MD AMLRPMJ, ?Q F?NNCLQ UGRF RSKMP ACJJQ RF?R KCR?QR?QGXC. "LMRFCP CV?KNJC MD CVRCPL?J QGEL?JGLE RF?R JC?BQ RM ?NMNRMQGQ MAASPQ GL 5-ACJJ BCTCJMNKCLR. 5-ACJJQ ?PC GKKSLC ACJJQ RF?R @GLB RM DMPCGEL K?APMKMJCASJCQ ?LB N?PRGAJCQ, ?LB R?PECR RFCK DMP BCQRPSARGML @W RFC GKKSLC QWQRCK. /MPK?JJW, 5-ACJJQ BM LMR R?PECR tQCJDu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igure 9.15). %SPGLE RFC AMSPQC MD LMPK?J BCTCJMNKCLR, RFCQC SLLCCBCB ACJJQ KSQR @C CJGKGL?RCB, CL?@JGLE DSJJW QCN?P?RCB DGLECPQ ?LB RMCQ RM DMPK. " ACJJ QGEL?JGLE KCAF?LGQK RPGEECPQ ?NMNRMQGQ, UFGAF BCQRPMWQ RFC ACJJQ @CRUCCL RFC BCTCJMNGLE BGEGRQ.
262
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/<ID@E8K@FE F= K?< .@>E8C C8J:8;<

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phosphatases ?PC CLXWKCQ RF?R PCKMTC RFC NFMQNF?RC EPMSN ?RR?AFCB RM NPMRCGLQ @W IGL?QCQ GL ? NPMACQQ A?JJCB BCNFMQNFMPWJ?RGML. $WAJGA ".1 (A".1) GQ BCEP?BCB GLRM ".1 @W phosphodiesterase, ?LB RFC PCJC?QC MD A?JAGSK QRMPCQ GQ PCTCPQCB @W RFC $?2+ NSKNQ RF?R ?PC JMA?RCB GL RFC CVRCPL?J ?LB GLRCPL?J KCK@P?LCQ MD RFC ACJJ.
9.4 R .@>E8C@E> @E .@E>C<-C<CC<; *I>8E@JDJ

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C FMU QGLEJC-ACJJCB WC?QRQ SQC ACJJ QGEL?JGLE RM AMKKSLGA?RC UGRF MLC ?LMRFCP z 3CJ?RC RFC PMJC MD OSMPSK QCLQGLE RM RFC ?@GJGRW MD QMKC @?ARCPG? RM DMPK @GMDGJKQ 8GRFGL-ACJJ QGEL?JGLE ?JJMUQ @?ARCPG? RM PCQNMLB RM CLTGPMLKCLR?J ASCQ, QSAF ?Q LSRPGCLR JCTCJQ, QMKC QGLEJC-ACJJCB MPE?LGQKQ ?JQM PCJC?QC KMJCASJCQ RM QGEL?J RM C?AF MRFCP.
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263
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5FC GLGRG?RGML MD ? QGEL?JGLE N?RFU?W GQ ? PCQNMLQC RM CVRCPL?J QRGKSJG. 5FGQ PCQNMLQC A?L R?IC K?LW BGDDCPCLR DMPKQ, GLAJSBGLE NPMRCGL QWLRFCQGQ, ? AF?LEC GL RFC ACJJs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
9.4 .@>E8C@E> @E .@E>C<-C<CC<; *I>8E@JDJ

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
A-/ C*)) C/$*) ,0 ./$*).

1. Figure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igure 9.10 *L ACPR?GL A?LACPQ, RFC (51?QC ?ARGTGRW MD RFC 3"4 (-NPMRCGL GQ GLFG@GRCB. 5FGQ KC?LQ RF?R RFC 3"4 NPMRCGL A?L LM JMLECP FWBPMJWXC (51 GLRM (%1. 8F?R CDDCAR UMSJB RFGQ F?TC ML BMULQRPC?K ACJJSJ?P CTCLRQ 3. Figure 9.17 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR OSMPSK QCLQGLE GQ D?JQC >. "SRMGLBSACP KSQR @GLB RM PCACNRMP RM RSPL ML RP?LQAPGNRGML MD ECLCQ PCQNMLQG@JC DMP RFC NPMBSARGML MD KMPC ?SRMGLBSACP. ?. 5FC PCACNRMP QR?WQ GL RFC @?ARCPG?J ACJJ, @SR RFC ?SRMGLBSACP BGDDSQCQ MSR. @. "SRMGLBSACP A?L MLJW ?AR ML ? BGDDCPCLR ACJJ: GR A?LLMR ?AR ML RFC ACJJ GL UFGAF GR GQ K?BC.
270
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A. "SRMGLBSACP RSPLQ ML ECLCQ RF?R CL?@JC RFC @?ARCPG? RM DMPK ? @GMDGJK.
4. Figure 9.18 8F?R ?BT?LR?EC KGEFR @GMDGJK NPMBSARGML AMLDCP ML RFC . (<9,<: GLQGBC RFC A?RFCRCP
- 1$ 2 ,0 ./$*).
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k# GLBSAC ECLC CVNPCQQGML >. " QK?JJ, FWBPMNFM@GA JGE?LB @GLBQ RM /'-k#, ?ARGT?RGLE GR. ?. 1FMQNFMPWJ?RGML MD RFC GLFG@GRMP *k-# BGQQMAG?RCQ RFC AMKNJCV @CRUCCL GR ?LB /'-k#, ?LB ?JJMUQ /'-k# RM CLRCP RFC LSAJCSQ ?LB QRGKSJ?RC RP?LQAPGNRGML. @. /'-k# GQ NFMQNFMPWJ?RCB ?LB GQ RFCL DPCC RM CLRCP RFC LSAJCSQ ?LB @GLB %/". A. /'-k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
7. 8FW ?PC GML AF?LLCJQ LCACQQ?PW RM RP?LQNMPR GMLQ GLRM MP MSR MD ? ACJJ >. *MLQ ?PC RMM J?PEC RM BGDDSQC RFPMSEF RFC KCK@P?LC. ?. *MLQ ?PC AF?PECB N?PRGAJCQ ?LB A?LLMR BGDDSQC RFPMSEF RFC FWBPMNFM@GA GLRCPGMP MD RFC KCK@P?LC. @. *MLQ BM LMR LCCB GML AF?LLCJQ RM KMTC RFPMSEF RFC KCK@P?LC. A. *MLQ @GLB RM A?PPGCP NPMRCGLQ GL RFC @JMMBQRPC?K, UFGAF KSQR @C PCKMTCB @CDMPC RP?LQNMPR GLRM RFC ACJJ. 8. &LBMAPGLC QGEL?JQ ?PC RP?LQKGRRCB KMPC QJMUJW RF?L N?P?APGLC QGEL?JQ @CA?SQC >>>>>>>>>>>. >. RFC JGE?LBQ ?PC RP?LQNMPRCB RFPMSEF RFC @JMMBQRPC?K ?LB RP?TCJ EPC?RCP BGQR?LACQ ?. RFC R?PECR ?LB QGEL?JGLE ACJJQ ?PC AJMQC RMECRFCP @. RFC JGE?LBQ ?PC BCEP?BCB P?NGBJW A. RFC JGE?LBQ BML'R @GLB RM A?PPGCP NPMRCGLQ BSPGLE RP?LQNMPR 9. 8FCPC BM %"( ?LB *13 MPGEGL?RC >. 5FCW ?PC DMPKCB @W NFMQNFMPWJ?RGML MD A".1. ?. 5FCW ?PC JGE?LBQ CVNPCQQCB @W QGEL?JGLE ACJJQ. @. 5FCW ?PC FMPKMLCQ RF?R BGDDSQC RFPMSEF RFC NJ?QK? KCK@P?LC RM QRGKSJ?RC NPMRCGL NPMBSARGML. A. 5FCW ?PC RFC AJC?T?EC NPMBSARQ MD RFC GLMQGRMJ NFMQNFMJGNGB, 1*12.
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271
16. 2SMPSK QCLQGLE GQ RPGEECPCB RM @CEGL UFCL >>>>>>>>>>>. >. RPC?RKCLR UGRF ?LRG@GMRGAQ MAASPQ ?. @?ARCPG? PCJC?QC EPMURF FMPKMLCQ
@. @?ARCPG?J NPMRCGL CVNPCQQGML GQ QUGRAFCB ML A. ? QSDDGAGCLR LSK@CP MD @?ARCPG? ?PC NPCQCLR
C-$/$CA' /#$)&$)" ,0 ./$*).

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
272
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10.1: C<CC D@M@J@FE 10.2: /?< C<CC CP:C< 10.3: CFEKIFC F= K?< C<CC CP:C< 10.4: C8E:<I 8E; K?< C<CC CP:C< 10.5: +IFB8IPFK@: C<CC D@M@J@FE
$EKIF;L:K@FE
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
10.1 R C<CC D@M@J@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC QRPSARSPC MD NPMI?PWMRGA ?LB CSI?PWMRGA ECLMKCQ z %GQRGLESGQF @CRUCCL AFPMKMQMKCQ, ECLCQ, ?LB RP?GRQ z %CQAPG@C RFC KCAF?LGQKQ MD AFPMKMQMKC AMKN?ARGML 5FC AMLRGLSGRW MD JGDC DPMK MLC ACJJ RM ?LMRFCP F?Q GRQ DMSLB?RGML GL RFC PCNPMBSARGML MD ACJJQ @W U?W MD RFC ACJJ AWAJC. 5FC cell cycle GQ ?L MPBCPJW QCOSCLAC MD CTCLRQ RF?R BCQAPG@CQ RFC QR?ECQ MD ? ACJJsQ JGDC DPMK RFC BGTGQGML MD ? QGLEJC N?PCLR ACJJ RM RFC NPMBSARGML MD RUM LCU B?SEFRCP ACJJQ. 5FC KCAF?LGQKQ GLTMJTCB GL RFC ACJJ AWAJC ?PC FGEFJW PCESJ?RCB.
274
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#CDMPC BGQASQQGLE RFC QRCNQ ? ACJJ KSQR SLBCPR?IC RM PCNJGA?RC, ? BCCNCP SLBCPQR?LBGLE MD RFC QRPSARSPC ?LB DSLARGML MD ? ACJJsQ ECLCRGA GLDMPK?RGML GQ LCACQQ?PW. " ACJJsQ %/", N?AI?ECB ?Q ? BMS@JC-QRP?LBCB %/" KMJCASJC, GQ A?JJCB GRQ genome. *L NPMI?PWMRCQ, RFC ECLMKC GQ AMKNMQCB MD ? QGLEJC, BMS@JCQRP?LBCB %/" KMJCASJC GL RFC DMPK MD ? JMMN MP AGPAJC (Figure 10.2). 5FC PCEGML GL RFC ACJJ AMLR?GLGLE RFGQ ECLCRGA K?RCPG?J GQ A?JJCB ? LSAJCMGB. 4MKC NPMI?PWMRCQ ?JQM F?TC QK?JJCP JMMNQ MD %/" A?JJCB NJ?QKGBQ RF?R ?PC LMR CQQCLRG?J DMP LMPK?J EPMURF. #?ARCPG? A?L CVAF?LEC RFCQC NJ?QKGBQ UGRF MRFCP @?ARCPG?, QMKCRGKCQ PCACGTGLE @CLCDGAG?J LCU ECLCQ RF?R RFC PCAGNGCLR A?L ?BB RM RFCGP AFPMKMQMK?J %/". "LRG@GMRGA PCQGQR?LAC GQ MLC RP?GR RF?R MDRCL QNPC?BQ RFPMSEF ? @?ARCPG?J AMJMLW RFPMSEF NJ?QKGB CVAF?LEC.
!@>LI< 10.2 0OLH>OVLQBP, FK@IRAFKD ?>@QBOF> >KA >O@E>B>, E>SB > PFKDIB, @FO@RI>O @EOLJLPLJB IL@>QBA FK > @BKQO>I OBDFLK @>IIBA QEB KR@IBLFA.
*L CSI?PWMRCQ, RFC ECLMKC AMLQGQRQ MD QCTCP?J BMS@JC-QRP?LBCB JGLC?P %/" KMJCASJCQ (Figure 10.3). &?AF QNCAGCQ MD CSI?PWMRCQ F?Q ? AF?P?ARCPGQRGA LSK@CP MD AFPMKMQMKCQ GL RFC LSAJCG MD GRQ ACJJQ. )SK?L @MBW ACJJQ F?TC 46 AFPMKMQMKCQ, UFGJC FSK?L gametes (QNCPK MP CEEQ) F?TC 23 AFPMKMQMKCQ C?AF. " RWNGA?J @MBW ACJJ, MP QMK?RGA ACJJ, AMLR?GLQ RUM K?RAFCB QCRQ MD AFPMKMQMKCQ, ? AMLDGESP?RGML ILMUL ?Q diploid. 5FC JCRRCP 5 GQ SQCB RM PCNPCQCLR ? QGLEJC QCR MD AFPMKMQMKCQ; RFCPCDMPC, ? BGNJMGB MPE?LGQK GQ BCQGEL?RCB 25. )SK?L ACJJQ RF?R AMLR?GL MLC QCR MD AFPMKMQMKCQ ?PC A?JJCB E?KCRCQ, MP QCV ACJJQ; RFCQC ?PC CEEQ ?LB QNCPK, ?LB ?PC BCQGEL?RCB 15, MP haploid.
!@>LI< 10.3 4EBOB >OB 23 M>FOP LC ELJLILDLRP @EOLJLPLJBP FK > CBJ>IB ERJ>K PLJ>QF@ @BII. 4EB @LKABKPBA @EOLJLPLJBP >OB SFBTBA TFQEFK QEB KR@IBRP (QLM), OBJLSBA COLJ > @BII FK JFQLPFP >KA PMOB>A LRQ LK > PIFAB (OFDEQ), >KA >OQFCF@F>IIV >OO>KDBA >@@LOAFKD QL IBKDQE (IBCQ); >K >OO>KDBJBKQ IFHB QEFP FP @>IIBA > H>OVLQVMB. )K QEFP FJ>DB, QEB @EOLJLPLJBP TBOB BUMLPBA QL CIRLOBP@BKQ PQ>FKP CLO AFCCBOBKQF>QFLK LC QEB AFCCBOBKQ @EOLJLPLJBP. ! JBQELA LC PQ>FKFKD @>IIBA x@EOLJLPLJB M>FKQFKDy BJMILVP CIRLOBP@BKQ AVBP QE>Q EFDEIFDEQ @EOLJLPLJBP FK AFCCBOBKQ @LILOP. (@OBAFQ: .>QFLK>I (RJ>K 'BKLJB 0OLGB@Q/.)()
.?RAFCB N?GPQ MD AFPMKMQMKCQ GL ? BGNJMGB MPE?LGQK ?PC A?JJCB homologous (tQ?KC ILMUJCBECu) chromosomes. )MKMJMEMSQ AFPMKMQMKCQ ?PC RFC Q?KC JCLERF ?LB F?TC QNCAGDGA LSAJCMRGBC QCEKCLRQ A?JJCB genes GL CV?ARJW RFC Q?KC JMA?RGML, MP locus. (CLCQ, RFC DSLARGML?J SLGRQ MD AFPMKMQMKCQ,
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BCRCPKGLC QNCAGDGA AF?P?ARCPGQRGAQ @W AMBGLE DMP QNCAGDGA NPMRCGLQ. 5P?GRQ ?PC RFC T?PG?RGMLQ MD RFMQC AF?P?ARCPGQRGAQ. 'MP CV?KNJC, F?GP AMJMP GQ ? AF?P?ARCPGQRGA UGRF RP?GRQ RF?R ?PC @JMLBC, @PMUL, MP @J?AI. &?AF AMNW MD ? FMKMJMEMSQ N?GP MD AFPMKMQMKCQ MPGEGL?RCQ DPMK ? BGDDCPCLR N?PCLR; RFCPCDMPC, RFC ECLCQ RFCKQCJTCQ ?PC LMR GBCLRGA?J. 5FC T?PG?RGML MD GLBGTGBS?JQ UGRFGL ? QNCAGCQ GQ BSC RM RFC QNCAGDGA AMK@GL?RGML MD RFC ECLCQ GLFCPGRCB DPMK @MRF N?PCLRQ. &TCL ? QJGEFRJW ?JRCPCB QCOSCLAC MD LSAJCMRGBCQ UGRFGL ? ECLC A?L PCQSJR GL ?L ?JRCPL?RGTC RP?GR. 'MP CV?KNJC, RFCPC ?PC RFPCC NMQQG@JC ECLC QCOSCLACQ ML RFC FSK?L AFPMKMQMKC RF?R AMBC DMP @JMMB RWNC: QCOSCLAC ", QCOSCLAC #, ?LB QCOSCLAC 0. #CA?SQC ?JJ BGNJMGB FSK?L ACJJQ F?TC RUM AMNGCQ MD RFC AFPMKMQMKC RF?R BCRCPKGLCQ @JMMB RWNC, RFC @JMMB RWNC (RFC RP?GR) GQ BCRCPKGLCB @W UFGAF RUM TCPQGMLQ MD RFC K?PICP ECLC ?PC GLFCPGRCB. *R GQ NMQQG@JC RM F?TC RUM AMNGCQ MD RFC Q?KC ECLC QCOSCLAC ML @MRF FMKMJMEMSQ AFPMKMQMKCQ, UGRF MLC ML C?AF (DMP CV?KNJC, "", ##, MP 00), MP RUM BGDDCPCLR QCOSCLACQ, QSAF ?Q "#. .GLMP T?PG?RGMLQ MD RP?GRQ, QSAF ?Q @JMMB RWNC, CWC AMJMP, ?LB F?LBCBLCQQ, AMLRPG@SRC RM RFC L?RSP?J T?PG?RGML DMSLB UGRFGL ? QNCAGCQ. )MUCTCP, GD RFC CLRGPC %/" QCOSCLAC DPMK ?LW N?GP MD FSK?L FMKMJMEMSQ AFPMKMQMKCQ GQ AMKN?PCB, RFC BGDDCPCLAC GQ JCQQ RF?L MLC NCPACLR. 5FC QCV AFPMKMQMKCQ, 9 ?LB :, ?PC RFC QGLEJC CVACNRGML RM RFC PSJC MD FMKMJMEMSQ AFPMKMQMKC SLGDMPKGRW: 0RFCP RF?L ? QK?JJ ?KMSLR MD FMKMJMEW RF?R GQ LCACQQ?PW RM ?AASP?RCJW NPMBSAC E?KCRCQ, RFC ECLCQ DMSLB ML RFC 9 ?LB : AFPMKMQMKCQ ?PC BGDDCPCLR.
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*D RFC %/" DPMK ?JJ 46 AFPMKMQMKCQ GL ? FSK?L ACJJ LSAJCSQ U?Q J?GB MSR CLB RM CLB, GR UMSJB KC?QSPC ?NNPMVGK?RCJW RUM KCRCPQ; FMUCTCP, GRQ BG?KCRCP UMSJB @C MLJW 2 LK. $MLQGBCPGLE RF?R RFC QGXC MD ? RWNGA?J FSK?L ACJJ GQ ?@MSR 10 ]K (100,000 ACJJQ JGLCB SN RM COS?J MLC KCRCP), %/" KSQR @C RGEFRJW N?AI?ECB RM DGR GL RFC ACJJsQ LSAJCSQ. "R RFC Q?KC RGKC, GR KSQR ?JQM @C PC?BGJW ?AACQQG@JC DMP RFC ECLCQ RM @C CVNPCQQCB. %SPGLE QMKC QR?ECQ MD RFC ACJJ AWAJC, RFC JMLE QRP?LBQ MD %/" ?PC AMLBCLQCB GLRM AMKN?AR AFPMKMQMKCQ. 5FCPC ?PC ? LSK@CP MD U?WQ RF?R AFPMKMQMKCQ ?PC AMKN?ARCB. *L RFC DGPQR JCTCJ MD AMKN?ARGML, QFMPR QRPCRAFCQ MD RFC %/" BMS@JC FCJGV UP?N ?PMSLB ? AMPC MD CGEFR histone proteins ?R PCESJ?P GLRCPT?JQ ?JMLE RFC CLRGPC JCLERF MD RFC AFPMKMQMKC (Figure 10.4). 5FC %/"FGQRMLC AMKNJCV GQ A?JJCB AFPMK?RGL. 5FC @C?BJGIC, FGQRMLC %/" AMKNJCV GQ A?JJCB ? nucleosome, ?LB %/" AMLLCARGLE RFC LSAJCMQMKCQ GQ A?JJCB JGLICP %/". " %/" KMJCASJC GL RFGQ DMPK GQ ?@MSR QCTCL RGKCQ QFMPRCP RF?L RFC BMS@JC FCJGV UGRFMSR RFC FGQRMLCQ, ?LB RFC @C?BQ ?PC ?@MSR 10 LK GL BG?KCRCP, GL AMLRP?QR UGRF RFC 2-LK BG?KCRCP MD ? %/" BMS@JC FCJGV. 5FC LCVR JCTCJ MD AMKN?ARGML MAASPQ ?Q RFC LSAJCMQMKCQ ?LB RFC JGLICP %/" @CRUCCL RFCK ?PC AMGJCB GLRM ? 30-LK AFPMK?RGL DG@CP. 5FGQ AMGJGLE DSPRFCP QFMPRCLQ RFC AFPMKMQMKC QM RF?R GR GQ LMU ?@MSR 50 RGKCQ QFMPRCP RF?L RFC CVRCLBCB DMPK. *L RFC RFGPB JCTCJ MD N?AIGLE, ? T?PGCRW MD DG@PMSQ NPMRCGLQ GQ SQCB RM N?AI RFC AFPMK?RGL. 5FCQC DG@PMSQ NPMRCGLQ ?JQM CLQSPC RF?R C?AF AFPMKMQMKC GL ? LML-BGTGBGLE ACJJ MAASNGCQ ? N?PRGASJ?P ?PC? MD RFC LSAJCSQ RF?R BMCQ LMR MTCPJ?N UGRF RF?R MD ?LW MRFCP AFPMKMQMKC (QCC RFC RMN GK?EC GL Figure 10.3).
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%/" PCNJGA?RCQ GL RFC 4 NF?QC MD GLRCPNF?QC. "DRCP PCNJGA?RGML, RFC AFPMKMQMKCQ ?PC AMKNMQCB MD RUM JGLICB QGQRCP chromatids. 8FCL DSJJW AMKN?AR, RFC N?GPQ MD GBCLRGA?JJW N?AICB AFPMKMQMKCQ ?PC @MSLB RM C?AF MRFCP @W AMFCQGL NPMRCGLQ. 5FC AMLLCARGML @CRUCCL RFC QGQRCP AFPMK?RGBQ GQ AJMQCQR GL ? PCEGML A?JJCB RFC centromere. 5FC AMLHMGLCB QGQRCP AFPMK?RGBQ, UGRF ? BG?KCRCP MD ?@MSR 1 ]K, ?PC TGQG@JC SLBCP ? JGEFR KGAPMQAMNC. 5FC ACLRPMKCPGA PCEGML GQ FGEFJW AMLBCLQCB ?LB RFSQ UGJJ ?NNC?P ?Q ? AMLQRPGARCB ?PC?.
This animation (http://openstaxcollege.org/l/Packaged_DNA) GJJSQRP?RCQ RFC BGDDCPCLR JCTCJQ MD AFPMKMQMKC N?AIGLE.
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC RFPCC QR?ECQ MD GLRCPNF?QC z %GQASQQ RFC @CF?TGMP MD AFPMKMQMKCQ BSPGLE I?PWMIGLCQGQ z &VNJ?GL FMU RFC AWRMNJ?QKGA AMLRCLR GQ BGTGBCB BSPGLE AWRMIGLCQGQ z %CDGLC RFC OSGCQACLR (0 NF?QC 5FC cell cycle GQ ?L MPBCPCB QCPGCQ MD CTCLRQ GLTMJTGLE ACJJ EPMURF ?LB ACJJ BGTGQGML RF?R NPMBSACQ RUM LCU B?SEFRCP ACJJQ. $CJJQ ML RFC N?RF RM ACJJ BGTGQGML NPMACCB RFPMSEF ? QCPGCQ MD NPCAGQCJW RGKCB ?LB A?PCDSJJW PCESJ?RCB QR?ECQ MD EPMURF, %/" PCNJGA?RGML, ?LB BGTGQGML RF?R NPMBSACQ RUM GBCLRGA?J (AJMLC) ACJJQ. 5FC ACJJ AWAJC F?Q RUM K?HMP NF?QCQ: GLRCPNF?QC ?LB RFC KGRMRGA NF?QC (Figure 10.5). %SPGLE interphase, RFC ACJJ EPMUQ ?LB %/" GQ PCNJGA?RCB. %SPGLE RFC mitotic phase, RFC PCNJGA?RCB %/" ?LB AWRMNJ?QKGA AMLRCLRQ ?PC QCN?P?RCB, ?LB RFC ACJJ BGTGBCQ.
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%SPGLE GLRCPNF?QC, RFC ACJJ SLBCPEMCQ LMPK?J EPMURF NPMACQQCQ UFGJC ?JQM NPCN?PGLE DMP ACJJ BGTGQGML. *L MPBCP DMP ? ACJJ RM KMTC DPMK GLRCPNF?QC GLRM RFC KGRMRGA NF?QC, K?LW GLRCPL?J ?LB CVRCPL?J AMLBGRGMLQ KSQR @C KCR. 5FC RFPCC QR?ECQ MD GLRCPNF?QC ?PC A?JJCB (1, 4, ?LB (2. G1 P.'8+ (F/789 G'5) 5FC DGPQR QR?EC MD GLRCPNF?QC GQ A?JJCB RFC G1 phase (DGPQR E?N) @CA?SQC, DPMK ? KGAPMQAMNGA ?QNCAR, JGRRJC AF?LEC GQ TGQG@JC. )MUCTCP, BSPGLE RFC (1 QR?EC, RFC ACJJ GQ OSGRC ?ARGTC ?R RFC @GMAFCKGA?J JCTCJ. 5FC ACJJ GQ ?AASKSJ?RGLE RFC @SGJBGLE @JMAIQ MD AFPMKMQMK?J %/" ?LB RFC ?QQMAG?RCB NPMRCGLQ ?Q UCJJ ?Q ?AASKSJ?RGLE QSDDGAGCLR CLCPEW PCQCPTCQ RM AMKNJCRC RFC R?QI MD PCNJGA?RGLE C?AF AFPMKMQMKC GL RFC LSAJCSQ. ! P.'8+ (!>39.+8/8 4, DNA) 5FPMSEFMSR GLRCPNF?QC, LSAJC?P %/" PCK?GLQ GL ? QCKG-AMLBCLQCB AFPMK?RGL AMLDGESP?RGML. *L RFC S phase, %/" PCNJGA?RGML A?L NPMACCB RFPMSEF RFC KCAF?LGQKQ RF?R PCQSJR GL RFC DMPK?RGML MD GBCLRGA?J
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N?GPQ MD %/" KMJCASJCQqQGQRCP AFPMK?RGBQqRF?R ?PC DGPKJW ?RR?AFCB RM RFC ACLRPMKCPGA PCEGML. 5FC ACLRPMQMKC GQ BSNJGA?RCB BSPGLE RFC 4 NF?QC. 5FC RUM ACLRPMQMKCQ UGJJ EGTC PGQC RM RFC mitotic spindle, RFC ?NN?P?RSQ RF?R MPAFCQRP?RCQ RFC KMTCKCLR MD AFPMKMQMKCQ BSPGLE KGRMQGQ. "R RFC ACLRCP MD C?AF ?LGK?J ACJJ, RFC ACLRPMQMKCQ MD ?LGK?J ACJJQ ?PC ?QQMAG?RCB UGRF ? N?GP MD PMB-JGIC M@HCARQ, RFC centrioles, UFGAF ?PC ?R PGEFR ?LEJCQ RM C?AF MRFCP. $CLRPGMJCQ FCJN MPE?LGXC ACJJ BGTGQGML. $CLRPGMJCQ ?PC LMR NPCQCLR GL RFC ACLRPMQMKCQ MD MRFCP CSI?PWMRGA QNCAGCQ, QSAF ?Q NJ?LRQ ?LB KMQR DSLEG. G2 P.'8+ (!+)43* G'5) *L RFC G2 phase, RFC ACJJ PCNJCLGQFCQ GRQ CLCPEW QRMPCQ ?LB QWLRFCQGXCQ NPMRCGLQ LCACQQ?PW DMP AFPMKMQMKC K?LGNSJ?RGML. 4MKC ACJJ MPE?LCJJCQ ?PC BSNJGA?RCB, ?LB RFC AWRMQICJCRML GQ BGQK?LRJCB RM NPMTGBC PCQMSPACQ DMP RFC KGRMRGA NF?QC. 5FCPC K?W @C ?BBGRGML?J ACJJ EPMURF BSPGLE (2. 5FC DGL?J NPCN?P?RGMLQ DMP RFC KGRMRGA NF?QC KSQR @C AMKNJCRCB @CDMPC RFC ACJJ GQ ?@JC RM CLRCP RFC DGPQR QR?EC MD KGRMQGQ.
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5FC KGRMRGA NF?QC GQ ? KSJRGQRCN NPMACQQ BSPGLE UFGAF RFC BSNJGA?RCB AFPMKMQMKCQ ?PC ?JGELCB, QCN?P?RCB, ?LB KMTC GLRM RUM LCU, GBCLRGA?J B?SEFRCP ACJJQ. 5FC DGPQR NMPRGML MD RFC KGRMRGA NF?QC GQ A?JJCB karyokinesis, MP LSAJC?P BGTGQGML. 5FC QCAMLB NMPRGML MD RFC KGRMRGA NF?QC, A?JJCB AWRMIGLCQGQ, GQ RFC NFWQGA?J QCN?P?RGML MD RFC AWRMNJ?QKGA AMKNMLCLRQ GLRM RFC RUM B?SEFRCP ACJJQ.
3CTGQGR RFC QR?ECQ MD KGRMQGQ ?R RFGQ site (http://openstaxcollege.org/l/Cell_cycle_mito) . K'7>40/3+8/8 (M/948/8) ,?PWMIGLCQGQ, ?JQM ILMUL ?Q mitosis, GQ BGTGBCB GLRM ? QCPGCQ MD NF?QCQqNPMNF?QC, NPMKCR?NF?QC, KCR?NF?QC, ?L?NF?QC, ?LB RCJMNF?QCqRF?R PCQSJR GL RFC BGTGQGML MD RFC ACJJ LSAJCSQ (Figure 10.6). ,?PWMIGLCQGQ GQ ?JQM A?JJCB KGRMQGQ.
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289
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C FMU A?LACP GQ A?SQCB @W SLAMLRPMJJCB ACJJ EPMURF z 6LBCPQR?LB FMU NPMRM-MLAMECLCQ ?PC LMPK?J ACJJ ECLCQ RF?R, UFCL KSR?RCB, @CAMKC MLAMECLCQ z %CQAPG@C FMU RSKMP QSNNPCQQMPQ DSLARGML z &VNJ?GL FMU KSR?LR RSKMP QSNNPCQQMPQ A?SQC A?LACP $?LACP AMKNPGQCQ K?LW BGDDCPCLR BGQC?QCQ A?SQCB @W ? AMKKML KCAF?LGQK: SLAMLRPMJJCB ACJJ EPMURF. %CQNGRC RFC PCBSLB?LAW ?LB MTCPJ?NNGLE JCTCJQ MD ACJJ AWAJC AMLRPMJ, CPPMPQ BM MAASP. 0LC MD RFC APGRGA?J NPMACQQCQ KMLGRMPCB @W RFC ACJJ AWAJC AFCAINMGLR QSPTCGJJ?LAC KCAF?LGQK GQ RFC NPMNCP PCNJGA?RGML MD %/" BSPGLE RFC 4 NF?QC. &TCL UFCL ?JJ MD RFC ACJJ AWAJC AMLRPMJQ ?PC DSJJW DSLARGML?J, ? QK?JJ NCPACLR?EC MD PCNJGA?RGML CPPMPQ (KSR?RGMLQ) UGJJ @C N?QQCB ML RM RFC B?SEFRCP ACJJQ. *D AF?LECQ RM RFC %/" LSAJCMRGBC QCOSCLAC MAASP UGRFGL ? AMBGLE NMPRGML MD ? ECLC ?LB ?PC LMR AMPPCARCB, ? ECLC KSR?RGML PCQSJRQ. "JJ A?LACPQ QR?PR UFCL ? ECLC KSR?RGML EGTCQ PGQC RM ? D?SJRW NPMRCGL RF?R NJ?WQ ? ICW PMJC GL ACJJ PCNPMBSARGML. 5FC AF?LEC GL RFC ACJJ RF?R PCQSJRQ DPMK RFC K?JDMPKCB NPMRCGL K?W @C KGLMP: NCPF?NQ ? QJGEFR BCJ?W GL RFC @GLBGLE MD $BI RM AWAJGL MP ?L 3@ NPMRCGL RF?R BCR?AFCQ DPMK GRQ R?PECR %/" UFGJC QRGJJ NFMQNFMPWJ?RCB. &TCL KGLMP KGQR?ICQ, FMUCTCP, K?W ?JJMU QS@QCOSCLR KGQR?ICQ RM MAASP KMPC PC?BGJW. 0TCP ?LB MTCP, QK?JJ SLAMPPCARCB CPPMPQ ?PC N?QQCB DPMK RFC N?PCLR ACJJ RM RFC B?SEFRCP ACJJQ ?LB ?KNJGDGCB ?Q C?AF ECLCP?RGML NPMBSACQ KMPC LML-DSLARGML?J NPMRCGLQ DPMK SLAMPPCARCB %/" B?K?EC. &TCLRS?JJW, RFC N?AC MD RFC ACJJ AWAJC QNCCBQ SN ?Q RFC CDDCARGTCLCQQ MD RFC AMLRPMJ ?LB PCN?GP KCAF?LGQKQ BCAPC?QCQ. 6LAMLRPMJJCB EPMURF MD RFC KSR?RCB ACJJQ MSRN?ACQ RFC EPMURF MD LMPK?J ACJJQ GL RFC ?PC?, ?LB ? RSKMP (t-MK?u) A?L PCQSJR.
+IFKF-FE:F><E<J
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290
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291
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10.5 R +IFB8IPFK@: C<CC D@M@J@FE

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292
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tCKNCP?RSPC-sCLQGRGTC KSR?LR Z) "0 G?8J< BGQRGLAR DPMK RFC (1 NF?QC MD GLRCPNF?QC; ? ACJJ GL (0 GQ LMR NPCN?PGLE RM BGTGBC "1 G?8J< (?JQM, DGPQR E?N) DGPQR NF?QC MD GLRCPNF?QC ACLRCPCB ML ACJJ EPMURF BSPGLE KGRMQGQ "2 G?8J< (?JQM, QCAMLB E?N) RFGPB NF?QC MD GLRCPNF?QC BSPGLE UFGAF RFC ACJJ SLBCPEMCQ DGL?J NPCN?P?RGMLQ DMP KGRMQGQ >8D<K< F?NJMGB PCNPMBSARGTC ACJJ MP QCV ACJJ (QNCPK, NMJJCL EP?GL, MP CEE) ><E< NFWQGA?J ?LB DSLARGML?J SLGR MD FCPCBGRW, ? QCOSCLAC MD %/" RF?R AMBCQ DMP ? NPMRCGL. ><EFD< RMR?J ECLCRGA GLDMPK?RGML MD ? ACJJ MP MPE?LGQK ?8GCF@; ACJJ, LSAJCSQ, MP MPE?LGQK AMLR?GLGLE MLC QCR MD AFPMKMQMKCQ (5) ?@JKFE< MLC MD QCTCP?J QGKGJ?P, FGEFJW AMLQCPTCB, JMU KMJCASJ?P UCGEFR, @?QGA NPMRCGLQ DMSLB GL RFC AFPMK?RGL MD ?JJ CSI?PWMRGA ACJJQ; ?QQMAG?RCQ UGRF %/" RM DMPK LSAJCMQMKCQ
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295
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
296
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sQ RP?GRQ ?PC BCRCPKGLCB @W RFC ECLCQ GLFCPGRCB DPMK C?AF N?PCLR. %SNJGA?RCB AFPMKMQMKCQ ?PC AMKNMQCB MD RUM QGQRCP AFPMK?RGBQ. $FPMKMQMKCQ ?PC AMKN?ARCB SQGLE ? T?PGCRW MD KCAF?LGQKQ BSPGLE ACPR?GL QR?ECQ MD RFC ACJJ AWAJC. 4CTCP?J AJ?QQCQ MD NPMRCGL ?PC GLTMJTCB GL RFC MPE?LGX?RGML ?LB N?AIGLE MD RFC AFPMKMQMK?J %/" GLRM ? FGEFJW AMLBCLQCB QRPSARSPC. 5FC AMLBCLQGLE AMKNJCV AMKN?ARQ AFPMKMQMKCQ, ?LB RFC PCQSJRGLE AMLBCLQCB QRPSARSPC GQ LCACQQ?PW DMP AFPMKMQMK?J QCEPCE?RGML BSPGLE KGRMQGQ.
10.2 /?< C<CC CP:C<

5FC ACJJ AWAJC GQ ?L MPBCPJW QCOSCLAC MD CTCLRQ. $CJJQ ML RFC N?RF RM ACJJ BGTGQGML NPMACCB RFPMSEF ? QCPGCQ MD NPCAGQCJW RGKCB ?LB A?PCDSJJW PCESJ?RCB QR?ECQ. *L CSI?PWMRCQ, RFC ACJJ AWAJC AMLQGQRQ MD ? JMLE NPCN?P?RMPW NCPGMB, A?JJCB GLRCPNF?QC. *LRCPNF?QC GQ BGTGBCB GLRM (1, 4, ?LB (2 NF?QCQ. 5FC KGRMRGA NF?QC @CEGLQ UGRF I?PWMIGLCQGQ (KGRMQGQ), UFGAF AMLQGQRQ MD DGTC QR?ECQ: NPMNF?QC, NPMKCR?NF?QC, KCR?NF?QC, ?L?NF?QC, ?LB RCJMNF?QC. 5FC DGL?J QR?EC MD RFC KGRMRGA NF?QC GQ AWRMIGLCQGQ, BSPGLE UFGAF RFC AWRMNJ?QKGA AMKNMLCLRQ MD RFC B?SEFRCP ACJJQ ?PC QCN?P?RCB CGRFCP @W ?L ?ARGL PGLE (?LGK?J ACJJQ) MP @W ACJJ NJ?RC DMPK?RGML (NJ?LR ACJJQ).
10.3 CFEKIFC F= K?< C<CC CP:C<

&?AF QRCN MD RFC ACJJ AWAJC GQ KMLGRMPCB @W GLRCPL?J AMLRPMJQ A?JJCB AFCAINMGLRQ. 5FCPC ?PC RFPCC K?HMP AFCAINMGLRQ GL RFC ACJJ AWAJC: MLC LC?P RFC CLB MD (1, ? QCAMLB ?R RFC (2/. RP?LQGRGML, ?LB RFC RFGPB BSPGLE KCR?NF?QC. 1MQGRGTC PCESJ?RMP KMJCASJCQ ?JJMU RFC ACJJ AWAJC RM ?BT?LAC RM RFC LCVR QR?EC. /CE?RGTC PCESJ?RMP KMJCASJCQ KMLGRMP ACJJSJ?P AMLBGRGMLQ ?LB A?L F?JR RFC AWAJC SLRGJ QNCAGDGA PCOSGPCKCLRQ ?PC KCR.
10.4 C8E:<I 8E; K?< C<CC CP:C<

$?LACP GQ RFC PCQSJR MD SLAFCAICB ACJJ BGTGQGML A?SQCB @W ? @PC?IBMUL MD RFC KCAF?LGQKQ RF?R PCESJ?RC RFC ACJJ AWAJC. 5FC JMQQ MD AMLRPMJ @CEGLQ UGRF ? AF?LEC GL RFC %/" QCOSCLAC MD ? ECLC RF?R AMBCQ DMP MLC MD RFC PCESJ?RMPW KMJCASJCQ. '?SJRW GLQRPSARGMLQ JC?B RM ? NPMRCGL RF?R BMCQ LMR DSLARGML ?Q GR QFMSJB. "LW BGQPSNRGML MD RFC KMLGRMPGLE QWQRCK A?L ?JJMU MRFCP KGQR?ICQ RM @C N?QQCB ML RM RFC B?SEFRCP ACJJQ. &?AF QSAACQQGTC ACJJ BGTGQGML UGJJ EGTC PGQC RM B?SEFRCP ACJJQ UGRF CTCL KMPC ?AASKSJ?RCB B?K?EC. &TCLRS?JJW, ?JJ AFCAINMGLRQ @CAMKC LMLDSLARGML?J, ?LB P?NGBJW PCNPMBSAGLE ACJJQ APMUB MSR LMPK?J ACJJQ, PCQSJRGLE GL ? RSKMP MP JCSICKG? (@JMMB A?LACP).
10.5 +IFB8IPFK@: C<CC D@M@J@FE

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1. Figure 10.6 8FGAF MD RFC DMJJMUGLE GQ RFC AMPPCAR MPBCP MD CTCLRQ GL KGRMQGQ >. 4GQRCP AFPMK?RGBQ JGLC SN ?R RFC KCR?NF?QC NJ?RC. 5FC IGLCRMAFMPC @CAMKCQ ?RR?AFCB RM RFC KGRMRGA QNGLBJC. 5FC LSAJCSQ PCDMPKQ ?LB RFC ACJJ
BGTGBCQ. $MFCQGL NPMRCGLQ @PC?I BMUL ?LB RFC QGQRCP AFPMK?RGBQ QCN?P?RC. ?. 5FC IGLCRMAFMPC @CAMKCQ ?RR?AFCB RM RFC KGRMRGA QNGLBJC. $MFCQGL NPMRCGLQ @PC?I BMUL ?LB RFC QGQRCP AFPMK?RGBQ QCN?P?RC. 4GQRCP AFPMK?RGBQ JGLC SN ?R RFC
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297
KCR?NF?QC NJ?RC. 5FC LSAJCSQ PCDMPKQ ?LB RFC ACJJ BGTGBCQ. @. 5FC IGLCRMAFMPC @CAMKCQ ?RR?AFCB RM RFC AMFCQGL NPMRCGLQ. 4GQRCP AFPMK?RGBQ JGLC SN ?R RFC KCR?NF?QC NJ?RC. 5FC IGLCRMAFMPC @PC?IQ BMUL ?LB RFC QGQRCP AFPMK?RGBQ QCN?P?RC. 5FC LSAJCSQ PCDMPKQ ?LB RFC ACJJ BGTGBCQ. A. 5FC IGLCRMAFMPC @CAMKCQ ?RR?AFCB RM RFC KGRMRGA QNGLBJC. 4GQRCP AFPMK?RGBQ JGLC SN ?R RFC KCR?NF?QC NJ?RC. $MFCQGL NPMRCGLQ @PC?I BMUL ?LB RFC QGQRCP AFPMK?RGBQ QCN?P?RC. 5FC LSAJCSQ PCDMPKQ ?LB RFC ACJJ BGTGBCQ.
2. Figure 10.13 3@ ?LB MRFCP NPMRCGLQ RF?R LCE?RGTCJW PCESJ?RC RFC ACJJ AWAJC ?PC QMKCRGKCQ A?JJCB RSKMP QSNNPCQQMPQ. 8FW BM WMS RFGLI RFC L?KC RSKMP QSNNPCQQMP KGEFR @C ?L ?NNPMNPG?RC DMP RFCQC NPMRCGLQ 3. Figure 10.14 )SK?L N?NGJJMK?TGPSQ A?L A?SQC ACPTGA?J A?LACP. 5FC TGPSQ CLAMBCQ &6, ? NPMRCGL RF?R @GLBQ N53. #?QCB ML RFGQ D?AR ?LB UF?R WMS ILMU ?@MSR N53, UF?R CDDCAR BM WMS RFGLI &6 @GLBGLE F?Q ML N53 ?ARGTGRW >. &6 ?ARGT?RCQ N53 ?. &6 GL?ARGT?RCQ N53 @. &6 KSR?RCQ N53 A. &6 @GLBGLE K?PIQ N53 DMP BCEP?B?RGML
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4. " BGNJMGB ACJJ F?Q>>>>>>> RFC LSK@CP MD AFPMKMQMKCQ ?Q ? F?NJMGB ACJJ. >. MLC-DMSPRF ?. F?JD @. RUGAC A. DMSP RGKCQ 5. "L MPE?LGQKs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
6. 5FC DGPQR JCTCJ MD %/" MPE?LGX?RGML GL ? CSI?PWMRGA ACJJ GQ K?GLR?GLCB @W UFGAF KMJCASJC >. ?. @. A. AMFCQGL AMLBCLQGL AFPMK?RGL FGQRMLC
7. *BCLRGA?J AMNGCQ MD AFPMK?RGL FCJB RMECRFCP @W AMFCQGL ?R RFC ACLRPMKCPC ?PC A?JJCB >>>>>. >. ?. @. A. FGQRMLCQ. LSAJCMQMKCQ. AFPMK?RGL. QGQRCP AFPMK?RGBQ.
14. 5FC AFPMKMQMKCQ @CAMKC TGQG@JC SLBCP ? JGEFR KGAPMQAMNC BSPGLE UFGAF QR?EC MD KGRMQGQ >. ?. @. A. NPMNF?QC NPMKCR?NF?QC KCR?NF?QC ?L?NF?QC
8. $FPMKMQMKCQ ?PC BSNJGA?RCB BSPGLE UF?R QR?EC MD RFC ACJJ AWAJC >. (1 NF?QC ?. 4 NF?QC @. NPMNF?QC A. NPMKCR?NF?QC 9. 8FGAF MD RFC DMJJMUGLE CTCLRQ BMCQ LMR MAASP BSPGLE QMKC QR?ECQ MD GLRCPNF?QC >. %/" BSNJGA?RGML ?. MPE?LCJJC BSNJGA?RGML @. GLAPC?QC GL ACJJ QGXC A. QCN?P?RGML MD QGQRCP AFPMK?RGBQ 10. 5FC KGRMRGA QNGLBJCQ ?PGQC DPMK UFGAF ACJJ QRPSARSPC >. ACLRPMKCPC
15. 5FC DSQGLE MD (MJEG TCQGAJCQ ?R RFC KCR?NF?QC NJ?RC MD BGTGBGLE NJ?LR ACJJQ DMPKQ UF?R QRPSARSPC >. ?. @. A. ACJJ NJ?RC ?ARGL PGLE AJC?T?EC DSPPMU KGRMRGA QNGLBJC
16. "R UFGAF MD RFC ACJJ AWAJC AFCAINMGLRQ BM CVRCPL?J DMPACQ F?TC RFC EPC?RCQR GLDJSCLAC >. (1 AFCAINMGLR
298
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NMJCQ MD RFC ACJJ RM DMPK RFC KCR?NF?QC NJ?RC. #CA?SQC RFCPC GQ ?L COS?J AF?LAC RF?R ? KGAPMRS@SJC DG@CP UGJJ CLAMSLRCP ? K?RCPL?JJW MP N?RCPL?JJW GLFCPGRCB AFPMKMQMKC, RFC ?PP?LECKCLR MD RFC RCRP?BQ ?R RFC KCR?NF?QC NJ?RC GQ P?LBMK. "LW K?RCPL?JJW GLFCPGRCB AFPMKMQMKC K?W D?AC CGRFCP NMJC. "LW N?RCPL?JJW GLFCPGRCB AFPMKMQMKC K?W ?JQM D?AC CGRFCP NMJC. 5FC MPGCLR?RGML MD C?AF RCRP?B GQ GLBCNCLBCLR MD RFC MPGCLR?RGML MD RFC MRFCP 22 RCRP?BQ. 5FGQ CTCLRqRFC P?LBMK (MP GLBCNCLBCLR) ?QQMPRKCLR MD FMKMJMEMSQ AFPMKMQMKCQ ?R RFC KCR?NF?QC NJ?RCqGQ RFC QCAMLB KCAF?LGQK RF?R GLRPMBSACQ T?PG?RGML GLRM RFC E?KCRCQ MP QNMPCQ. *L C?AF ACJJ RF?R SLBCPEMCQ KCGMQGQ, RFC ?PP?LECKCLR MD RFC RCRP?BQ GQ BGDDCPCLR. 5FC LSK@CP MD T?PG?RGMLQ GQ BCNCLBCLR ML RFC LSK@CP MD AFPMKMQMKCQ K?IGLE SN ? QCR. 5FCPC ?PC RUM NMQQG@GJGRGCQ DMP MPGCLR?RGML ?R RFC KCR?NF?QC NJ?RC; RFC NMQQG@JC LSK@CP MD ?JGELKCLRQ RFCPCDMPC COS?JQ 25, UFCPC 5 GQ RFC LSK@CP MD AFPMKMQMKCQ NCP QCR. )SK?LQ F?TC 23 AFPMKMQMKC N?GPQ, UFGAF PCQSJRQ GL MTCP CGEFR KGJJGML (223) NMQQG@JC ECLCRGA?JJW-BGQRGLAR E?KCRCQ. 5FGQ LSK@CP BMCQ LMR GLAJSBC RFC T?PG?@GJGRW RF?R U?Q NPCTGMSQJW APC?RCB GL RFC QGQRCP AFPMK?RGBQ @W APMQQMTCP. (GTCL RFCQC RUM KCAF?LGQKQ, GR GQ FGEFJW SLJGICJW RF?R ?LW RUM F?NJMGB ACJJQ PCQSJRGLE DPMK KCGMQGQ UGJJ F?TC RFC Q?KC ECLCRGA AMKNMQGRGML ( Figure 11.4). 5M QSKK?PGXC RFC ECLCRGA AMLQCOSCLACQ MD KCGMQGQ *, RFC K?RCPL?J ?LB N?RCPL?J ECLCQ ?PC PCAMK@GLCB @W APMQQMTCP CTCLRQ RF?R MAASP @CRUCCL C?AF FMKMJMEMSQ N?GP BSPGLE NPMNF?QC *. *L ?BBGRGML, RFC P?LBMK ?QQMPRKCLR MD RCRP?BQ ML RFC KCR?NF?QC NJ?RC NPMBSACQ ? SLGOSC AMK@GL?RGML MD K?RCPL?J ?LB N?RCPL?J AFPMKMQMKCQ RF?R UGJJ K?IC RFCGP U?W GLRM RFC E?KCRCQ.
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MD QCEPCE?RGML GLBGA?RCQ RF?R RFC E?KCRCQ DMP RFC EPCCL/UPGLIJCB NJ?LR ?JJ ?PC @9, ?LB RFC E?KCRCQ DMP RFC WCJJMU/PMSLB NJ?LR ?PC ?JJ &R. 5FCPCDMPC, RFC '1 ECLCP?RGML MD MDDQNPGLE ?JJ ?PC &@R9 (Figure 12.16).
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.CLBCJsQ UMPI QSEECQRCB RF?R RP?GRQ ?PC GLFCPGRCB GLBCNCLBCLRJW MD C?AF MRFCP. .MPE?L GBCLRGDGCB ? 1:1 AMPPCQNMLBCLAC @CRUCCL ? QCEPCE?RGLE RP?GR ?LB RFC 9 AFPMKMQMKC, QSEECQRGLE RF?R RFC P?LBMK QCEPCE?RGML MD AFPMKMQMKCQ U?Q RFC NFWQGA?J @?QGQ MD .CLBCJsQ KMBCJ. 5FGQ ?JQM BCKMLQRP?RCB RF?R JGLICB ECLCQ BGQPSNR .CLBCJsQ NPCBGARCB MSRAMKCQ. 5FC D?AR RF?R C?AF AFPMKMQMKC A?L A?PPW K?LW JGLICB ECLCQ CVNJ?GLQ FMU GLBGTGBS?JQ A?L F?TC K?LW KMPC RP?GRQ RF?L RFCW F?TC AFPMKMQMKCQ. )MUCTCP, M@QCPT?RGMLQ @W PCQC?PAFCPQ GL .MPE?LsQ J?@MP?RMPW QSEECQRCB RF?R ?JJCJCQ NMQGRGMLCB ML RFC Q?KC AFPMKMQMKC UCPC LMR ?JU?WQ GLFCPGRCB RMECRFCP. %SPGLE KCGMQGQ, JGLICB ECLCQ QMKCFMU @CA?KC SLJGLICB. H42414-4:8 +)42(/3'9/43 *L 1909, 'P?LQ +?LQQCL M@QCPTCB AFG?QK?R?qRFC NMGLR ?R UFGAF AFPMK?RGBQ ?PC GL AMLR?AR UGRF C?AF MRFCP ?LB K?W CVAF?LEC QCEKCLRQqNPGMP RM RFC DGPQR BGTGQGML MD KCGMQGQ. )C QSEECQRCB RF?R ?JJCJCQ @CAMKC SLJGLICB ?LB AFPMKMQMKCQ NFWQGA?JJW CVAF?LEC QCEKCLRQ. "Q AFPMKMQMKCQ AMLBCLQCB ?LB N?GPCB UGRF RFCGP FMKMJMEQ, RFCW ?NNC?PCB RM GLRCP?AR ?R BGQRGLAR NMGLRQ. +?LQQCL QSEECQRCB RF?R RFCQC NMGLRQ AMPPCQNMLBCB RM PCEGMLQ GL UFGAF AFPMKMQMKC QCEKCLRQ UCPC CVAF?LECB. *R GQ LMU ILMUL RF?R RFC N?GPGLE ?LB GLRCP?ARGML @CRUCCL FMKMJMEMSQ AFPMKMQMKCQ, ILMUL ?Q QWL?NQGQ, BMCQ KMPC RF?L QGKNJW MPE?LGXC RFC FMKMJMEQ DMP KGEP?RGML RM QCN?P?RC B?SEFRCP ACJJQ. 8FCL QWL?NQCB, FMKMJMEMSQ AFPMKMQMKCQ SLBCPEM PCAGNPMA?J NFWQGA?J CVAF?LECQ ?R RFCGP ?PKQ GL ? NPMACQQ A?JJCB homologous recombination, MP KMPC QGKNJW, tAPMQQGLE MTCP.u 5M @CRRCP SLBCPQR?LB RFC RWNC MD CVNCPGKCLR?J PCQSJRQ RF?R PCQC?PAFCPQ UCPC M@R?GLGLE ?R RFGQ RGKC, AMLQGBCP ? FCRCPMXWEMSQ GLBGTGBS?J RF?R GLFCPGRCB BMKGL?LR K?RCPL?J ?JJCJCQ DMP RUM ECLCQ ML RFC Q?KC AFPMKMQMKC (QSAF ?Q AB) ?LB RUM PCACQQGTC N?RCPL?J ?JJCJCQ DMP RFMQC Q?KC ECLCQ (QSAF ?Q ()). *D RFC ECLCQ ?PC JGLICB, MLC UMSJB CVNCAR RFGQ GLBGTGBS?J RM NPMBSAC E?KCRCQ RF?R ?PC CGRFCP AB MP () UGRF ? 1:1 P?RGM. *D RFC ECLCQ ?PC SLJGLICB, RFC GLBGTGBS?J QFMSJB NPMBSAC AB, A), (B, ?LB () E?KCRCQ UGRF COS?J DPCOSCLAGCQ, ?AAMPBGLE RM RFC .CLBCJG?L AMLACNR MD GLBCNCLBCLR ?QQMPRKCLR. #CA?SQC RFCW AMPPCQNMLB RM LCU ?JJCJC AMK@GL?RGMLQ, RFC ECLMRWNCQ "@ ?LB ?# ?PC nonparental types RF?R PCQSJR DPMK FMKMJMEMSQ PCAMK@GL?RGML BSPGLE KCGMQGQ. Parental types ?PC NPMECLW RF?R CVFG@GR RFC Q?KC ?JJCJGA AMK@GL?RGML ?Q RFCGP N?PCLRQ. .MPE?L ?LB FGQ AMJJC?ESCQ, FMUCTCP, DMSLB RF?R UFCL QSAF FCRCPMXWEMSQ GLBGTGBS?JQ UCPC RCQR APMQQCB RM ? FMKMXWEMSQ PCACQQGTC N?PCLR (A(B) ` (())), @MRF N?PCLR?J ?LB LMLN?PCLR?J A?QCQ MAASPPCB. 'MP CV?KNJC, 950 MDDQNPGLE KGEFR @C PCAMTCPCB RF?R UCPC CGRFCP A(B) MP (()), @SR 50 MDDQNPGLE UMSJB ?JQM @C M@R?GLCB RF?R UCPC CGRFCP A()) MP ((B). 5FCQC PCQSJRQ QSEECQRCB RF?R JGLI?EC MAASPPCB KMQR MDRCL, @SR ? QGELGDGA?LR KGLMPGRW MD MDDQNPGLE UCPC RFC NPMBSARQ MD PCAMK@GL?RGML.
356
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t?JJ-LGEFRCPu RM K?RFCK?RGA?JJW CJSAGB?RC RFC NPM@JCK MD JGLI?EC ?LB PCAMK@GL?RGML. *L 1913, "JDPCB 4RSPRCT?LR, ? QRSBCLR GL .MPE?LsQ J?@MP?RMPW, E?RFCPCB PCQSJRQ DPMK PCQC?PAFCPQ GL RFC J?@MP?RMPW, ?LB RMMI RFCK FMKC MLC LGEFR RM KSJJ RFCK MTCP. #W RFC LCVR KMPLGLE, FC F?B APC?RCB RFC DGPQR tAFPMKMQMKC K?N,u ? JGLC?P PCNPCQCLR?RGML MD ECLC MPBCP ?LB PCJ?RGTC BGQR?LAC ML ? AFPMKMQMKC (Figure 13.4).
#(!04%2 13 X -/$%2. 5.$%234!.$).'3 /& ).(%2)4!.#%
357
!@>LI< 13.4 4EFP DBKBQF@ J>M LOABOP D96:67/03( DBKBP LK QEB ?>PFP LC OB@LJ?FK>QFLK COBNRBK@V.
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igure 13.4, @W SQGLE PCAMK@GL?RGML DPCOSCLAW RM NPCBGAR ECLCRGA BGQR?LAC, RFC PCJ?RGTC MPBCP MD ECLCQ ML AFPMKMQMKC 2 AMSJB @C GLDCPPCB. 5FC T?JSCQ QFMUL PCNPCQCLR K?N BGQR?LACQ GL ACLRGKMPE?LQ (A.), UFGAF AMPPCQNMLB RM PCAMK@GL?RGML DPCOSCLAGCQ (GL NCPACLR). 5FCPCDMPC, RFC ECLCQ DMP @MBW AMJMP ?LB UGLE QGXC UCPC 65.5 y 48.5 = 17 A. ?N?PR, GLBGA?RGLE RF?R RFC K?RCPL?J ?LB N?RCPL?J ?JJCJCQ DMP RFCQC ECLCQ PCAMK@GLC GL 17 NCPACLR MD MDDQNPGLE, ML ?TCP?EC. 5M AMLQRPSAR ? AFPMKMQMKC K?N, 4RSPRCT?LR ?QQSKCB RF?R ECLCQ UCPC MPBCPCB QCPG?JJW ML RFPC?BJGIC AFPMKMQMKCQ. )C ?JQM ?QQSKCB RF?R RFC GLAGBCLAC MD PCAMK@GL?RGML @CRUCCL RUM FMKMJMEMSQ AFPMKMQMKCQ AMSJB MAASP UGRF COS?J JGICJGFMMB ?LWUFCPC ?JMLE RFC JCLERF MD RFC AFPMKMQMKC. 0NCP?RGLE SLBCP RFCQC ?QQSKNRGMLQ, 4RSPRCT?LR NMQRSJ?RCB RF?R ?JJCJCQ RF?R UCPC D?P ?N?PR ML ? AFPMKMQMKC UCPC KMPC JGICJW RM BGQQMAG?RC BSPGLE KCGMQGQ QGKNJW @CA?SQC RFCPC U?Q ? J?PECP PCEGML MTCP UFGAF PCAMK@GL?RGML AMSJB MAASP. $MLTCPQCJW, ?JJCJCQ RF?R UCPC AJMQC RM C?AF MRFCP ML RFC AFPMKMQMKC UCPC JGICJW RM @C GLFCPGRCB RMECRFCP. 5FC ?TCP?EC LSK@CP MD APMQQMTCPQ @CRUCCL RUM ?JJCJCQqRF?R GQ, RFCGP recombination frequencyqAMPPCJ?RCB UGRF RFCGP ECLCRGA BGQR?LAC DPMK C?AF MRFCP, PCJ?RGTC RM RFC JMA?RGMLQ MD MRFCP ECLCQ ML RF?R AFPMKMQMKC. $MLQGBCPGLE RFC CV?KNJC APMQQ @CRUCCL A(B) ?LB (()) ?@MTC, RFC DPCOSCLAW MD PCAMK@GL?RGML AMSJB @C A?JASJ?RCB ?Q 50/1000 = 0.05. 5F?R GQ, RFC JGICJGFMMB MD ? APMQQMTCP @CRUCCL ECLCQ A/( ?LB B/) U?Q 0.05, MP 5 NCPACLR. 4SAF ? PCQSJR UMSJB GLBGA?RC RF?R RFC ECLCQ UCPC BCDGLGRGTCJW JGLICB, @SR RF?R RFCW UCPC D?P CLMSEF ?N?PR DMP APMQQMTCPQ RM MAA?QGML?JJW MAASP. 4RSPRCT?LR BGTGBCB FGQ ECLCRGA K?N GLRM K?N SLGRQ, MP centimorgans (cM), GL UFGAF ? PCAMK@GL?RGML DPCOSCLAW MD 0.01 AMPPCQNMLBQ RM 1 A.. #W PCNPCQCLRGLE ?JJCJCQ GL ? JGLC?P K?N, 4RSPRCT?LR QSEECQRCB RF?R ECLCQ A?L P?LEC DPMK @CGLE NCPDCARJW JGLICB (PCAMK@GL?RGML DPCOSCLAW = 0) RM @CGLE NCPDCARJW SLJGLICB (PCAMK@GL?RGML DPCOSCLAW = 0.5) UFCL ECLCQ ?PC ML BGDDCPCLR AFPMKMQMKCQ MP ECLCQ ?PC QCN?P?RCB TCPW D?P ?N?PR ML RFC Q?KC AFPMKMQMKC. 1CPDCARJW SLJGLICB ECLCQ AMPPCQNMLB RM RFC DPCOSCLAGCQ NPCBGARCB @W .CLBCJ RM ?QQMPR GLBCNCLBCLRJW GL ? BGFW@PGB APMQQ. " PCAMK@GL?RGML DPCOSCLAW MD 0.5 GLBGA?RCQ RF?R 50 NCPACLR MD MDDQNPGLE ?PC PCAMK@GL?LRQ ?LB RFC MRFCP 50 NCPACLR ?PC N?PCLR?J RWNCQ. 5F?R GQ, CTCPW RWNC MD ?JJCJC AMK@GL?RGML GQ PCNPCQCLRCB UGRF COS?J DPCOSCLAW. 5FGQ PCNPCQCLR?RGML ?JJMUCB 4RSPRCT?LR RM ?BBGRGTCJW A?JASJ?RC BGQR?LACQ @CRUCCL QCTCP?J ECLCQ ML RFC Q?KC AFPMKMQMKC. )MUCTCP, ?Q RFC ECLCRGA BGQR?LACQ ?NNPM?AFCB 0.50, FGQ NPCBGARGMLQ @CA?KC JCQQ ?AASP?RC @CA?SQC GR U?Q LMR AJC?P UFCRFCP RFC ECLCQ UCPC TCPW D?P ?N?PR ML RFC Q?KC AFPMKMQMKC MP ML BGDDCPCLR AFPMKMQMKCQ. *L 1931, #?P@?P? .A$JGLRMAI ?LB )?PPGCR $PCGEFRML BCKMLQRP?RCB RFC APMQQMTCP MD FMKMJMEMSQ AFPMKMQMKCQ GL AMPL NJ?LRQ. 8CCIQ J?RCP, FMKMJMEMSQ PCAMK@GL?RGML GL D96:67/03( U?Q BCKMLQRP?RCB KGAPMQAMNGA?JJW @W $SPR 4RCPL. 4RCPL M@QCPTCB QCTCP?J 9-JGLICB NFCLMRWNCQ RF?R UCPC ?QQMAG?RCB UGRF ?
358
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QRPSARSP?JJW SLSQS?J ?LB BGQQGKGJ?P 9 AFPMKMQMKC N?GP GL UFGAF MLC 9 U?Q KGQQGLE ? QK?JJ RCPKGL?J QCEKCLR, ?LB RFC MRFCP 9 U?Q DSQCB RM ? NGCAC MD RFC : AFPMKMQMKC. #W APMQQGLE DJGCQ, M@QCPTGLE RFCGP MDDQNPGLE, ?LB RFCL TGQS?JGXGLE RFC MDDQNPGLEs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
3CTGCU 4RSPRCT?LRsQ NPMACQQ RM APC?RC ? ECLCRGA K?N ML RFC @?QGQ MD PCAMK@GL?RGML DPCOSCLAGCQ here (http://openstaxcollege.org/l/gene_crossover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
13.2 R C?IFDFJFD8C B8J@J F= $E?<I@K<; D@JFI;<IJ

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C FMU ? I?PWMEP?K GQ APC?RCB z &VNJ?GL FMU LMLBGQHSLARGML JC?BQ RM BGQMPBCPQ GL AFPMKMQMKC LSK@CP z $MKN?PC BGQMPBCPQ A?SQCB @W ?LCSNJMGBW z %CQAPG@C FMU CPPMPQ GL AFPMKMQMKC QRPSARSPC MAASP RFPMSEF GLTCPQGMLQ ?LB RP?LQJMA?RGMLQ *LFCPGRCB BGQMPBCPQ A?L ?PGQC UFCL AFPMKMQMKCQ @CF?TC ?@LMPK?JJW BSPGLE KCGMQGQ. $FPMKMQMKC BGQMPBCPQ A?L @C BGTGBCB GLRM RUM A?RCEMPGCQ: ?@LMPK?JGRGCQ GL AFPMKMQMKC LSK@CP ?LB AFPMKMQMK?J QRPSARSP?J PC?PP?LECKCLRQ. #CA?SQC CTCL QK?JJ QCEKCLRQ MD AFPMKMQMKCQ A?L QN?L K?LW ECLCQ, AFPMKMQMK?J BGQMPBCPQ ?PC AF?P?ARCPGQRGA?JJW BP?K?RGA ?LB MDRCL D?R?J.
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5FC GQMJ?RGML ?LB KGAPMQAMNGA M@QCPT?RGML MD AFPMKMQMKCQ DMPKQ RFC @?QGQ MD AWRMECLCRGAQ ?LB GQ RFC NPGK?PW KCRFMB @W UFGAF AJGLGAG?LQ BCRCAR AFPMKMQMK?J ?@LMPK?JGRGCQ GL FSK?LQ. " karyotype GQ RFC LSK@CP ?LB ?NNC?P?LAC MD AFPMKMQMKCQ, ?LB GLAJSBCQ RFCGP JCLERF, @?LBGLE N?RRCPL, ?LB ACLRPMKCPC NMQGRGML. 5M M@R?GL ? TGCU MD ?L GLBGTGBS?JsQ I?PWMRWNC, AWRMJMEGQRQ NFMRMEP?NF RFC AFPMKMQMKCQ ?LB RFCL ASR ?LB N?QRC C?AF AFPMKMQMKC GLRM ? AF?PR, MP karyogram, ?JQM ILMUL ?Q ?L GBCMEP?K (Figure 13.5).
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359
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*L ? EGTCL QNCAGCQ, AFPMKMQMKCQ A?L @C GBCLRGDGCB @W RFCGP LSK@CP, QGXC, ACLRPMKCPC NMQGRGML, ?LB @?LBGLE N?RRCPL. *L ? FSK?L I?PWMRWNC, autosomes MP t@MBW AFPMKMQMKCQu (?JJ MD RFC LMLpQCV AFPMKMQMKCQ) ?PC ECLCP?JJW MPE?LGXCB GL ?NNPMVGK?RC MPBCP MD QGXC DPMK J?PECQR (AFPMKMQMKC 1) RM QK?JJCQR (AFPMKMQMKC 22). 5FC 9 ?LB : AFPMKMQMKCQ ?PC LMR ?SRMQMKCQ. )MUCTCP, AFPMKMQMKC 21 GQ ?ARS?JJW QFMPRCP RF?L AFPMKMQMKC 22. 5FGQ U?Q BGQAMTCPCB ?DRCP RFC L?KGLE MD %MUL QWLBPMKC ?Q RPGQMKW 21, PCDJCARGLE FMU RFGQ BGQC?QC PCQSJRQ DPMK NMQQCQQGLE MLC CVRP? AFPMKMQMKC 21 (RFPCC RMR?J). /MR U?LRGLE RM AF?LEC RFC L?KC MD RFGQ GKNMPR?LR BGQC?QC, AFPMKMQMKC 21 PCR?GLCB GRQ LSK@CPGLE, BCQNGRC BCQAPG@GLE RFC QFMPRCQR QCR MD AFPMKMQMKCQ. 5FC AFPMKMQMKC t?PKQu NPMHCARGLE DPMK CGRFCP CLB MD RFC ACLRPMKCPC K?W @C BCQGEL?RCB ?Q QFMPR MP JMLE, BCNCLBGLE ML RFCGP PCJ?RGTC JCLERFQ. 5FC QFMPR ?PK GQ ?@@PCTG?RCB 7 (DMP tNCRGRCu), UFCPC?Q RFC JMLE ?PK GQ ?@@PCTG?RCB 8 (@CA?SQC GR DMJJMUQ tNu ?JNF?@CRGA?JJW). &?AF ?PK GQ DSPRFCP QS@BGTGBCB ?LB BCLMRCB @W ? LSK@CP. 6QGLE RFGQ L?KGLE QWQRCK, JMA?RGMLQ ML AFPMKMQMKCQ A?L @C BCQAPG@CB AMLQGQRCLRJW GL RFC QAGCLRGDGA JGRCP?RSPC.
360
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!@>LI< 13.12 7EBK LKB @EOLJLPLJB RKABODLBP >K FKSBOPFLK ?RQ QEB LQEBO ALBP KLQ, LKB @EOLJLPLJB JRPQ CLOJ >K FKSBOQBA ILLM QL OBQ>FK MLFKQ-CLO-MLFKQ FKQBO>@QFLK AROFKD PVK>MPFP. 4EFP FKSBOPFLK M>FOFKD FP BPPBKQF>I QL J>FKQ>FKFKD DBKB >IFDKJBKQ AROFKD JBFLPFP >KA QL >IILT CLO OB@LJ?FK>QFLK.
366
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uROCK1 >KA " P14u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translocation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igure 13.13).
!@>LI< 13.13 ! OB@FMOL@>I QO>KPIL@>QFLK L@@ROP TEBK > PBDJBKQ LC $.! FP QO>KPCBOOBA COLJ LKB @EOLJLPLJB QL >KLQEBO, KLKELJLILDLRP @EOLJLPLJB. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .>QFLK>I (RJ>K 'BKLJB 2BPB>O@E/53!)
1. 6FLI>FKB 'LFAQP BQ >I., x3BDJBKQ>I ARMIF@>QFLK >PPL@F>QBA TFQE QEB ERJ>K-PMB@FCF@ FKSBOPFLK LC @EOLJLPLJB 18: > CROQEBO BU>JMIB LC QEB FJM>@Q LC PBDJBKQ>I ARMIF@>QFLKP LK H>OVLQVMB >KA DBKLJB BSLIRQFLK FK MOFJ>QBP,y H<4(5 G,5,;0*:. 115 (2004):116-122
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5FC $FPMKMQMK?J 5FCMPW MD GLFCPGR?LAC, NPMNMQCB @W 4SRRML ?LB #MTCPG, QR?RCQ RF?R AFPMKMQMKCQ ?PC RFC TCFGAJCQ MD ECLCRGA FCPCBGRW. /CGRFCP .CLBCJG?L ECLCRGAQ LMP ECLC JGLI?EC GQ NCPDCARJW ?AASP?RC; GLQRC?B, AFPMKMQMKC @CF?TGMP GLTMJTCQ QCEPCE?RGML, GLBCNCLBCLR ?QQMPRKCLR, ?LB MAA?QGML?JJW, JGLI?EC. 4RSPRCT?LR BCTGQCB ? KCRFMB RM ?QQCQQ PCAMK@GL?RGML DPCOSCLAW ?LB GLDCP RFC PCJ?RGTC NMQGRGMLQ ?LB BGQR?LACQ MD JGLICB ECLCQ ML ? AFPMKMQMKC ML RFC @?QGQ MD RFC ?TCP?EC LSK@CP MD APMQQMTCPQ GL RFC GLRCPTCLGLE PCEGML @CRUCCL RFC ECLCQ. 4RSPRCT?LR AMPPCARJW NPCQSKCB RF?R ECLCQ ?PC ?PP?LECB GL QCPG?J
368
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MPBCP ML AFPMKMQMKCQ ?LB RF?R PCAMK@GL?RGML @CRUCCL FMKMJMEQ A?L MAASP ?LWUFCPC ML ? AFPMKMQMKC UGRF COS?J JGICJGFMMB. 8FCPC?Q JGLI?EC A?SQCQ ?JJCJCQ ML RFC Q?KC AFPMKMQMKC RM @C GLFCPGRCB RMECRFCP, FMKMJMEMSQ PCAMK@GL?RGML @G?QCQ ?JJCJCQ RMU?PB ?L GLFCPGR?LAC N?RRCPL MD GLBCNCLBCLR ?QQMPRKCLR.
13.2 C?IFDFJFD8C B8J@J F= $E?<I@K<; D@JFI;<IJ

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1. Figure 13.3 *L ? RCQR APMQQ DMP RUM AF?P?ARCPGQRGAQ QSAF ?Q RFC MLC QFMUL FCPC, A?L RFC NPCBGARCB DPCOSCLAW MD PCAMK@GL?LR MDDQNPGLE @C 60 NCPACLR 8FW MP UFW LMR 2. Figure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igure 13.6 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR LMLBGQHSLARGML GQ RPSC >. /MLBGQHSLARGML MLJW PCQSJRQ GL E?KCRCQ UGRF L+1 MP Lp1 AFPMKMQMKCQ. ?. /MLBGQHSLARGML MAASPPGLE BSPGLE KCGMQGQ ** PCQSJRQ GL 50 NCPACLR LMPK?J E?KCRCQ. @. /MLBGQHSLARGML BSPGLE KCGMQGQ * PCQSJRQ GL 50 NCPACLR LMPK?J E?KCRCQ. A. /MLBGQHSLARGML ?JU?WQ PCQSJRQ GL DMSP BGDDCPCLR IGLBQ MD E?KCRCQ.
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4. 9-JGLICB PCACQQGTC RP?GRQ GL FSK?LQ (MP GL D96:67/03() ?PC M@QCPTCB >>>>>>>>. >. GL KMPC K?JCQ RF?L DCK?JCQ ?. GL KMPC DCK?JCQ RF?L K?JCQ @. GL K?JCQ ?LB DCK?JCQ COS?JJW A. GL BGDDCPCLR BGQRPG@SRGMLQ BCNCLBGLE ML RFC RP?GR 5. 5FC DGPQR QSEECQRGML RF?R AFPMKMQMKCQ K?W NFWQGA?JJW CVAF?LEC QCEKCLRQ A?KC DPMK RFC KGAPMQAMNGA GBCLRGDGA?RGML MD >>>>>>>>. >. QWL?NQGQ ?. QGQRCP AFPMK?RGBQ @. AFG?QK?R? A. ?JJCJCQ 6. 8FGAF PCAMK@GL?RGML DPCOSCLAW AMPPCQNMLBQ RM GLBCNCLBCLR ?QQMPRKCLR ?LB RFC ?@QCLAC MD JGLI?EC >. 0 ?. 0.25 @. 0.50 A. 0.75 7. 8FGAF PCAMK@GL?RGML DPCOSCLAW AMPPCQNMLBQ RM NCPDCAR JGLI?EC ?LB TGMJ?RCQ RFC J?U MD GLBCNCLBCLR ?QQMPRKCLR >. 0 ?. 0.25 @. 0.50 A. 0.75 8. 8FGAF MD RFC DMJJMUGLE AMBCQ BCQAPG@CQ NMQGRGML 12 ML RFC JMLE ?PK MD AFPMKMQMKC 13 >. ?. @. A. 13N12 13O12 12N13 12O13
9. *L ?EPGASJRSPC, NMJWNJMGB APMNQ (JGIC AMDDCC, QRP?U@CPPGCQ, MP @?L?L?Q) RCLB RM NPMBSAC >>>>>>>>. >. KMPC SLGDMPKGRW
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14. &VNJ?GL FMU RFC $FPMKMQMK?J 5FCMPW MD *LFCPGR?LAC FCJNCB RM ?BT?LAC MSP SLBCPQR?LBGLE MD ECLCRGAQ. 15. 6QGLE BG?EP?KQ, GJJSQRP?RC FMU LMLBGQHSLARGML A?L PCQSJR GL ?L ?LCSNJMGB XWEMRC.
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL RFC NPMACQQ MD %/" PCNJGA?RGML GL NPMI?PWMRCQ z %GQASQQ RFC PMJC MD BGDDCPCLR CLXWKCQ ?LB NPMRCGLQ GL QSNNMPRGLE RFGQ NPMACQQ %/" PCNJGA?RGML F?Q @CCL CVRPCKCJW UCJJ QRSBGCB GL NPMI?PWMRCQ NPGK?PGJW @CA?SQC MD RFC QK?JJ QGXC MD RFC ECLMKC ?LB RFC KSR?LRQ RF?R ?PC ?T?GJ?@JC. E. *630 F?Q 4.6 KGJJGML @?QC N?GPQ GL ? QGLEJC AGPASJ?P AFPMKMQMKC ?LB ?JJ MD GR ECRQ PCNJGA?RCB GL ?NNPMVGK?RCJW 42 KGLSRCQ, QR?PRGLE DPMK ? QGLEJC MPGEGL MD PCNJGA?RGML ?LB NPMACCBGLE ?PMSLB RFC AGPAJC GL @MRF BGPCARGMLQ. 5FGQ KC?LQ RF?R ?NNPMVGK?RCJW 1000 LSAJCMRGBCQ ?PC ?BBCB NCP QCAMLB. 5FC NPMACQQ GQ OSGRC P?NGB ?LB MAASPQ UGRFMSR K?LW KGQR?ICQ. %/" PCNJGA?RGML CKNJMWQ ? J?PEC LSK@CP MD NPMRCGLQ ?LB CLXWKCQ, C?AF MD UFGAF NJ?WQ ? APGRGA?J PMJC BSPGLE RFC NPMACQQ. 0LC MD RFC ICW NJ?WCPQ GQ RFC CLXWKC %/" NMJWKCP?QC, ?JQM ILMUL ?Q %/" NMJ, UFGAF ?BBQ LSAJCMRGBCQ MLC @W MLC RM RFC EPMUGLE %/" AF?GL RF?R ?PC AMKNJCKCLR?PW RM RFC RCKNJ?RC QRP?LB. 5FC ?BBGRGML MD LSAJCMRGBCQ PCOSGPCQ CLCPEW; RFGQ CLCPEW GQ M@R?GLCB DPMK RFC LSAJCMRGBCQ RF?R F?TC RFPCC NFMQNF?RCQ ?RR?AFCB RM RFCK, QGKGJ?P RM "51 UFGAF F?Q RFPCC NFMQNF?RC EPMSNQ ?RR?AFCB. 8FCL RFC @MLB @CRUCCL RFC NFMQNF?RCQ GQ @PMICL, RFC CLCPEW PCJC?QCB GQ SQCB RM DMPK RFC NFMQNFMBGCQRCP @MLB @CRUCCL RFC GLAMKGLE LSAJCMRGBC ?LB RFC EPMUGLE AF?GL. *L NPMI?PWMRCQ, RFPCC K?GL RWNCQ MD NMJWKCP?QCQ ?PC ILMUL: %/" NMJ *, %/" NMJ **, ?LB %/" NMJ ***. *R GQ LMU ILMUL RF?R %/" NMJ *** GQ RFC CLXWKC PCOSGPCB DMP %/" QWLRFCQGQ; %/" NMJ * ?LB %/" NMJ ** ?PC NPGK?PGJW PCOSGPCB DMP PCN?GP. )MU BMCQ RFC PCNJGA?RGML K?AFGLCPW ILMU UFCPC RM @CEGL *R RSPLQ MSR RF?R RFCPC ?PC QNCAGDGA LSAJCMRGBC QCOSCLACQ A?JJCB MPGEGLQ MD PCNJGA?RGML UFCPC PCNJGA?RGML @CEGLQ. *L E. *630, UFGAF F?Q ? QGLEJC MPGEGL MD PCNJGA?RGML ML GRQ MLC AFPMKMQMKC (?Q BM KMQR NPMI?PWMRCQ), GR GQ ?NNPMVGK?RCJW 245 @?QC N?GPQ JMLE ?LB GQ PGAF GL "5 QCOSCLACQ. 5FC MPGEGL MD PCNJGA?RGML GQ PCAMELGXCB @W ACPR?GL NPMRCGLQ RF?R @GLB RM RFGQ QGRC. "L CLXWKC A?JJCB helicase SLUGLBQ RFC %/" @W @PC?IGLE RFC FWBPMECL @MLBQ @CRUCCL RFC LGRPMECLMSQ @?QC N?GPQ. "51 FWBPMJWQGQ GQ PCOSGPCB DMP RFGQ NPMACQQ. "Q RFC %/" MNCLQ SN, :-QF?NCB QRPSARSPCQ A?JJCB replication forks ?PC DMPKCB. 5UM PCNJGA?RGML DMPIQ ?PC DMPKCB ?R RFC MPGEGL MD PCNJGA?RGML ?LB RFCQC ECR CVRCLBCB @G- BGPCARGML?JJW ?Q PCNJGA?RGML NPMACCBQ. Single-strand binding
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RFC DMPK?RGML MD NFMQNFMBGCQRCP JGLI?EC @CRUCCL RFC 3'-0) CLB MD MLC LSAJCMRGBC ?LB RFC 5' NFMQNF?RC CLB MD RFC MRFCP DP?EKCLR. 0LAC RFC AFPMKMQMKC F?Q @CCL AMKNJCRCJW PCNJGA?RCB, RFC RUM %/" AMNGCQ KMTC GLRM RUM BGDDCPCLR ACJJQ BSPGLE ACJJ BGTGQGML. 5FC NPMACQQ MD %/" PCNJGA?RGML A?L @C QSKK?PGXCB ?Q DMJJMUQ: 1. %/" SLUGLBQ ?R RFC MPGEGL MD PCNJGA?RGML. 2. )CJGA?QC MNCLQ SN RFC %/"-DMPKGLE PCNJGA?RGML DMPIQ; RFCQC ?PC CVRCLBCB @GBGPCARGML?JJW. 3. 4GLEJC-QRP?LB @GLBGLE NPMRCGLQ AM?R RFC %/" ?PMSLB RFC PCNJGA?RGML DMPI RM NPCTCLR PCUGLBGLE MD RFC %/". 4. 5MNMGQMKCP?QC @GLBQ ?R RFC PCEGML ?FC?B MD RFC PCNJGA?RGML DMPI RM NPCTCLR QSNCPAMGJGLE. 5. 1PGK?QC QWLRFCQGXCQ 3/" NPGKCPQ AMKNJCKCLR?PW RM RFC %/" QRP?LB. 6. %/" NMJWKCP?QC QR?PRQ ?BBGLE LSAJCMRGBCQ RM RFC 3'-0) CLB MD RFC NPGKCP. 7. &JMLE?RGML MD @MRF RFC J?EEGLE ?LB RFC JC?BGLE QRP?LB AMLRGLSCQ. 8. 3/" NPGKCPQ ?PC PCKMTCB @W CVMLSAJC?QC ?ARGTGRW. 9. (?NQ ?PC DGJJCB @W %/" NMJ @W ?BBGLE B/51Q. 10. 5FC E?N @CRUCCL RFC RUM %/" DP?EKCLRQ GQ QC?JCB @W %/" JGE?QC, UFGAF FCJNQ GL RFC DMPK?RGML MD NFMQNFMBGCQRCP @MLBQ. Table 14.1 QSKK?PGXCQ RFC CLXWKCQ GLTMJTCB GL NPMI?PWMRGA %/" PCNJGA?RGML ?LB RFC DSLARGMLQ MD C?AF.
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3CTGCU RFC DSJJ NPMACQQ MD %/" PCNJGA?RGML here (http://openstaxcollege.org/l/replication_DNA) .
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14.5 R D)A -<GC@:8K@FE @E LB8IPFK<J

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %GQASQQ RFC QGKGJ?PGRGCQ ?LB BGDDCPCLACQ @CRUCCL %/" PCNJGA?RGML GL CSI?PWMRCQ ?LB NPMI?PWMRCQ z 4R?RC RFC PMJC MD RCJMKCP?QC GL %/" PCNJGA?RGML &SI?PWMRGA ECLMKCQ ?PC KSAF KMPC AMKNJCV ?LB J?PECP GL QGXC RF?L NPMI?PWMRGA ECLMKCQ. 5FC FSK?L ECLMKC F?Q RFPCC @GJJGML @?QC N?GPQ NCP F?NJMGB QCR MD AFPMKMQMKCQ, ?LB 6 @GJJGML @?QC N?GPQ ?PC PCNJGA?RCB BSPGLE RFC 4 NF?QC MD RFC ACJJ AWAJC. 5FCPC ?PC KSJRGNJC MPGEGLQ MD PCNJGA?RGML ML RFC CSI?PWMRGA AFPMKMQMKC; FSK?LQ A?L F?TC SN RM 100,000 MPGEGLQ MD PCNJGA?RGML. 5FC P?RC MD PCNJGA?RGML GQ ?NNPMVGK?RCJW 100 LSAJCMRGBCQ NCP QCAMLB, KSAF QJMUCP RF?L NPMI?PWMRGA PCNJGA?RGML. *L WC?QR, UFGAF GQ ? CSI?PWMRC, QNCAG?J QCOSCLACQ ILMUL ?Q "SRMLMKMSQJW 3CNJGA?RGLE 4COSCLACQ ("34) ?PC DMSLB ML RFC AFPMKMQMKCQ. 5FCQC ?PC COSGT?JCLR RM RFC MPGEGL MD PCNJGA?RGML GL E. *630. 5FC LSK@CP MD %/" NMJWKCP?QCQ GL CSI?PWMRCQ GQ KSAF KMPC RF?L NPMI?PWMRCQ: 14 ?PC ILMUL, MD UFGAF DGTC ?PC ILMUL RM F?TC K?HMP PMJCQ BSPGLE PCNJGA?RGML ?LB F?TC @CCL UCJJ QRSBGCB. 5FCW ?PC ILMUL ?Q NMJ C, NMJ D, NMJ E, NMJ F, ?LB NMJ G. 5FC CQQCLRG?J QRCNQ MD PCNJGA?RGML ?PC RFC Q?KC ?Q GL NPMI?PWMRCQ. #CDMPC PCNJGA?RGML A?L QR?PR, RFC %/" F?Q RM @C K?BC ?T?GJ?@JC ?Q RCKNJ?RC. &SI?PWMRGA %/" GQ @MSLB RM @?QGA NPMRCGLQ ILMUL ?Q FGQRMLCQ RM DMPK QRPSARSPCQ A?JJCB LSAJCMQMKCQ. 5FC AFPMK?RGL (RFC AMKNJCV @CRUCCL %/" ?LB NPMRCGLQ) K?W SLBCPEM QMKC AFCKGA?J KMBGDGA?RGMLQ, QM RF?R RFC %/" K?W @C ?@JC RM QJGBC MDD RFC NPMRCGLQ MP @C ?AACQQG@JC RM RFC CLXWKCQ MD RFC %/" PCNJGA?RGML K?AFGLCPW. "R RFC MPGEGL MD PCNJGA?RGML, ? NPCPCNJGA?RGML AMKNJCV GQ K?BC UGRF MRFCP GLGRG?RMP NPMRCGLQ. 0RFCP NPMRCGLQ ?PC RFCL PCAPSGRCB RM QR?PR RFC PCNJGA?RGML NPMACQQ (Table 14.2). " FCJGA?QC SQGLE RFC CLCPEW DPMK "51 FWBPMJWQGQ MNCLQ SN RFC %/" FCJGV. 3CNJGA?RGML DMPIQ ?PC DMPKCB ?R C?AF PCNJGA?RGML MPGEGL ?Q RFC %/" SLUGLBQ. 5FC MNCLGLE MD RFC BMS@JC FCJGV A?SQCQ MTCPUGLBGLE, MP QSNCPAMGJGLE, GL RFC %/" ?FC?B MD RFC PCNJGA?RGML DMPI. 5FCQC ?PC PCQMJTCB UGRF RFC ?ARGML MD RMNMGQMKCP?QCQ. 1PGKCPQ ?PC DMPKCB @W RFC CLXWKC NPGK?QC, ?LB SQGLE RFC NPGKCP, %/" NMJ A?L QR?PR QWLRFCQGQ. 8FGJC RFC JC?BGLE QRP?LB GQ AMLRGLSMSQJW QWLRFCQGXCB @W RFC CLXWKC NMJ F, RFC J?EEGLE QRP?LB GQ QWLRFCQGXCB @W NMJ G. " QJGBGLE AJ?KN NPMRCGL ILMUL ?Q 1$/" (1PMJGDCP?RGLE $CJJ /SAJC?P "LRGECL) FMJBQ RFC %/" NMJ GL NJ?AC QM RF?R GR BMCQ LMR QJGBC MDD RFC %/". 3/?QC ) PCKMTCQ RFC 3/" NPGKCP, UFGAF GQ RFCL PCNJ?ACB UGRF %/" LSAJCMRGBCQ. 5FC 0I?X?IG DP?EKCLRQ GL RFC J?EEGLE QRP?LB ?PC HMGLCB RMECRFCP ?DRCP RFC PCNJ?ACKCLR MD RFC 3/" NPGKCPQ UGRF %/". 5FC E?NQ RF?R PCK?GL ?PC QC?JCB @W %/" JGE?QC, UFGAF DMPKQ RFC NFMQNFMBGCQRCP @MLB.
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2. +?QICJGMDD CR ?J., t5CJMKCP?QC PC?ARGT?RGML PCTCPQCQ RGQQSC BCECLCP?RGML GL ?ECB RCJMKCP?QC-BCDGAGCLR KGAC,u N(;<9, 469 (2011): 102-7.
388
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %GQASQQ RFC BGDDCPCLR RWNCQ MD KSR?RGMLQ GL %/" z &VNJ?GL %/" PCN?GP KCAF?LGQKQ %/" PCNJGA?RGML GQ ? FGEFJW ?AASP?RC NPMACQQ, @SR KGQR?ICQ A?L MAA?QGML?JJW MAASP, QSAF ?Q ? %/" NMJWKCP?QC GLQCPRGLE ? UPMLE @?QC. 6LAMPPCARCB KGQR?ICQ K?W QMKCRGKCQ JC?B RM QCPGMSQ AMLQCOSCLACQ, QSAF ?Q A?LACP. 3CN?GP KCAF?LGQKQ AMPPCAR RFC KGQR?ICQ. *L P?PC A?QCQ, KGQR?ICQ ?PC LMR AMPPCARCB, JC?BGLE RM KSR?RGMLQ; GL MRFCP A?QCQ, PCN?GP CLXWKCQ ?PC RFCKQCJTCQ KSR?RCB MP BCDCARGTC. .MQR MD RFC KGQR?ICQ BSPGLE %/" PCNJGA?RGML ?PC NPMKNRJW AMPPCARCB @W %/" NMJWKCP?QC @W NPMMDPC?BGLE RFC @?QC RF?R F?Q @CCL HSQR ?BBCB (Figure 14.17). *L proofreading, RFC %/" NMJ PC?BQ RFC LCUJW ?BBCB @?QC @CDMPC ?BBGLE RFC LCVR MLC, QM ? AMPPCARGML A?L @C K?BC. 5FC NMJWKCP?QC AFCAIQ UFCRFCP RFC LCUJW ?BBCB @?QC F?Q N?GPCB AMPPCARJW UGRF RFC @?QC GL RFC RCKNJ?RC QRP?LB. *D GR GQ RFC PGEFR @?QC, RFC LCVR LSAJCMRGBC GQ ?BBCB. *D ?L GLAMPPCAR @?QC F?Q @CCL ?BBCB, RFC CLXWKC K?ICQ ? ASR ?R RFC NFMQNFMBGCQRCP @MLB ?LB PCJC?QCQ RFC UPMLE LSAJCMRGBC. 5FGQ GQ NCPDMPKCB @W RFC CVMLSAJC?QC ?ARGML MD %/" NMJ ***. 0LAC RFC GLAMPPCAR LSAJCMRGBC F?Q @CCL PCKMTCB, ? LCU MLC UGJJ @C ?BBCB ?E?GL.
!@>LI< 14.17 0OLLCOB>AFKD ?V $.! MLIVJBO>PB @LOOB@QP BOOLOP AROFKD OBMIF@>QFLK.
4MKC CPPMPQ ?PC LMR AMPPCARCB BSPGLE PCNJGA?RGML, @SR ?PC GLQRC?B AMPPCARCB ?DRCP PCNJGA?RGML GQ AMKNJCRCB; RFGQ RWNC MD PCN?GP GQ ILMUL ?Q mismatch repair (Figure 14.18). 5FC CLXWKCQ PCAMELGXC RFC GLAMPPCARJW ?BBCB LSAJCMRGBC ?LB CVAGQC GR; RFGQ GQ RFCL PCNJ?ACB @W RFC AMPPCAR @?QC. *D RFGQ PCK?GLQ SLAMPPCARCB, GR K?W JC?B RM KMPC NCPK?LCLR B?K?EC. )MU BM KGQK?RAF PCN?GP CLXWKCQ PCAMELGXC UFGAF MD RFC RUM @?QCQ GQ RFC GLAMPPCAR MLC *L E. *630, ?DRCP PCNJGA?RGML, RFC LGRPMECLMSQ @?QC ?BCLGLC ?AOSGPCQ
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
!@>LI< 14.18 )K JFPJ>Q@E OBM>FO, QEB FK@LOOB@QIV >AABA ?>PB FP ABQB@QBA >CQBO OBMIF@>QFLK. 4EB JFPJ>Q@E OBM>FO MOLQBFKP ABQB@Q QEFP ?>PB >KA OBJLSB FQ COLJ QEB KBTIV PVKQEBPFWBA PQO>KA ?V KR@IB>PB >@QFLK. 4EB D>M FP KLT CFIIBA TFQE QEB @LOOB@QIV M>FOBA ?>PB.
*L ?LMRFCP RWNC MD PCN?GP KCAF?LGQK, nucleotide excision repair, CLXWKCQ PCNJ?AC GLAMPPCAR @?QCQ @W K?IGLE ? ASR ML @MRF RFC 3' ?LB 5' CLBQ MD RFC GLAMPPCAR @?QC (Figure 14.19). 5FC QCEKCLR MD %/" GQ PCKMTCB ?LB PCNJ?ACB UGRF RFC AMPPCARJW N?GPCB LSAJCMRGBCQ @W RFC ?ARGML MD %/" NMJ. 0LAC RFC @?QCQ ?PC DGJJCB GL, RFC PCK?GLGLE E?N GQ QC?JCB UGRF ? NFMQNFMBGCQRCP JGLI?EC A?R?JWXCB @W %/" JGE?QC. 5FGQ PCN?GP KCAF?LGQK GQ MDRCL CKNJMWCB UFCL 67 CVNMQSPC A?SQCQ RFC DMPK?RGML MD NWPGKGBGLC BGKCPQ.
!@>LI< 14.19 .R@IBLQFAB BU@FPFLK OBM>FOP QEVJFKB AFJBOP. 7EBK BUMLPBA QL 56, QEVJFKBP IVFKD >AG>@BKQ QL B>@E LQEBO @>K CLOJ QEVJFKB AFJBOP. )K KLOJ>I @BIIP, QEBV >OB BU@FPBA >KA OBMI>@BA.
" UCJJ-QRSBGCB CV?KNJC MD KGQR?ICQ LMR @CGLE AMPPCARCB GQ QCCL GL NCMNJC QSDDCPGLE DPMK VCPMBCPK? NGEKCLRMQ? (Figure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
390
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!@>LI< 14.20 8BOLABOJ> MFDJBKQLP> FP > @LKAFQFLK FK TEF@E QEVJFKB AFJBOFW>QFLK COLJ BUMLPROB QL 56 FP KLQ OBM>FOBA. %UMLPROB QL PRKIFDEQ OBPRIQP FK PHFK IBPFLKP. (@OBAFQ: *>JBP (>IMBOK BQ >I.)
&PPMPQ BSPGLE %/" PCNJGA?RGML ?PC LMR RFC MLJW PC?QML UFW KSR?RGMLQ ?PGQC GL %/". Mutations, T?PG?RGMLQ GL RFC LSAJCMRGBC QCOSCLAC MD ? ECLMKC, A?L ?JQM MAASP @CA?SQC MD B?K?EC RM %/". 4SAF KSR?RGMLQ K?W @C MD RUM RWNCQ: GLBSACB MP QNMLR?LCMSQ. Induced mutations ?PC RFMQC RF?R PCQSJR DPMK ?L CVNMQSPC RM AFCKGA?JQ, 67 P?WQ, V-P?WQ, MP QMKC MRFCP CLTGPMLKCLR?J ?ECLR. Spontaneous mutations MAASP UGRFMSR ?LW CVNMQSPC RM ?LW CLTGPMLKCLR?J ?ECLR; RFCW ?PC ? PCQSJR MD L?RSP?J PC?ARGMLQ R?IGLE NJ?AC UGRFGL RFC @MBW. .SR?RGMLQ K?W F?TC ? UGBC P?LEC MD CDDCARQ. 4MKC KSR?RGMLQ ?PC LMR CVNPCQQCB; RFCQC ?PC ILMUL ?Q silent mutations. Point mutations ?PC RFMQC KSR?RGMLQ RF?R ?DDCAR ? QGLEJC @?QC N?GP. 5FC KMQR AMKKML LSAJCMRGBC KSR?RGMLQ ?PC QS@QRGRSRGMLQ, GL UFGAF MLC @?QC GQ PCNJ?ACB @W ?LMRFCP. 5FCQC A?L @C MD RUM RWNCQ, CGRFCP RP?LQGRGMLQ MP RP?LQTCPQGMLQ. Transition substitution PCDCPQ RM ? NSPGLC MP NWPGKGBGLC @CGLE PCNJ?ACB @W ? @?QC MD RFC Q?KC IGLB; DMP CV?KNJC, ? NSPGLC QSAF ?Q ?BCLGLC K?W @C PCNJ?ACB @W RFC NSPGLC ES?LGLC. Transversion substitution PCDCPQ RM ? NSPGLC @CGLE PCNJ?ACB @W ? NWPGKGBGLC, MP TGAC TCPQ?; DMP CV?KNJC, AWRMQGLC, ? NWPGKGBGLC, GQ PCNJ?ACB @W ?BCLGLC, ? NSPGLC. .SR?RGMLQ A?L ?JQM @C RFC PCQSJR MD RFC ?BBGRGML MD ? @?QC, ILMUL ?Q ?L GLQCPRGML, MP RFC PCKMT?J MD ? @?QC, ?JQM ILMUL ?Q BCJCRGML. 4MKCRGKCQ ? NGCAC MD %/" DPMK MLC AFPMKMQMKC K?W ECR RP?LQJMA?RCB RM ?LMRFCP AFPMKMQMKC MP RM ?LMRFCP PCEGML MD RFC Q?KC AFPMKMQMKC; RFGQ GQ ?JQM ILMUL ?Q RP?LQJMA?RGML. 5FCQC KSR?RGML RWNCQ ?PC QFMUL GL Figure 14.21.
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391
!@>LI< 14.21 -RQ>QFLKP @>K IB>A QL @E>KDBP FK QEB MOLQBFK PBNRBK@B BK@LABA ?V QEB $.!.
! CO>JBPEFCQ JRQ>QFLK QE>Q OBPRIQP FK QEB FKPBOQFLK LC QEOBB KR@IBLQFABP FP LCQBK IBPP ABIBQBOFLRP QE>K > JRQ>QFLK QE>Q OBPRIQP FK QEB FKPBOQFLK LC LKB KR@IBLQFAB. 7EV .SR?RGMLQ GL PCN?GP ECLCQ F?TC @CCL ILMUL RM A?SQC A?LACP. .?LW KSR?RCB PCN?GP ECLCQ F?TC @CCL GKNJGA?RCB GL ACPR?GL DMPKQ MD N?LAPC?RGA A?LACP, AMJML A?LACP, ?LB AMJMPCAR?J A?LACP. .SR?RGMLQ A?L ?DDCAR CGRFCP QMK?RGA ACJJQ MP ECPK ACJJQ. *D K?LW KSR?RGMLQ ?AASKSJ?RC GL ? QMK?RGA ACJJ, RFCW K?W JC?B RM NPM@JCKQ QSAF ?Q RFC SLAMLRPMJJCB ACJJ BGTGQGML M@QCPTCB GL A?LACP. *D ? KSR?RGML R?ICQ NJ?AC GL ECPK ACJJQ, RFC KSR?RGML UGJJ @C N?QQCB ML RM RFC LCVR ECLCP?RGML, ?Q GL RFC A?QC MD FCKMNFGJG? ?LB VCPMBCPK? NGEKCLRMQ?.
392
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KI8EJM<IJ@FE JL9JK@KLK@FE UFCL ? NSPGLC GQ PCNJ?ACB @W ? NWPGKGBGLC MP ? NWPGKGBGLC GQ PCNJ?ACB @W ? NSPGLC
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14.1 #@JKFI@:8C B8J@J F= (F;<IE 0E;<IJK8E;@E>
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
14.2 D)A .KIL:KLI< 8E; .<HL<E:@E>

5FC ASPPCLRJW ?AACNRCB KMBCJ MD RFC BMS@JC-FCJGV QRPSARSPC MD %/" U?Q NPMNMQCB @W 8?RQML ?LB $PGAI. 4MKC MD RFC Q?JGCLR DC?RSPCQ ?PC RF?R RFC RUM QRP?LBQ RF?R K?IC SN RFC BMS@JC FCJGV ?PC AMKNJCKCLR?PW ?LB ?LRG-N?P?JJCJ GL L?RSPC. %CMVWPG@MQC QSE?PQ ?LB NFMQNF?RCQ DMPK RFC @?AI@MLC MD RFC QRPSARSPC, ?LB RFC LGRPMECLMSQ @?QCQ ?PC QR?AICB GLQGBC. 5FC BG?KCRCP MD RFC BMS@JC FCJGV, 2 LK, GQ SLGDMPK RFPMSEFMSR. " NSPGLC ?JU?WQ N?GPQ UGRF ? NWPGKGBGLC; " N?GPQ UGRF 5, ?LB ( N?GPQ UGRF $. 0LC RSPL MD RFC FCJGV F?Q RCL @?QC N?GPQ. %SPGLE ACJJ BGTGQGML, C?AF B?SEFRCP ACJJ PCACGTCQ ? AMNW MD RFC %/" @W ? NPMACQQ ILMUL ?Q %/" PCNJGA?RGML. 1PMI?PWMRCQ ?PC KSAF QGKNJCP RF?L CSI?PWMRCQ GL K?LW MD RFCGP DC?RSPCQ. .MQR NPMI?PWMRCQ AMLR?GL ? QGLEJC, AGPASJ?P AFPMKMQMKC. *L ECLCP?J, CSI?PWMRGA AFPMKMQMKCQ AMLR?GL ? JGLC?P %/" KMJCASJC N?AI?ECB GLRM LSAJCMQMKCQ, ?LB F?TC RUM BGQRGLAR PCEGMLQ RF?R A?L @C BGQRGLESGQFCB @W QR?GLGLE, PCDJCARGLE BGDDCPCLR QR?RCQ MD N?AI?EGLE ?LB AMKN?ARGML.
14.3 B8J@:J F= D)A -<GC@:8K@FE

5FC KMBCJ DMP %/" PCNJGA?RGML QSEECQRQ RF?R RFC RUM QRP?LBQ MD RFC BMS@JC FCJGV QCN?P?RC BSPGLE PCNJGA?RGML, ?LB C?AF QRP?LB QCPTCQ ?Q ? RCKNJ?RC DPMK UFGAF RFC LCU AMKNJCKCLR?PW QRP?LB GQ AMNGCB. *L AMLQCPT?RGTC PCNJGA?RGML, RFC N?PCLR?J %/" GQ AMLQCPTCB, ?LB RFC B?SEFRCP %/" GQ LCUJW QWLRFCQGXCB. 5FC QCKG-AMLQCPT?RGTC KCRFMB QSEECQRQ RF?R C?AF MD RFC RUM N?PCLR?J %/" QRP?LBQ ?ARQ ?Q RCKNJ?RC DMP LCU %/" RM @C QWLRFCQGXCB; ?DRCP PCNJGA?RGML, C?AF BMS@JC-QRP?LBCB %/" GLAJSBCQ MLC N?PCLR?J MP tMJBu QRP?LB ?LB MLC tLCUu QRP?LB. 5FC BGQNCPQGTC KMBC QSEECQRCB RF?R RFC RUM AMNGCQ MD RFC %/" UMSJB F?TC QCEKCLRQ MD N?PCLR?J %/" ?LB LCUJW QWLRFCQGXCB %/".
14.4 D)A -<GC@:8K@FE @E +IFB8IPFK<J

3CNJGA?RGML GL NPMI?PWMRCQ QR?PRQ DPMK ? QCOSCLAC DMSLB ML RFC AFPMKMQMKC A?JJCB RFC MPGEGL MD PCNJGA?RGMLqRFC NMGLR ?R UFGAF RFC %/" MNCLQ SN. )CJGA?QC MNCLQ SN RFC %/" BMS@JC FCJGV, PCQSJRGLE GL RFC DMPK?RGML MD RFC PCNJGA?RGML DMPI. 4GLEJC-QRP?LB @GLBGLE NPMRCGLQ @GLB RM RFC QGLEJC-QRP?LBCB %/" LC?P RFC PCNJGA?RGML DMPI RM ICCN RFC DMPI MNCL. 1PGK?QC QWLRFCQGXCQ ?L 3/" NPGKCP RM GLGRG?RC QWLRFCQGQ @W %/" NMJWKCP?QC, UFGAF A?L ?BB LSAJCMRGBCQ MLJW GL RFC 5' RM 3' BGPCARGML. 0LC QRP?LB GQ QWLRFCQGXCB AMLRGLSMSQJW GL RFC BGPCARGML MD RFC PCNJGA?RGML DMPI; RFGQ GQ A?JJCB RFC JC?BGLE QRP?LB. 5FC MRFCP QRP?LB GQ QWLRFCQGXCB GL ? BGPCARGML ?U?W DPMK RFC PCNJGA?RGML DMPI, GL QFMPR QRPCRAFCQ MD %/" ILMUL ?Q 0I?X?IG DP?EKCLRQ. 5FGQ QRP?LB GQ ILMUL ?Q RFC J?EEGLE QRP?LB. 0LAC PCNJGA?RGML GQ AMKNJCRCB, RFC 3/" NPGKCPQ ?PC PCNJ?ACB @W %/" LSAJCMRGBCQ ?LB RFC %/" GQ QC?JCB UGRF %/" JGE?QC, UFGAF APC?RCQ NFMQNFMBGCQRCP @MLBQ @CRUCCL RFC 3'-0) MD MLC CLB ?LB RFC 5' NFMQNF?RC MD RFC MRFCP QRP?LB.
14.5 D)A -<GC@:8K@FE @E LB8IPFK<J

3CNJGA?RGML GL CSI?PWMRCQ QR?PRQ ?R KSJRGNJC MPGEGLQ MD PCNJGA?RGML. 5FC KCAF?LGQK GQ OSGRC QGKGJ?P RM NPMI?PWMRCQ. " NPGKCP GQ PCOSGPCB RM GLGRG?RC QWLRFCQGQ, UFGAF GQ RFCL CVRCLBCB @W %/" NMJWKCP?QC ?Q GR ?BBQ LSAJCMRGBCQ MLC @W MLC RM RFC EPMUGLE AF?GL. 5FC JC?BGLE QRP?LB GQ QWLRFCQGXCB AMLRGLSMSQJW, UFCPC?Q RFC J?EEGLE QRP?LB GQ QWLRFCQGXCB GL QFMPR QRPCRAFCQ A?JJCB 0I?X?IG DP?EKCLRQ. 5FC 3/" NPGKCPQ ?PC PCNJ?ACB UGRF %/" LSAJCMRGBCQ; RFC %/" PCK?GLQ MLC AMLRGLSMSQ QRP?LB @W JGLIGLE RFC %/" DP?EKCLRQ UGRF %/" JGE?QC. 5FC CLBQ MD RFC AFPMKMQMKCQ NMQC ? NPM@JCK ?Q NMJWKCP?QC GQ SL?@JC RM CVRCLB RFCK UGRFMSR ? NPGKCP. 5CJMKCP?QC, ?L CLXWKC UGRF ?L GL@SGJR 3/" RCKNJ?RC, CVRCLBQ RFC CLBQ @W AMNWGLE RFC 3/" RCKNJ?RC ?LB CVRCLBGLE MLC CLB MD RFC AFPMKMQMKC. %/" NMJWKCP?QC A?L RFCL CVRCLB RFC %/" SQGLE RFC NPGKCP. *L RFGQ U?W, RFC CLBQ MD RFC AFPMKMQMKCQ ?PC NPMRCARCB.
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%/" NMJWKCP?QC A?L K?IC KGQR?ICQ UFGJC ?BBGLE LSAJCMRGBCQ. *R CBGRQ RFC %/" @W NPMMDPC?BGLE CTCPW LCUJW ?BBCB @?QC. *LAMPPCAR @?QCQ ?PC PCKMTCB ?LB PCNJ?ACB @W RFC AMPPCAR @?QC, ?LB RFCL ? LCU @?QC GQ ?BBCB. .MQR KGQR?ICQ ?PC AMPPCARCB BSPGLE PCNJGA?RGML, ?JRFMSEF UFCL RFGQ BMCQ LMR F?NNCL, RFC KGQK?RAF PCN?GP KCAF?LGQK GQ CKNJMWCB. .GQK?RAF PCN?GP CLXWKCQ PCAMELGXC RFC UPMLEJW GLAMPNMP?RCB @?QC ?LB CVAGQC GR DPMK RFC %/", PCNJ?AGLE GR UGRF RFC AMPPCAR @?QC. *L WCR ?LMRFCP RWNC MD PCN?GP, LSAJCMRGBC CVAGQGML PCN?GP, RFC GLAMPPCAR @?QC GQ PCKMTCB ?JMLE UGRF ? DCU @?QCQ ML RFC 5' ?LB 3' CLB, ?LB RFCQC ?PC PCNJ?ACB @W AMNWGLE RFC RCKNJ?RC UGRF RFC FCJN MD %/" NMJWKCP?QC. 5FC CLBQ MD RFC LCUJW QWLRFCQGXCB DP?EKCLR ?PC ?RR?AFCB RM RFC PCQR MD RFC %/" SQGLE %/" JGE?QC, UFGAF APC?RCQ ? NFMQNFMBGCQRCP @MLB. .MQR KGQR?ICQ ?PC AMPPCARCB, ?LB GD RFCW ?PC LMR, RFCW K?W PCQSJR GL ? KSR?RGML BCDGLCB ?Q ? NCPK?LCLR AF?LEC GL RFC %/" QCOSCLAC. .SR?RGMLQ A?L @C MD K?LW RWNCQ, QSAF ?Q QS@QRGRSRGML, BCJCRGML, GLQCPRGML, ?LB RP?LQJMA?RGML. .SR?RGMLQ GL PCN?GP ECLCQ K?W JC?B RM QCPGMSQ AMLQCOSCLACQ QSAF ?Q A?LACP. .SR?RGMLQ A?L @C GLBSACB MP K?W MAASP QNMLR?LCMSQJW.
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1. Figure 14.10 *L CSI?PWMRGA ACJJQ, %/" ?LB 3/" QWLRFCQGQ MAASP GL ? QCN?P?RC AMKN?PRKCLR DPMK NPMRCGL QWLRFCQGQ. *L NPMI?PWMRGA ACJJQ, @MRF NPMACQQCQ MAASP RMECRFCP. 8F?R ?BT?LR?ECQ KGEFR RFCPC @C RM QCN?P?RGLE RFC NPMACQQCQ 8F?R ?BT?LR?ECQ KGEFR RFCPC @C RM F?TGLE RFCK MAASP RMECRFCP 2. Figure 14.14 :MS GQMJ?RC ? ACJJ QRP?GL GL UFGAF RFC HMGLGLE RMECRFCP MD 0I?X?IG DP?EKCLRQ GQ GKN?GPCB ?LB QSQNCAR RF?R ? KSR?RGML F?Q MAASPPCB GL ?L CLXWKC DMSLB ?R RFC PCNJGA?RGML DMPI. 8FGAF CLXWKC GQ KMQR JGICJW RM @C KSR?RCB 3. Figure 14.21 " DP?KCQFGDR KSR?RGML RF?R PCQSJRQ GL RFC GLQCPRGML MD RFPCC LSAJCMRGBCQ GQ MDRCL JCQQ BCJCRCPGMSQ RF?L ? KSR?RGML RF?R PCQSJRQ GL RFC GLQCPRGML MD MLC LSAJCMRGBC. 8FW
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15. 5FC GLGRG?J KCAF?LGQK DMP PCN?GPGLE LSAJCMRGBC CPPMPQ GL %/" GQ >>>>>>>>. >. KGQK?RAF PCN?GP ?. %/" NMJWKCP?QC NPMMDPC?BGLE @. LSAJCMRGBC CVAGQGML PCN?GP A. RFWKGLC BGKCPQ
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16. &VNJ?GL (PGDDGRF'Q RP?LQDMPK?RGML CVNCPGKCLRQ. 8F?R BGB FC AMLAJSBC DPMK RFCK 17. 8FW UCPC P?BGM?ARGTC QSJDSP ?LB NFMQNFMPMSQ SQCB RM J?@CJ @?ARCPGMNF?EC GL )CPQFCW ?LB $F?QC'Q CVNCPGKCLRQ 18. 1PMTGBC ? @PGCD QSKK?PW MD RFC 4?LECP QCOSCLAGLE KCRFMB. 19. %CQAPG@C RFC QRPSARSPC ?LB AMKNJCKCLR?PW @?QC N?GPGLE MD %/". 20. )MU BGB RFC QAGCLRGDGA AMKKSLGRW JC?PL RF?R %/" PCNJGA?RGML R?ICQ NJ?AC GL ? QCKGAMLQCPT?RGTC D?QFGML 21. %/" PCNJGA?RGML GQ @GBGPCARGML?J ?LB BGQAMLRGLSMSQ; CVNJ?GL WMSP SLBCPQR?LBGLE MD RFMQC AMLACNRQ. 22. 8F?R ?PC 0I?X?IG DP?EKCLRQ ?LB FMU RFCW ?PC DMPKCB 23. *D RFC P?RC MD PCNJGA?RGML GL ? N?PRGASJ?P NPMI?PWMRC GQ 900 LSAJCMRGBCQ NCP QCAMLB, FMU JMLE UMSJB GR R?IC 1.2 KGJJGML @?QC N?GP ECLMKCQ RM K?IC RUM AMNGCQ 24. &VNJ?GL RFC CTCLRQ R?IGLE NJ?AC ?R RFC PCNJGA?RGML DMPI. *D RFC ECLC DMP FCJGA?QC GQ KSR?RCB, UF?R N?PR MD PCNJGA?RGML UGJJ @C ?DDCARCB 25. 8F?R GQ RFC PMJC MD ? NPGKCP GL %/" PCNJGA?RGML 8F?R UMSJB F?NNCL GD WMS DMPEMR RM ?BB ? NPGKCP GL ? RS@C AMLR?GLGLE RFC PC?ARGML KGV DMP ? %/" QCOSCLAGLE PC?ARGML 26. )MU BM RFC JGLC?P AFPMKMQMKCQ GL CSI?PWMRCQ CLQSPC RF?R GRQ CLBQ ?PC PCNJGA?RCB AMKNJCRCJW 27. 8F?R GQ RFC AMLQCOSCLAC MD KSR?RGML MD ? KGQK?RAF PCN?GP CLXWKC )MU UGJJ RFGQ ?DDCAR RFC DSLARGML MD ? ECLC
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able 15.1). 5FC P3/" KMJCASJCQ ?PC AMLQGBCPCB QRPSARSP?J 3/"Q @CA?SQC RFCW F?TC ? ACJJSJ?P PMJC @SR ?PC LMR RP?LQJ?RCB GLRM NPMRCGL. 5FC P3/"Q ?PC AMKNMLCLRQ MD RFC PG@MQMKC ?LB ?PC CQQCLRG?J RM RFC NPMACQQ MD RP?LQJ?RGML. 3/" NMJWKCP?QC * QWLRFCQGXCQ ?JJ MD RFC P3/"Q CVACNR DMP RFC 54 P3/" KMJCASJC. 5FC t4u BCQGEL?RGML ?NNJGCQ RM t4TCB@CPEu SLGRQ, ? LML?BBGRGTC T?JSC RF?R AF?P?ARCPGXCQ RFC QNCCB ?R UFGAF ? N?PRGAJC QCBGKCLRQ BSPGLE ACLRPGDSE?RGML.
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406
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407
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408
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RFC K?HMP QF?PC MD CSI?PWMRGA ECLCQ, QM RFGQ QCARGML UGJJ DMASQ ML FMU RFGQ NMJWKCP?QC ?AAMKNJGQFCQ CJMLE?RGML ?LB RCPKGL?RGML. "JRFMSEF RFC CLXWK?RGA NPMACQQ MD CJMLE?RGML GQ CQQCLRG?JJW RFC Q?KC GL CSI?PWMRCQ ?LB NPMI?PWMRCQ, RFC %/" RCKNJ?RC GQ KMPC AMKNJCV. 8FCL CSI?PWMRGA ACJJQ ?PC LMR BGTGBGLE, RFCGP ECLCQ CVGQR ?Q ? BGDDSQC K?QQ MD %/" ?LB NPMRCGLQ A?JJCB AFPMK?RGL. 5FC %/" GQ RGEFRJW N?AI?ECB ?PMSLB AF?PECB FGQRMLC NPMRCGLQ ?R PCNC?RCB GLRCPT?JQ. 5FCQC %/"pFGQRMLC AMKNJCVCQ, AMJJCARGTCJW A?JJCB LSAJCMQMKCQ, ?PC PCESJ?PJW QN?ACB ?LB GLAJSBC 146 LSAJCMRGBCQ MD %/" UMSLB ?PMSLB CGEFR FGQRMLCQ JGIC RFPC?B ?PMSLB ? QNMMJ. 'MP NMJWLSAJCMRGBC QWLRFCQGQ RM MAASP, RFC RP?LQAPGNRGML K?AFGLCPW LCCBQ RM KMTC FGQRMLCQ MSR MD RFC U?W CTCPW RGKC GR CLAMSLRCPQ ? LSAJCMQMKC. 5FGQ GQ ?AAMKNJGQFCB @W ? QNCAG?J NPMRCGL AMKNJCV A?JJCB FACT, UFGAF QR?LBQ DMP tD?AGJGR?RCQ AFPMK?RGL RP?LQAPGNRGML.u 5FGQ AMKNJCV NSJJQ FGQRMLCQ ?U?W DPMK RFC %/" RCKNJ?RC ?Q RFC NMJWKCP?QC KMTCQ ?JMLE GR. 0LAC RFC NPC-K3/" GQ QWLRFCQGXCB, RFC '"$5 AMKNJCV PCNJ?ACQ RFC FGQRMLCQ RM PCAPC?RC RFC LSAJCMQMKCQ. 5FC RCPKGL?RGML MD RP?LQAPGNRGML GQ BGDDCPCLR DMP RFC BGDDCPCLR NMJWKCP?QCQ. 6LJGIC GL NPMI?PWMRCQ, CJMLE?RGML @W 3/" NMJWKCP?QC ** GL CSI?PWMRCQ R?ICQ NJ?AC 1,000p2,000 LSAJCMRGBCQ @CWMLB RFC CLB MD RFC ECLC @CGLE RP?LQAPG@CB. 5FGQ NPC-K3/" R?GJ GQ QS@QCOSCLRJW PCKMTCB @W AJC?T?EC BSPGLE K3/" NPMACQQGLE. 0L RFC MRFCP F?LB, 3/" NMJWKCP?QCQ * ?LB *** PCOSGPC RCPKGL?RGML QGEL?JQ. (CLCQ RP?LQAPG@CB @W 3/" NMJWKCP?QC * AMLR?GL ? QNCAGDGA 18-LSAJCMRGBC QCOSCLAC RF?R GQ PCAMELGXCB @W ? RCPKGL?RGML NPMRCGL. 5FC NPMACQQ MD RCPKGL?RGML GL 3/" NMJWKCP?QC *** GLTMJTCQ ?L K3/" F?GPNGL QGKGJ?P RM PFMGLBCNCLBCLR RCPKGL?RGML MD RP?LQAPGNRGML GL NPMI?PWMRCQ.
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409
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uO>QEBO QE>K MOLQBFKPu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methylguanosine cap GQ ?BBCB RM RFC 5' CLB MD RFC EPMUGLE RP?LQAPGNR @W ? NFMQNF?RC JGLI?EC. 5FGQ KMGCRW (DSLARGML?J EPMSN) NPMRCARQ RFC L?QACLR K3/" DPMK BCEP?B?RGML. *L ?BBGRGML, D?ARMPQ GLTMJTCB GL NPMRCGL QWLRFCQGQ PCAMELGXC RFC A?N RM FCJN GLGRG?RC RP?LQJ?RGML @W PG@MQMKCQ.
410
#(!04%2 15 X '%.%3 !.$ 02/4%).3
3' P41>-A "'/1 0LAC CJMLE?RGML GQ AMKNJCRC, RFC NPC-K3/" GQ AJC?TCB @W ?L CLBMLSAJC?QC @CRUCCL ?L ""6""" AMLQCLQSQ QCOSCLAC ?LB ? (6-PGAF QCOSCLAC, JC?TGLE RFC ""6""" QCOSCLAC ML RFC NPC-K3/". "L CLXWKC A?JJCB NMJW-" NMJWKCP?QC RFCL ?BBQ ? QRPGLE MD ?NNPMVGK?RCJW 200 " PCQGBSCQ, A?JJCB RFC polyA tail. 5FGQ KMBGDGA?RGML DSPRFCP NPMRCARQ RFC NPC-K3/" DPMK BCEP?B?RGML ?LB QGEL?JQ RFC CVNMPR MD RFC ACJJSJ?P D?ARMPQ RF?R RFC RP?LQAPGNR LCCBQ RM RFC AWRMNJ?QK. P7+-2 NA !51/)/3&SI?PWMRGA ECLCQ ?PC AMKNMQCB MD exons, UFGAF AMPPCQNMLB RM NPMRCGL-AMBGLE QCOSCLACQ (,?-ML QGELGDGCQ RF?R RFCW ?PC ,?NPCQQCB), ?LB 05;CPTCLGLE QCOSCLACQ A?JJCB introns (05;-PML BCLMRCQ RFCGP 05;CPTCLGLE PMJC), UFGAF K?W @C GLTMJTCB GL ECLC PCESJ?RGML @SR ?PC PCKMTCB DPMK RFC NPC-K3/" BSPGLE NPMACQQGLE. *LRPML QCOSCLACQ GL K3/" BM LMR CLAMBC DSLARGML?J NPMRCGLQ. 5FC BGQAMTCPW MD GLRPMLQ A?KC ?Q ? QSPNPGQC RM PCQC?PAFCPQ GL RFC 1970Q UFM CVNCARCB RF?R NPCK3/"Q UMSJB QNCAGDW NPMRCGL QCOSCLACQ UGRFMSR DSPRFCP NPMACQQGLE, ?Q RFCW F?B M@QCPTCB GL NPMI?PWMRCQ. 5FC ECLCQ MD FGEFCP CSI?PWMRCQ TCPW MDRCL AMLR?GL MLC MP KMPC GLRPMLQ. 5FCQC PCEGMLQ K?W AMPPCQNMLB RM PCESJ?RMPW QCOSCLACQ; FMUCTCP, RFC @GMJMEGA?J QGELGDGA?LAC MD F?TGLE K?LW GLRPMLQ MP F?TGLE TCPW JMLE GLRPMLQ GL ? ECLC GQ SLAJC?P. *R GQ NMQQG@JC RF?R GLRPMLQ QJMU BMUL ECLC CVNPCQQGML @CA?SQC GR R?ICQ JMLECP RM RP?LQAPG@C NPC-K3/"Q UGRF JMRQ MD GLRPMLQ. "JRCPL?RGTCJW, GLRPMLQ K?W @C LMLDSLARGML?J QCOSCLAC PCKL?LRQ JCDR MTCP DPMK RFC DSQGML MD ?LAGCLR ECLCQ RFPMSEFMSR CTMJSRGML. 5FGQ GQ QSNNMPRCB @W RFC D?AR RF?R QCN?P?RC CVMLQ MDRCL CLAMBC QCN?P?RC NPMRCGL QS@SLGRQ MP BMK?GLQ. 'MP RFC KMQR N?PR, RFC QCOSCLACQ MD GLRPMLQ A?L @C KSR?RCB UGRFMSR SJRGK?RCJW ?DDCARGLE RFC NPMRCGL NPMBSAR. "JJ MD ? NPC-K3/"sQ GLRPMLQ KSQR @C AMKNJCRCJW ?LB NPCAGQCJW PCKMTCB @CDMPC NPMRCGL QWLRFCQGQ. *D RFC NPMACQQ CPPQ @W CTCL ? QGLEJC LSAJCMRGBC, RFC PC?BGLE DP?KC MD RFC PCHMGLCB CVMLQ UMSJB QFGDR, ?LB RFC PCQSJRGLE NPMRCGL UMSJB @C BWQDSLARGML?J. 5FC NPMACQQ MD PCKMTGLE GLRPMLQ ?LB PCAMLLCARGLE CVMLQ GQ A?JJCB splicing (Figure 15.13). *LRPMLQ ?PC PCKMTCB ?LB BCEP?BCB UFGJC RFC NPC-K3/" GQ QRGJJ GL RFC LSAJCSQ. 4NJGAGLE MAASPQ @W ? QCOSCLAC-QNCAGDGA KCAF?LGQK RF?R CLQSPCQ GLRPMLQ UGJJ @C PCKMTCB ?LB CVMLQ PCHMGLCB UGRF RFC ?AASP?AW ?LB NPCAGQGML MD ? QGLEJC LSAJCMRGBC. 5FC QNJGAGLE MD NPC-K3/"Q GQ AMLBSARCB @W AMKNJCVCQ MD NPMRCGLQ ?LB 3/" KMJCASJCQ A?JJCB QNJGACMQMKCQ.
!@>LI< 15.13 0OB-J2.! PMIF@FKD FKSLISBP QEB MOB@FPB OBJLS>I LC FKQOLKP COLJ QEB MOFJ>OV 2.! QO>KP@OFMQ. 4EB PMIF@FKD MOL@BPP FP @>Q>IVWBA ?V MOLQBFK @LJMIBUBP @>IIBA PMIF@BLPLJBP QE>Q >OB @LJMLPBA LC MOLQBFKP >KA 2.! JLIB@RIBP @>IIBA PK2.!P. 3MIF@BLPLJBP OB@LDKFWB PBNRBK@BP >Q QEB 5' >KA 3' BKA LC QEB FKQOLK.
%OOLOP FK PMIF@FKD >OB FJMIF@>QBA FK @>K@BOP >KA LQEBO ERJ>K AFPB>PBP. 7E>Q HFKAP LC JRQ>QFLKP JFDEQ IB>A QL PMIF@FKD BOOLOP 4EFKH LC AFCCBOBKQ MLPPF?IB LRQ@LJBP FC PMIF@FKD BOOLOP L@@RO.
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411
/MRC RF?R KMPC RF?L 70 GLBGTGBS?J GLRPMLQ A?L @C NPCQCLR, ?LB C?AF F?Q RM SLBCPEM RFC NPMACQQ MD QNJGAGLEqGL ?BBGRGML RM 5' A?NNGLE ?LB RFC ?BBGRGML MD ? NMJW-" R?GJqHSQR RM ECLCP?RC ? QGLEJC, RP?LQJ?R?@JC K3/" KMJCASJC.
4CC FMU GLRPMLQ ?PC PCKMTCB BSPGLE 3/" QNJGAGLE at this website (http://openstaxcollege.org/l/ RNA_splicing) .
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p$)3 KMGCRW (KCRFWJ DSLARGML?J EPMSN) GQ ?BBCB DMP QR?@GJGRW. 1PC-R3/" KMJCASJCQ ?JQM SLBCPEM KCRFWJ?RGML. "Q UGRF NPC-K3/"Q, QS@SLGR CVAGQGML MAASPQ GL CSI?PWMRGA NPC-3/"Q BCQRGLCB RM @CAMKC R3/"Q MP P3/"Q. .?RSPC P3/"Q K?IC SN ?NNPMVGK?RCJW 50 NCPACLR MD C?AF PG@MQMKC. 4MKC MD ? PG@MQMKCsQ 3/" KMJCASJCQ ?PC NSPCJW QRPSARSP?J, UFCPC?Q MRFCPQ F?TC A?R?JWRGA MP @GLBGLE ?ARGTGRGCQ. .?RSPC R3/"Q R?IC ML ? RFPCC-BGKCLQGML?J QRPSARSPC RFPMSEF GLRP?KMJCASJ?P FWBPMECL @MLBGLE RM NMQGRGML RFC ?KGLM ?AGB @GLBGLE QGRC ?R MLC CLB ?LB RFC anticodon ?R RFC MRFCP CLB (Figure 15.14). 5FC ?LRGAMBML GQ ? RFPCCLSAJCMRGBC QCOSCLAC GL ? R3/" RF?R GLRCP?ARQ UGRF ?L K3/" AMBML RFPMSEF AMKNJCKCLR?PW @?QC N?GPGLE.
412
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC BGDDCPCLR QRCNQ GL NPMRCGL QWLRFCQGQ z %GQASQQ RFC PMJC MD PG@MQMKCQ GL NPMRCGL QWLRFCQGQ 5FC QWLRFCQGQ MD NPMRCGLQ AMLQSKCQ KMPC MD ? ACJJsQ CLCPEW RF?L ?LW MRFCP KCR?@MJGA NPMACQQ. *L RSPL, NPMRCGLQ ?AAMSLR DMP KMPC K?QQ RF?L ?LW MRFCP AMKNMLCLR MD JGTGLE MPE?LGQKQ (UGRF RFC CVACNRGML MD U?RCP), ?LB NPMRCGLQ NCPDMPK TGPRS?JJW CTCPW DSLARGML MD ? ACJJ. 5FC NPMACQQ MD RP?LQJ?RGML, MP NPMRCGL QWLRFCQGQ, GLTMJTCQ RFC BCAMBGLE MD ?L K3/" KCQQ?EC GLRM ? NMJWNCNRGBC NPMBSAR. "KGLM ?AGBQ ?PC AMT?JCLRJW QRPSLE RMECRFCP @W GLRCPJGLIGLE NCNRGBC @MLBQ GL JCLERFQ P?LEGLE DPMK ?NNPMVGK?RCJW 50 ?KGLM ?AGB PCQGBSCQ RM KMPC RF?L 1,000. &?AF GLBGTGBS?J ?KGLM ?AGB F?Q ?L ?KGLM EPMSN (/)2) ?LB ? A?P@MVWJ ($00)) EPMSN. 1MJWNCNRGBCQ ?PC DMPKCB UFCL RFC ?KGLM EPMSN MD MLC ?KGLM ?AGB DMPKQ ?L ?KGBC (G.C., NCNRGBC) @MLB UGRF RFC A?P@MVWJ EPMSN MD ?LMRFCP ?KGLM ?AGB (Figure 15.15). 5FGQ PC?ARGML GQ A?R?JWXCB @W PG@MQMKCQ ?LB ECLCP?RCQ MLC U?RCP KMJCASJC.
#(!04%2 15 X '%.%3 !.$ 02/4%).3
413
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*L ?BBGRGML RM RFC K3/" RCKNJ?RC, K?LW KMJCASJCQ ?LB K?APMKMJCASJCQ AMLRPG@SRC RM RFC NPMACQQ MD RP?LQJ?RGML. 5FC AMKNMQGRGML MD C?AF AMKNMLCLR K?W T?PW ?APMQQ QNCAGCQ; DMP GLQR?LAC, PG@MQMKCQ K?W AMLQGQR MD BGDDCPCLR LSK@CPQ MD P3/"Q ?LB NMJWNCNRGBCQ BCNCLBGLE ML RFC MPE?LGQK. )MUCTCP, RFC ECLCP?J QRPSARSPCQ ?LB DSLARGMLQ MD RFC NPMRCGL QWLRFCQGQ K?AFGLCPW ?PC AMKN?P?@JC DPMK @?ARCPG? RM FSK?L ACJJQ. 5P?LQJ?RGML PCOSGPCQ RFC GLNSR MD ?L K3/" RCKNJ?RC, PG@MQMKCQ, R3/"Q, ?LB T?PGMSQ CLXWK?RGA D?ARMPQ.
$JGAI RFPMSEF RFC QRCNQ MD RFGQ PBS interactive (http://openstaxcollege.org/l/prokary_protein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polysome. 9 NA8 5FC R3/"Q ?PC QRPSARSP?J 3/" KMJCASJCQ RF?R UCPC RP?LQAPG@CB DPMK ECLCQ @W 3/" NMJWKCP?QC ***. %CNCLBGLE ML RFC QNCAGCQ, 40 RM 60 RWNCQ MD R3/"Q CVGQR GL RFC AWRMNJ?QK. 4CPTGLE ?Q ?B?NRMPQ, QNCAGDGA R3/"Q @GLB RM QCOSCLACQ ML RFC K3/" RCKNJ?RC ?LB ?BB RFC AMPPCQNMLBGLE ?KGLM ?AGB RM RFC
414
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5FC ECLCRGA AMBC PCDCPQ RM RFC %/" ?JNF?@CR (", 5, $, (), RFC 3/" ?JNF?@CR (", 6, $, (), ?LB RFC NMJWNCNRGBC ?JNF?@CR (20 ?KGLM ?AGBQ). 5FC $CLRP?J %MEK? BCQAPG@CQ RFC DJMU MD ECLCRGA GLDMPK?RGML GL RFC ACJJ DPMK ECLCQ RM K3/" RM NPMRCGLQ. (CLCQ ?PC SQCB RM K?IC K3/" @W RFC NPMACQQ MD RP?LQAPGNRGML; K3/" GQ SQCB RM QWLRFCQGXC NPMRCGLQ @W RFC NPMACQQ MD RP?LQJ?RGML. 5FC ECLCRGA AMBC GQ
420
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BCECLCP?RC @CA?SQC 64 RPGNJCR AMBMLQ GL K3/" QNCAGDW MLJW 20 ?KGLM ?AGBQ ?LB RFPCC LMLQCLQC AMBMLQ. "JKMQR CTCPW QNCAGCQ ML RFC NJ?LCR SQCQ RFC Q?KC ECLCRGA AMBC.
15.2 +IFB8IPFK@: /I8EJ:I@GK@FE

*L NPMI?PWMRCQ, K3/" QWLRFCQGQ GQ GLGRG?RCB ?R ? NPMKMRCP QCOSCLAC ML RFC %/" RCKNJ?RC AMKNPGQGLE RUM AMLQCLQSQ QCOSCLACQ RF?R PCAPSGR 3/" NMJWKCP?QC. 5FC NPMI?PWMRGA NMJWKCP?QC AMLQGQRQ MD ? AMPC CLXWKC MD DMSP NPMRCGL QS@SLGRQ ?LB ? H NPMRCGL RF?R ?QQGQRQ MLJW UGRF GLGRG?RGML. &JMLE?RGML QWLRFCQGXCQ K3/" GL RFC 5' RM 3' BGPCARGML ?R ? P?RC MD 40 LSAJCMRGBCQ NCP QCAMLB. 5CPKGL?RGML JG@CP?RCQ RFC K3/" ?LB MAASPQ CGRFCP @W PFM NPMRCGL GLRCP?ARGML MP @W RFC DMPK?RGML MD ?L K3/" F?GPNGL.
15.3 LB8IPFK@: /I8EJ:I@GK@FE

5P?LQAPGNRGML GL CSI?PWMRCQ GLTMJTCQ MLC MD RFPCC RWNCQ MD NMJWKCP?QCQ, BCNCLBGLE ML RFC ECLC @CGLE RP?LQAPG@CB. 3/" NMJWKCP?QC ** RP?LQAPG@CQ ?JJ MD RFC NPMRCGL-AMBGLE ECLCQ, UFCPC?Q 3/" NMJWKCP?QC * RP?LQAPG@CQ P3/" ECLCQ, ?LB 3/" NMJWKCP?QC *** RP?LQAPG@CQ P3/", R3/", ?LB QK?JJ LSAJC?P 3/" ECLCQ. 5FC GLGRG?RGML MD RP?LQAPGNRGML GL CSI?PWMRCQ GLTMJTCQ RFC @GLBGLE MD QCTCP?J RP?LQAPGNRGML D?ARMPQ RM AMKNJCV NPMKMRCP QCOSCLACQ RF?R ?PC SQS?JJW JMA?RCB SNQRPC?K MD RFC ECLC @CGLE AMNGCB. 5FC K3/" GQ QWLRFCQGXCB GL RFC 5' RM 3' BGPCARGML, ?LB RFC '"$5 AMKNJCV KMTCQ ?LB PC?QQCK@JCQ LSAJCMQMKCQ ?Q RFC NMJWKCP?QC N?QQCQ @W. 8FCPC?Q 3/" NMJWKCP?QCQ * ?LB *** RCPKGL?RC RP?LQAPGNRGML @W NPMRCGL- MP 3/" F?GPNGL-BCNCLBCLR KCRFMBQ, 3/" NMJWKCP?QC ** RP?LQAPG@CQ DMP 1,000 MP KMPC LSAJCMRGBCQ @CWMLB RFC ECLC RCKNJ?RC ?LB AJC?TCQ RFC CVACQQ BSPGLE NPC-K3/" NPMACQQGLE.
15.4 -)A +IF:<JJ@E> @E LB8IPFK<J

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
15.5 -@9FJFD<J 8E; +IFK<@E .PEK?<J@J

5FC NJ?WCPQ GL RP?LQJ?RGML GLAJSBC RFC K3/" RCKNJ?RC, PG@MQMKCQ, R3/"Q, ?LB T?PGMSQ CLXWK?RGA D?ARMPQ. 5FC QK?JJ PG@MQMK?J QS@SLGR DMPKQ ML RFC K3/" RCKNJ?RC CGRFCP ?R RFC 4FGLC-%?JE?PLM QCOSCLAC (NPMI?PWMRCQ) MP RFC 5' A?N (CSI?PWMRCQ). 5P?LQJ?RGML @CEGLQ ?R RFC GLGRG?RGLE "6( ML RFC K3/", QNCAGDWGLE KCRFGMLGLC. 5FC DMPK?RGML MD NCNRGBC @MLBQ MAASPQ @CRUCCL QCOSCLRG?J ?KGLM ?AGBQ QNCAGDGCB @W RFC K3/" RCKNJ?RC ?AAMPBGLE RM RFC ECLCRGA AMBC. $F?PECB R3/"Q CLRCP RFC PG@MQMK?J " QGRC, ?LB RFCGP ?KGLM ?AGB @MLBQ UGRF RFC ?KGLM ?AGB ?R RFC 1 QGRC. 5FC CLRGPC K3/" GQ RP?LQJ?RCB GL RFPCC-LSAJCMRGBC tQRCNQu MD RFC PG@MQMKC. 8FCL ? LMLQCLQC AMBML GQ CLAMSLRCPCB, ? PCJC?QC D?ARMP @GLBQ ?LB BGQQMAG?RCQ RFC AMKNMLCLRQ ?LB DPCCQ RFC LCU NPMRCGL. 'MJBGLE MD RFC NPMRCGL MAASPQ BSPGLE ?LB ?DRCP RP?LQJ?RGML.
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1. Figure 15.11 " QAGCLRGQR QNJGACQ ? CSI?PWMRGA NPMKMRCP GL DPMLR MD ? @?ARCPG?J ECLC ?LB GLQCPRQ RFC ECLC GL ? @?ARCPG?J AFPMKMQMKC. 8MSJB WMS CVNCAR RFC @?ARCPG? RM RP?LQAPG@C RFC ECLC 2. Figure 15.13 &PPMPQ GL QNJGAGLE ?PC GKNJGA?RCB GL A?LACPQ ?LB MRFCP FSK?L BGQC?QCQ. 8F?R IGLBQ MD KSR?RGMLQ KGEFR JC?B RM QNJGAGLE CPPMPQ 5FGLI MD BGDDCPCLR NMQQG@JC MSRAMKCQ GD QNJGAGLE CPPMPQ MAASP. 3. Figure 15.16 .?LW ?LRG@GMRGAQ GLFG@GR @?ARCPG?J NPMRCGL QWLRFCQGQ. 'MP CV?KNJC, RCRP?AWAJGLC @JMAIQ RFC " QGRC ML RFC @?ARCPG?J PG@MQMKC, ?LB AFJMP?KNFCLGAMJ @JMAIQ NCNRGBWJ RP?LQDCP. 8F?R QNCAGDGA CDDCAR UMSJB WMS CVNCAR C?AF MD RFCQC ?LRG@GMRGAQ RM F?TC ML NPMRCGL QWLRFCQGQ 5CRP?AWAJGLC UMSJB BGPCARJW ?DDCAR: >. R3/" @GLBGLE RM RFC PG@MQMKC ?. PG@MQMKC ?QQCK@JW @. EPMURF MD RFC NPMRCGL AF?GL $FJMP?KNFCLGAMJ UMSJB BGPCARJW ?DDCAR >. R3/" @GLBGLE RM RFC PG@MQMKC ?. PG@MQMKC ?QQCK@JW @. EPMURF MD RFC NPMRCGL AF?GL
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4. 5FC "6$ ?LB "6" AMBMLQ GL K3/" @MRF QNCAGDW GQMJCSAGLC. 8F?R DC?RSPC MD RFC ECLCRGA AMBC CVNJ?GLQ RFGQ >. AMKNJCKCLR?PGRW ?. LMLQCLQC AMBMLQ @. SLGTCPQ?JGRW A. BCECLCP?AW 5. )MU K?LW LSAJCMRGBCQ ?PC GL 12 K3/" AMBMLQ >. 12 ?. 24 @. 36 A. 48 6. 8FGAF QS@SLGR MD RFC E. *630 NMJWKCP?QC AMLDCPQ QNCAGDGAGRW RM RP?LQAPGNRGML >. C ?. D @. D' A. H 7. 5FC -10 ?LB -35 PCEGMLQ MD NPMI?PWMRGA NPMKMRCPQ ?PC A?JJCB AMLQCLQSQ QCOSCLACQ @CA?SQC >>>>>>>>. >. RFCW ?PC GBCLRGA?J GL ?JJ @?ARCPG?J QNCAGCQ ?. RFCW ?PC QGKGJ?P GL ?JJ @?ARCPG?J QNCAGCQ @. RFCW CVGQR GL ?JJ MPE?LGQKQ A. RFCW F?TC RFC Q?KC DSLARGML GL ?JJ MPE?LGQKQ 8. 8FGAF DC?RSPC MD NPMKMRCPQ A?L @C DMSLB GL @MRF NPMI?PWMRCQ ?LB CSI?PWMRCQ >. ($ @MV ?. 5"5" @MV @. MAR?KCP @MV A. -10 ?LB -35 QCOSCLACQ 9. 8F?R RP?LQAPGNRQ UGJJ @C KMQR ?DDCARCB @W JMU JCTCJQ MD i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14. *K?EGLC GD RFCPC UCPC 200 AMKKMLJW MAASPPGLE ?KGLM ?AGBQ GLQRC?B MD 20. (GTCL UF?R WMS ILMU ?@MSR RFC ECLCRGA AMBC, UF?R UMSJB @C RFC QFMPRCQR NMQQG@JC AMBML JCLERF &VNJ?GL. 15. %GQASQQ FMU BCECLCP?AW MD RFC ECLCRGA AMBC K?ICQ ACJJQ KMPC PM@SQR RM KSR?RGMLQ. 16. *D K3/" GQ AMKNJCKCLR?PW RM RFC %/" RCKNJ?RC QRP?LB ?LB RFC %/" RCKNJ?RC QRP?LB GQ AMKNJCKCLR?PW RM RFC %/" LMLRCKNJ?RC QRP?LB, RFCL UFW ?PC @?QC QCOSCLACQ MD K3/" ?LB RFC %/" LMLRCKNJ?RC QRP?LB LMR GBCLRGA?J $MSJB RFCW CTCP @C 17. *L WMSP MUL UMPBQ, BCQAPG@C RFC BGDDCPCLAC @CRUCCL PFM-BCNCLBCLR ?LB PFM-GLBCNCLBCLR RCPKGL?RGML MD RP?LQAPGNRGML GL NPMI?PWMRCQ. 18. 5P?LQAPG@C ?LB RP?LQJ?RC RFC DMJJMUGLE %/" QCOSCLAC (LMLRCKNJ?RC QRP?LB): 5'"5(($$((55"55""($"-3' 19. &VNJ?GL FMU QGLEJC LSAJCMRGBC AF?LECQ A?L F?TC T?QRJW BGDDCPCLR CDDCARQ ML NPMRCGL DSLARGML.
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&?AF QMK?RGA ACJJ GL RFC @MBW ECLCP?JJW AMLR?GLQ RFC Q?KC %/". " DCU CVACNRGMLQ GLAJSBC PCB @JMMB ACJJQ, UFGAF AMLR?GL LM %/" GL RFCGP K?RSPC QR?RC, ?LB QMKC GKKSLC QWQRCK ACJJQ RF?R PC?PP?LEC RFCGP %/" UFGJC NPMBSAGLE ?LRG@MBGCQ. *L ECLCP?J, FMUCTCP, RFC ECLCQ RF?R BCRCPKGLC UFCRFCP WMS F?TC EPCCL CWCQ, @PMUL F?GP, ?LB FMU D?QR WMS KCR?@MJGXC DMMB ?PC RFC Q?KC GL RFC ACJJQ GL WMSP CWCQ ?LB WMSP JGTCP, CTCL RFMSEF RFCQC MPE?LQ DSLARGML OSGRC BGDDCPCLRJW. *D C?AF ACJJ F?Q RFC Q?KC %/", FMU GQ GR RF?R ACJJQ MP MPE?LQ ?PC BGDDCPCLR 8FW BM ACJJQ GL RFC CWC BGDDCP QM BP?K?RGA?JJW DPMK ACJJQ GL RFC JGTCP 8FCPC?Q C?AF ACJJ QF?PCQ RFC Q?KC ECLMKC ?LB %/" QCOSCLAC, C?AF ACJJ BMCQ LMR RSPL ML, MP CVNPCQQ, RFC Q?KC QCR MD ECLCQ. &?AF ACJJ RWNC LCCBQ ? BGDDCPCLR QCR MD NPMRCGLQ RM NCPDMPK GRQ DSLARGML. 5FCPCDMPC, MLJW ? QK?JJ QS@QCR MD NPMRCGLQ GQ CVNPCQQCB GL ? ACJJ. 'MP RFC NPMRCGLQ RM @C CVNPCQQCB, RFC %/" KSQR @C RP?LQAPG@CB GLRM 3/" ?LB RFC 3/" KSQR @C RP?LQJ?RCB GLRM NPMRCGL. *L ? EGTCL ACJJ RWNC, LMR ?JJ ECLCQ CLAMBCB GL RFC %/" ?PC RP?LQAPG@CB GLRM 3/" MP RP?LQJ?RCB GLRM NPMRCGL @CA?SQC QNCAGDGA ACJJQ GL MSP @MBW F?TC QNCAGDGA DSLARGMLQ. 4NCAG?JGXCB NPMRCGLQ RF?R K?IC SN RFC CWC (GPGQ, JCLQ, ?LB AMPLC?) ?PC MLJW CVNPCQQCB GL RFC CWC, UFCPC?Q RFC QNCAG?JGXCB NPMRCGLQ GL RFC FC?PR (N?ACK?ICP ACJJQ, FC?PR KSQAJC, ?LB T?JTCQ) ?PC MLJW CVNPCQQCB GL RFC FC?PR. "R ?LW EGTCL RGKC, MLJW ? QS@QCR MD ?JJ MD RFC ECLCQ CLAMBCB @W MSP %/" ?PC CVNPCQQCB ?LB RP?LQJ?RCB GLRM NPMRCGLQ. 5FC CVNPCQQGML MD QNCAGDGA ECLCQ GQ ? FGEFJW PCESJ?RCB NPMACQQ UGRF K?LW JCTCJQ ?LB QR?ECQ MD AMLRPMJ. 5FGQ AMKNJCVGRW CLQSPCQ RFC NPMNCP CVNPCQQGML GL RFC NPMNCP ACJJ ?R RFC NPMNCP RGKC.
424
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %GQASQQ UFW CTCPW ACJJ BMCQ LMR CVNPCQQ ?JJ MD GRQ ECLCQ z %CQAPG@C FMU NPMI?PWMRGA ECLC PCESJ?RGML MAASPQ ?R RFC RP?LQAPGNRGML?J JCTCJ z %GQASQQ FMU CSI?PWMRGA ECLC PCESJ?RGML MAASPQ ?R RFC CNGECLCRGA, RP?LQAPGNRGML?J, NMQRRP?LQAPGNRGML?J, RP?LQJ?RGML?J, ?LB NMQR-RP?LQJ?RGML?J JCTCJQ 'MP ? ACJJ RM DSLARGML NPMNCPJW, LCACQQ?PW NPMRCGLQ KSQR @C QWLRFCQGXCB ?R RFC NPMNCP RGKC. "JJ ACJJQ AMLRPMJ MP PCESJ?RC RFC QWLRFCQGQ MD NPMRCGLQ DPMK GLDMPK?RGML CLAMBCB GL RFCGP %/". 5FC NPMACQQ MD RSPLGLE ML ? ECLC RM NPMBSAC 3/" ?LB NPMRCGL GQ A?JJCB gene expression. 8FCRFCP GL ? QGKNJC SLGACJJSJ?P MPE?LGQK MP ? AMKNJCV KSJRG-ACJJSJ?P MPE?LGQK, C?AF ACJJ AMLRPMJQ UFCL ?LB FMU GRQ ECLCQ ?PC CVNPCQQCB. 'MP RFGQ RM MAASP, RFCPC KSQR @C ? KCAF?LGQK RM AMLRPMJ UFCL ? ECLC GQ CVNPCQQCB RM K?IC 3/" ?LB NPMRCGL, FMU KSAF MD RFC NPMRCGL GQ K?BC, ?LB UFCL GR GQ RGKC RM QRMN K?IGLE RF?R NPMRCGL @CA?SQC GR GQ LM JMLECP LCCBCB. 5FC PCESJ?RGML MD ECLC CVNPCQQGML AMLQCPTCQ CLCPEW ?LB QN?AC. *R UMSJB PCOSGPC ? QGELGDGA?LR ?KMSLR MD CLCPEW DMP ?L MPE?LGQK RM CVNPCQQ CTCPW ECLC ?R ?JJ RGKCQ, QM GR GQ KMPC CLCPEW CDDGAGCLR RM RSPL ML RFC ECLCQ MLJW UFCL RFCW ?PC PCOSGPCB. *L ?BBGRGML, MLJW CVNPCQQGLE ? QS@QCR MD ECLCQ GL C?AF ACJJ Q?TCQ QN?AC @CA?SQC %/" KSQR @C SLUMSLB DPMK GRQ RGEFRJW AMGJCB QRPSARSPC RM RP?LQAPG@C ?LB RP?LQJ?RC RFC %/". $CJJQ UMSJB F?TC RM @C CLMPKMSQ GD CTCPW NPMRCGL UCPC CVNPCQQCB GL CTCPW ACJJ ?JJ RFC RGKC. 5FC AMLRPMJ MD ECLC CVNPCQQGML GQ CVRPCKCJW AMKNJCV. .?JDSLARGMLQ GL RFGQ NPMACQQ ?PC BCRPGKCLR?J RM RFC ACJJ ?LB A?L JC?B RM RFC BCTCJMNKCLR MD K?LW BGQC?QCQ, GLAJSBGLE A?LACP.
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5M SLBCPQR?LB FMU ECLC CVNPCQQGML GQ PCESJ?RCB, UC KSQR DGPQR SLBCPQR?LB FMU ? ECLC AMBCQ DMP ? DSLARGML?J NPMRCGL GL ? ACJJ. 5FC NPMACQQ MAASPQ GL @MRF NPMI?PWMRGA ?LB CSI?PWMRGA ACJJQ, HSQR GL QJGEFRJW BGDDCPCLR K?LLCPQ. 1PMI?PWMRGA MPE?LGQKQ ?PC QGLEJC-ACJJCB MPE?LGQKQ RF?R J?AI ? ACJJ LSAJCSQ, ?LB RFCGP %/" RFCPCDMPC DJM?RQ DPCCJW GL RFC ACJJ AWRMNJ?QK. 5M QWLRFCQGXC ? NPMRCGL, RFC NPMACQQCQ MD RP?LQAPGNRGML ?LB RP?LQJ?RGML MAASP ?JKMQR QGKSJR?LCMSQJW. 8FCL RFC PCQSJRGLE NPMRCGL GQ LM JMLECP LCCBCB, RP?LQAPGNRGML QRMNQ. "Q ? PCQSJR, RFC NPGK?PW KCRFMB RM AMLRPMJ UF?R RWNC MD NPMRCGL ?LB FMU KSAF MD C?AF NPMRCGL GQ CVNPCQQCB GL ? NPMI?PWMRGA ACJJ GQ RFC PCESJ?RGML MD %/" RP?LQAPGNRGML. "JJ MD RFC QS@QCOSCLR QRCNQ MAASP ?SRMK?RGA?JJW. 8FCL KMPC NPMRCGL GQ PCOSGPCB, KMPC RP?LQAPGNRGML MAASPQ. 5FCPCDMPC, GL NPMI?PWMRGA ACJJQ, RFC AMLRPMJ MD ECLC CVNPCQQGML GQ KMQRJW ?R RFC RP?LQAPGNRGML?J JCTCJ. &SI?PWMRGA ACJJQ, GL AMLRP?QR, F?TC GLRP?ACJJSJ?P MPE?LCJJCQ RF?R ?BB RM RFCGP AMKNJCVGRW. *L CSI?PWMRGA ACJJQ, RFC %/" GQ AMLR?GLCB GLQGBC RFC ACJJsQ LSAJCSQ ?LB RFCPC GR GQ RP?LQAPG@CB GLRM 3/". 5FC LCUJW QWLRFCQGXCB 3/" GQ RFCL RP?LQNMPRCB MSR MD RFC LSAJCSQ GLRM RFC AWRMNJ?QK, UFCPC PG@MQMKCQ RP?LQJ?RC RFC 3/" GLRM NPMRCGL. 5FC NPMACQQCQ MD RP?LQAPGNRGML ?LB RP?LQJ?RGML ?PC NFWQGA?JJW QCN?P?RCB @W RFC LSAJC?P KCK@P?LC; RP?LQAPGNRGML MAASPQ MLJW UGRFGL RFC LSAJCSQ, ?LB RP?LQJ?RGML MAASPQ MLJW MSRQGBC RFC LSAJCSQ GL RFC AWRMNJ?QK. 5FC PCESJ?RGML MD ECLC CVNPCQQGML A?L MAASP ?R ?JJ QR?ECQ MD RFC NPMACQQ (Figure 16.2). 3CESJ?RGML K?W MAASP UFCL RFC %/" GQ SLAMGJCB ?LB JMMQCLCB DPMK LSAJCMQMKCQ RM @GLB RP?LQAPGNRGML D?ARMPQ ( epigenetic JCTCJ), UFCL RFC 3/" GQ RP?LQAPG@CB (RP?LQAPGNRGML?J JCTCJ), UFCL RFC 3/" GQ NPMACQQCB ?LB CVNMPRCB RM RFC AWRMNJ?QK ?DRCP GR GQ RP?LQAPG@CB ( post-transcriptional JCTCJ), UFCL RFC 3/" GQ RP?LQJ?RCB GLRM NPMRCGL (RP?LQJ?RGML?J JCTCJ), MP ?DRCP RFC NPMRCGL F?Q @CCL K?BC ( posttranslational JCTCJ).
#(!04%2 16 X '%.% %802%33)/.
425
!@>LI< 16.2 0OLH>OVLQF@ QO>KP@OFMQFLK >KA QO>KPI>QFLK L@@RO PFJRIQ>KBLRPIV FK QEB @VQLMI>PJ, >KA OBDRI>QFLK L@@ROP >Q QEB QO>KP@OFMQFLK>I IBSBI. %RH>OVLQF@ DBKB BUMOBPPFLK FP OBDRI>QBA AROFKD QO>KP@OFMQFLK >KA 2.! MOL@BPPFKD, TEF@E Q>HB MI>@B FK QEB KR@IBRP, >KA AROFKD MOLQBFK QO>KPI>QFLK, TEF@E Q>HBP MI>@B FK QEB @VQLMI>PJ. &ROQEBO OBDRI>QFLK J>V L@@RO QEOLRDE MLPQ-QO>KPI>QFLK>I JLAFCF@>QFLKP LC MOLQBFKP.
5FC BGDDCPCLACQ GL RFC PCESJ?RGML MD ECLC CVNPCQQGML @CRUCCL NPMI?PWMRCQ ?LB CSI?PWMRCQ ?PC QSKK?PGXCB GL Table 16.1. 5FC PCESJ?RGML MD ECLC CVNPCQQGML GQ BGQASQQCB GL BCR?GJ GL QS@QCOSCLR KMBSJCQ.
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
LB8IPFK@: FI>8E@JDJ
426
#(!04%2 16 X '%.% %802%33)/.
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
16.2 R +IFB8IPFK@: "<E< -<>LC8K@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC QRCNQ GLTMJTCB GL NPMI?PWMRGA ECLC PCESJ?RGML z &VNJ?GL RFC PMJCQ MD ?ARGT?RMPQ, GLBSACPQ, ?LB PCNPCQQMPQ GL ECLC PCESJ?RGML 5FC %/" MD NPMI?PWMRCQ GQ MPE?LGXCB GLRM ? AGPASJ?P AFPMKMQMKC QSNCPAMGJCB GL RFC LSAJCMGB PCEGML MD RFC ACJJ AWRMNJ?QK. 1PMRCGLQ RF?R ?PC LCCBCB DMP ? QNCAGDGA DSLARGML, MP RF?R ?PC GLTMJTCB GL RFC Q?KC @GMAFCKGA?J N?RFU?W, ?PC CLAMBCB RMECRFCP GL @JMAIQ A?JJCB operons. 'MP CV?KNJC, ?JJ MD RFC ECLCQ LCCBCB RM SQC J?ARMQC ?Q ?L CLCPEW QMSPAC ?PC AMBCB LCVR RM C?AF MRFCP GL RFC J?ARMQC (MP 3(*) MNCPML. *L NPMI?PWMRGA ACJJQ, RFCPC ?PC RFPCC RWNCQ MD PCESJ?RMPW KMJCASJCQ RF?R A?L ?DDCAR RFC CVNPCQQGML MD MNCPMLQ: PCNPCQQMPQ, ?ARGT?RMPQ, ?LB GLBSACPQ. Repressors ?PC NPMRCGLQ RF?R QSNNPCQQ RP?LQAPGNRGML MD ? ECLC GL PCQNMLQC RM ?L CVRCPL?J QRGKSJSQ, UFCPC?Q activators ?PC NPMRCGLQ RF?R GLAPC?QC RFC RP?LQAPGNRGML MD ? ECLC GL PCQNMLQC RM ?L CVRCPL?J QRGKSJSQ. 'GL?JJW, GLBSACPQ ?PC QK?JJ KMJCASJCQ RF?R CGRFCP ?ARGT?RC MP PCNPCQQ RP?LQAPGNRGML BCNCLBGLE ML RFC LCCBQ MD RFC ACJJ ?LB RFC ?T?GJ?@GJGRW MD QS@QRP?RC.
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#?ARCPG? QSAF ?Q E. *630 LCCB ?KGLM ?AGBQ RM QSPTGTC. Tryptophan GQ MLC QSAF ?KGLM ?AGB RF?R E. *630 A?L GLECQR DPMK RFC CLTGPMLKCLR. E. *630 A?L ?JQM QWLRFCQGXC RPWNRMNF?L SQGLE CLXWKCQ RF?R ?PC CLAMBCB @W DGTC ECLCQ. 5FCQC DGTC ECLCQ ?PC LCVR RM C?AF MRFCP GL UF?R GQ A?JJCB RFC tryptophan (trp) operon (Figure 16.3). *D RPWNRMNF?L GQ NPCQCLR GL RFC CLTGPMLKCLR, RFCL E. *630 BMCQ LMR LCCB RM QWLRFCQGXC GR ?LB RFC QUGRAF AMLRPMJJGLE RFC ?ARGT?RGML MD RFC ECLCQ GL RFC ;97 MNCPML GQ QUGRAFCB MDD. )MUCTCP, UFCL RPWNRMNF?L ?T?GJ?@GJGRW GQ JMU, RFC QUGRAF AMLRPMJJGLE RFC MNCPML GQ RSPLCB ML, RP?LQAPGNRGML GQ GLGRG?RCB, RFC ECLCQ ?PC CVNPCQQCB, ?LB RPWNRMNF?L GQ QWLRFCQGXCB.
#(!04%2 16 X '%.% %802%33)/.
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!@>LI< 16.3 4EB CFSB DBKBP QE>Q >OB KBBABA QL PVKQEBPFWB QOVMQLME>K FK E. *630 >OB IL@>QBA KBUQ QL B>@E LQEBO FK QEB ;97 LMBOLK. 7EBK QOVMQLME>K FP MIBKQFCRI, QTL QOVMQLME>K JLIB@RIBP ?FKA QEB OBMOBPPLO MOLQBFK >Q QEB LMBO>QLO PBNRBK@B. 4EFP MEVPF@>IIV ?IL@HP QEB 2.! MLIVJBO>PB COLJ QO>KP@OF?FKD QEB QOVMQLME>K DBKBP. 7EBK QOVMQLME>K FP >?PBKQ, QEB OBMOBPPLO MOLQBFK ALBP KLQ ?FKA QL QEB LMBO>QLO >KA QEB DBKBP >OB QO>KP@OF?BA.
" %/" QCOSCLAC RF?R AMBCQ DMP NPMRCGLQ GQ PCDCPPCB RM ?Q RFC AMBGLE PCEGML. 5FC DGTC AMBGLE PCEGMLQ DMP RFC RPWNRMNF?L @GMQWLRFCQGQ CLXWKCQ ?PC ?PP?LECB QCOSCLRG?JJW ML RFC AFPMKMQMKC GL RFC MNCPML. +SQR @CDMPC RFC AMBGLE PCEGML GQ RFC transcriptional start site. 5FGQ GQ RFC PCEGML MD %/" RM UFGAF 3/" NMJWKCP?QC @GLBQ RM GLGRG?RC RP?LQAPGNRGML. 5FC NPMKMRCP QCOSCLAC GQ SNQRPC?K MD RFC RP?LQAPGNRGML?J QR?PR QGRC; C?AF MNCPML F?Q ? QCOSCLAC UGRFGL MP LC?P RFC NPMKMRCP RM UFGAF NPMRCGLQ (?ARGT?RMPQ MP PCNPCQQMPQ) A?L @GLB ?LB PCESJ?RC RP?LQAPGNRGML. " %/" QCOSCLAC A?JJCB RFC MNCP?RMP QCOSCLAC GQ CLAMBCB @CRUCCL RFC NPMKMRCP PCEGML ?LB RFC DGPQR ;97 AMBGLE ECLC. 5FGQ operator AMLR?GLQ RFC %/" AMBC RM UFGAF RFC PCNPCQQMP NPMRCGL A?L @GLB. 8FCL RPWNRMNF?L GQ NPCQCLR GL RFC ACJJ, RUM RPWNRMNF?L KMJCASJCQ @GLB RM RFC ;97 PCNPCQQMP, UFGAF AF?LECQ QF?NC RM @GLB RM RFC ;97 MNCP?RMP. #GLBGLE MD RFC RPWNRMNF?LpPCNPCQQMP AMKNJCV ?R RFC MNCP?RMP NFWQGA?JJW NPCTCLRQ RFC 3/" NMJWKCP?QC DPMK @GLBGLE, ?LB RP?LQAPG@GLE RFC BMULQRPC?K ECLCQ. 8FCL RPWNRMNF?L GQ LMR NPCQCLR GL RFC ACJJ, RFC PCNPCQQMP @W GRQCJD BMCQ LMR @GLB RM RFC MNCP?RMP; RFCPCDMPC, RFC MNCPML GQ ?ARGTC ?LB RPWNRMNF?L GQ QWLRFCQGXCB. #CA?SQC RFC PCNPCQQMP NPMRCGL ?ARGTCJW @GLBQ RM RFC MNCP?RMP RM ICCN RFC ECLCQ RSPLCB MDD, RFC ;97 MNCPML GQ LCE?RGTCJW PCESJ?RCB ?LB RFC NPMRCGLQ RF?R @GLB RM RFC MNCP?RMP RM QGJCLAC ;97 CVNPCQQGML ?PC negative regulators.
8?RAF this video (http://openstaxcollege.org/l/trp_operon) RM JC?PL KMPC ?@MSR RFC ;97 MNCPML.
C8K89FC@K< A:K@M8KFI +IFK<@E (CA+): AE A:K@M8KFI -<>LC8KFI

+SQR ?Q RFC ;97 MNCPML GQ LCE?RGTCJW PCESJ?RCB @W RPWNRMNF?L KMJCASJCQ, RFCPC ?PC NPMRCGLQ RF?R @GLB RM RFC MNCP?RMP QCOSCLACQ RF?R ?AR ?Q ? positive regulator RM RSPL ECLCQ ML ?LB ?ARGT?RC RFCK. 'MP CV?KNJC, UFCL EJSAMQC GQ QA?PAC, E. *630 @?ARCPG? A?L RSPL RM MRFCP QSE?P QMSPACQ DMP DSCJ. 5M BM RFGQ, LCU ECLCQ RM NPMACQQ RFCQC ?JRCPL?RC ECLCQ KSQR @C RP?LQAPG@CB. 8FCL EJSAMQC JCTCJQ BPMN, AWAJGA ".1 (A".1) @CEGLQ RM ?AASKSJ?RC GL RFC ACJJ. 5FC A".1 KMJCASJC GQ ? QGEL?JGLE KMJCASJC RF?R GQ GLTMJTCB GL EJSAMQC ?LB CLCPEW KCR?@MJGQK GL E. *630. 8FCL EJSAMQC JCTCJQ BCAJGLC GL RFC ACJJ, ?AASKSJ?RGLE A".1 @GLBQ RM RFC NMQGRGTC PCESJ?RMP catabolite activator protein (CAP), ? NPMRCGL RF?R @GLBQ RM RFC NPMKMRCPQ MD MNCPMLQ RF?R AMLRPMJ RFC NPMACQQGLE MD ?JRCPL?RGTC QSE?PQ. 8FCL A".1 @GLBQ RM $"1, RFC AMKNJCV @GLBQ RM RFC
428
#(!04%2 16 X '%.% %802%33)/.
NPMKMRCP PCEGML MD RFC ECLCQ RF?R ?PC LCCBCB RM SQC RFC ?JRCPL?RC QSE?P QMSPACQ (Figure 16.4). *L RFCQC MNCPMLQ, ? $"1 @GLBGLE QGRC GQ JMA?RCB SNQRPC?K MD RFC 3/" NMJWKCP?QC @GLBGLE QGRC GL RFC NPMKMRCP. 5FGQ GLAPC?QCQ RFC @GLBGLE ?@GJGRW MD 3/" NMJWKCP?QC RM RFC NPMKMRCP PCEGML ?LB RFC RP?LQAPGNRGML MD RFC ECLCQ.
!@>LI< 16.4 7EBK DIR@LPB IBSBIP C>II, E. *630 J>V RPB LQEBO PRD>OP CLO CRBI ?RQ JRPQ QO>KP@OF?B KBT DBKBP QL AL PL. !P DIR@LPB PRMMIFBP ?B@LJB IFJFQBA, @!-0 IBSBIP FK@OB>PB. 4EFP @!-0 ?FKAP QL QEB #!0 MOLQBFK, > MLPFQFSB OBDRI>QLO QE>Q ?FKAP QL >K LMBO>QLO OBDFLK RMPQOB>J LC QEB DBKBP OBNRFOBA QL RPB LQEBO PRD>O PLRO@BP.
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5FC RFGPB RWNC MD ECLC PCESJ?RGML GL NPMI?PWMRGA ACJJQ MAASPQ RFPMSEF inducible operons, UFGAF F?TC NPMRCGLQ RF?R @GLB RM ?ARGT?RC MP PCNPCQQ RP?LQAPGNRGML BCNCLBGLE ML RFC JMA?J CLTGPMLKCLR ?LB RFC LCCBQ MD RFC ACJJ. 5FC 3(* MNCPML GQ ? RWNGA?J GLBSAG@JC MNCPML. "Q KCLRGMLCB NPCTGMSQJW, E. *630 GQ ?@JC RM SQC MRFCP QSE?PQ ?Q CLCPEW QMSPACQ UFCL EJSAMQC AMLACLRP?RGMLQ ?PC JMU. 5M BM QM, RFC A".1p$"1 NPMRCGL AMKNJCV QCPTCQ ?Q ? NMQGRGTC PCESJ?RMP RM GLBSAC RP?LQAPGNRGML. 0LC QSAF QSE?P QMSPAC GQ J?ARMQC. 5FC lac operon CLAMBCQ RFC ECLCQ LCACQQ?PW RM ?AOSGPC ?LB NPMACQQ RFC J?ARMQC DPMK RFC JMA?J CLTGPMLKCLR. $"1 @GLBQ RM RFC MNCP?RMP QCOSCLAC SNQRPC?K MD RFC NPMKMRCP RF?R GLGRG?RCQ RP?LQAPGNRGML MD RFC 3(* MNCPML. )MUCTCP, DMP RFC 3(* MNCPML RM @C ?ARGT?RCB, RUM AMLBGRGMLQ KSQR @C KCR. 'GPQR, RFC JCTCJ MD EJSAMQC KSQR @C TCPW JMU MP LML-CVGQRCLR. 4CAMLB, J?ARMQC KSQR @C NPCQCLR. 0LJW UFCL EJSAMQC GQ ?@QCLR ?LB J?ARMQC GQ NPCQCLR UGJJ RFC 3(* MNCPML @C RP?LQAPG@CB (Figure 16.5). 5FGQ K?ICQ QCLQC DMP RFC ACJJ, @CA?SQC GR UMSJB @C CLCPECRGA?JJW U?QRCDSJ RM APC?RC RFC NPMRCGLQ RM NPMACQQ J?ARMQC GD EJSAMQC U?Q NJCLRGDSJ MP J?ARMQC U?Q LMR ?T?GJ?@JC.
#(!04%2 16 X '%.% %802%33)/.
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!@>LI< 16.5 4O>KP@OFMQFLK LC QEB 3(* LMBOLK FP @>OBCRIIV OBDRI>QBA PL QE>Q FQP BUMOBPPFLK LKIV L@@ROP TEBK DIR@LPB FP IFJFQBA >KA I>@QLPB FP MOBPBKQ QL PBOSB >P >K >IQBOK>QFSB CRBI PLRO@B.
)K E. *630, QEB ;97 LMBOLK FP LK ?V ABC>RIQ, TEFIB QEB 3(* LMBOLK FP LCC. 7EV AL VLR QEFKH QEFP FP QEB @>PB *D EJSAMQC GQ ?@QCLR, RFCL $"1 A?L @GLB RM RFC MNCP?RMP QCOSCLAC RM ?ARGT?RC RP?LQAPGNRGML. *D J?ARMQC GQ ?@QCLR, RFCL RFC PCNPCQQMP @GLBQ RM RFC MNCP?RMP RM NPCTCLR RP?LQAPGNRGML. *D CGRFCP MD RFCQC PCOSGPCKCLRQ GQ KCR, RFCL RP?LQAPGNRGML PCK?GLQ MDD. 0LJW UFCL @MRF AMLBGRGMLQ ?PC Q?RGQDGCB GQ RFC 3(* MNCPML RP?LQAPG@CB (Table 16.2).
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#(!04%2 16 X '%.% %802%33)/.
8?RAF ?L animated tutorial (http://openstaxcollege.org/l/lac_operon) ?@MSR RFC UMPIGLEQ MD 3(* MNCPML FCPC.
16.3 R LB8IPFK@: G@><E<K@: "<E< -<>LC8K@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL RFC NPMACQQ MD CNGECLCRGA PCESJ?RGML z %CQAPG@C FMU ?AACQQ RM %/" GQ AMLRPMJJCB @W FGQRMLC KMBGDGA?RGML &SI?PWMRGA ECLC CVNPCQQGML GQ KMPC AMKNJCV RF?L NPMI?PWMRGA ECLC CVNPCQQGML @CA?SQC RFC NPMACQQCQ MD RP?LQAPGNRGML ?LB RP?LQJ?RGML ?PC NFWQGA?JJW QCN?P?RCB. 6LJGIC NPMI?PWMRGA ACJJQ, CSI?PWMRGA ACJJQ A?L PCESJ?RC ECLC CVNPCQQGML ?R K?LW BGDDCPCLR JCTCJQ. &SI?PWMRGA ECLC CVNPCQQGML @CEGLQ UGRF AMLRPMJ MD ?AACQQ RM RFC %/". 5FGQ DMPK MD PCESJ?RGML, A?JJCB CNGECLCRGA PCESJ?RGML, MAASPQ CTCL @CDMPC RP?LQAPGNRGML GQ GLGRG?RCB.
G@><E<K@: CFEKIFC: -<>LC8K@E> A::<JJ KF "<E<J N@K?@E K?< C?IFDFJFD<

5FC FSK?L ECLMKC CLAMBCQ MTCP 20,000 ECLCQ; C?AF MD RFC 23 N?GPQ MD FSK?L AFPMKMQMKCQ CLAMBCQ RFMSQ?LBQ MD ECLCQ. 5FC %/" GL RFC LSAJCSQ GQ NPCAGQCJW UMSLB, DMJBCB, ?LB AMKN?ARCB GLRM AFPMKMQMKCQ QM RF?R GR UGJJ DGR GLRM RFC LSAJCSQ. *R GQ ?JQM MPE?LGXCB QM RF?R QNCAGDGA QCEKCLRQ A?L @C ?AACQQCB ?Q LCCBCB @W ? QNCAGDGA ACJJ RWNC. 5FC DGPQR JCTCJ MD MPE?LGX?RGML, MP N?AIGLE, GQ RFC UGLBGLE MD %/" QRP?LBQ ?PMSLB FGQRMLC NPMRCGLQ. )GQRMLCQ N?AI?EC ?LB MPBCP %/" GLRM QRPSARSP?J SLGRQ A?JJCB LSAJCMQMKC AMKNJCVCQ, UFGAF A?L AMLRPMJ RFC ?AACQQ MD NPMRCGLQ RM RFC %/" PCEGMLQ (Figure 16.6a). 6LBCP RFC CJCARPML KGAPMQAMNC, RFGQ UGLBGLE MD %/" ?PMSLB FGQRMLC NPMRCGLQ RM DMPK LSAJCMQMKCQ JMMIQ JGIC QK?JJ @C?BQ ML ? QRPGLE (Figure 16.6b). 5FCQC @C?BQ (FGQRMLC NPMRCGLQ) A?L KMTC ?JMLE RFC QRPGLE (%/") ?LB AF?LEC RFC QRPSARSPC MD RFC KMJCASJC.
!@>LI< 16.6 $.! FP CLIABA >OLRKA EFPQLKB MOLQBFKP QL @OB>QB (>) KR@IBLPLJB @LJMIBUBP. 4EBPB KR@IBLPLJBP @LKQOLI QEB >@@BPP LC MOLQBFKP QL QEB RKABOIVFKD $.!. 7EBK SFBTBA QEOLRDE >K BIB@QOLK JF@OLP@LMB (?), QEB KR@IBLPLJBP ILLH IFHB ?B>AP LK > PQOFKD. (@OBAFQ xJF@OLDO>MEy: JLAFCF@>QFLK LC TLOH ?V #EOFP 7LLA@L@H)
*D %/" CLAMBGLE ? QNCAGDGA ECLC GQ RM @C RP?LQAPG@CB GLRM 3/", RFC LSAJCMQMKCQ QSPPMSLBGLE RF?R PCEGML MD %/" A?L QJGBC BMUL RFC %/" RM MNCL RF?R QNCAGDGA AFPMKMQMK?J PCEGML ?LB ?JJMU DMP RFC RP?LQAPGNRGML?J K?AFGLCPW (3/" NMJWKCP?QC) RM GLGRG?RC RP?LQAPGNRGML (Figure 16.7). /SAJCMQMKCQ A?L
#(!04%2 16 X '%.% %802%33)/.
431
KMTC RM MNCL RFC AFPMKMQMKC QRPSARSPC RM CVNMQC ? QCEKCLR MD %/", @SR BM QM GL ? TCPW AMLRPMJJCB K?LLCP.
!@>LI< 16.7 .R@IBLPLJBP @>K PIFAB >ILKD $.!. 7EBK KR@IBLPLJBP >OB PM>@BA @ILPBIV QLDBQEBO (QLM), QO>KP@OFMQFLK C>@QLOP @>KKLQ ?FKA >KA DBKB BUMOBPPFLK FP QROKBA LCC. 7EBK QEB KR@IBLPLJBP >OB PM>@BA C>O >M>OQ (?LQQLJ), QEB $.! FP BUMLPBA. 4O>KP@OFMQFLK C>@QLOP @>K ?FKA, >IILTFKD DBKB BUMOBPPFLK QL L@@RO. -LAFCF@>QFLKP QL QEB EFPQLKBP >KA $.! >CCB@Q KR@IBLPLJB PM>@FKD.
)K CBJ>IBP, LKB LC QEB QTL 8 @EOLJLPLJBP FP FK>@QFS>QBA AROFKD BJ?OVLKF@ ABSBILMJBKQ ?B@>RPB LC BMFDBKBQF@ @E>KDBP QL QEB @EOLJ>QFK. 7E>Q FJM>@Q AL VLR QEFKH QEBPB @E>KDBP TLRIA E>SB LK KR@IBLPLJB M>@HFKD )MU RFC FGQRMLC NPMRCGLQ KMTC GQ BCNCLBCLR ML QGEL?JQ DMSLB ML @MRF RFC FGQRMLC NPMRCGLQ ?LB ML RFC %/". 5FCQC QGEL?JQ ?PC R?EQ ?BBCB RM FGQRMLC NPMRCGLQ ?LB %/" RF?R RCJJ RFC FGQRMLCQ GD ? AFPMKMQMK?J PCEGML QFMSJB @C MNCL MP AJMQCB (Figure 16.8 BCNGARQ KMBGDGA?RGMLQ RM FGQRMLC NPMRCGLQ ?LB %/"). 5FCQC R?EQ ?PC LMR NCPK?LCLR, @SR K?W @C ?BBCB MP PCKMTCB ?Q LCCBCB. 5FCW ?PC AFCKGA?J KMBGDGA?RGMLQ (NFMQNF?RC, KCRFWJ, MP ?ACRWJ EPMSNQ) RF?R ?PC ?RR?AFCB RM QNCAGDGA ?KGLM ?AGBQ GL RFC NPMRCGL MP RM RFC LSAJCMRGBCQ MD RFC %/". 5FC R?EQ BM LMR ?JRCP RFC %/" @?QC QCOSCLAC, @SR RFCW BM ?JRCP FMU RGEFRJW UMSLB RFC %/" GQ ?PMSLB RFC FGQRMLC NPMRCGLQ. %/" GQ ? LCE?RGTCJW AF?PECB KMJCASJC; RFCPCDMPC, AF?LECQ GL RFC AF?PEC MD RFC FGQRMLC UGJJ AF?LEC FMU RGEFRJW UMSLB RFC %/" KMJCASJC UGJJ @C. 8FCL SLKMBGDGCB, RFC FGQRMLC NPMRCGLQ F?TC ? J?PEC NMQGRGTC AF?PEC; @W ?BBGLE AFCKGA?J KMBGDGA?RGMLQ JGIC ?ACRWJ EPMSNQ, RFC AF?PEC @CAMKCQ JCQQ NMQGRGTC. 5FC %/" KMJCASJC GRQCJD A?L ?JQM @C KMBGDGCB. 5FGQ MAASPQ UGRFGL TCPW QNCAGDGA PCEGMLQ A?JJCB $N( GQJ?LBQ. 5FCQC ?PC QRPCRAFCQ UGRF ? FGEF DPCOSCLAW MD AWRMQGLC ?LB ES?LGLC BGLSAJCMRGBC %/" N?GPQ ($() DMSLB GL RFC NPMKMRCP PCEGMLQ MD ECLCQ. 8FCL RFGQ AMLDGESP?RGML CVGQRQ, RFC AWRMQGLC KCK@CP MD RFC N?GP A?L @C KCRFWJ?RCB (? KCRFWJ EPMSN GQ ?BBCB). 5FGQ KMBGDGA?RGML AF?LECQ FMU RFC %/" GLRCP?ARQ UGRF NPMRCGLQ, GLAJSBGLE RFC FGQRMLC NPMRCGLQ RF?R AMLRPMJ ?AACQQ RM RFC PCEGML. )GEFJW KCRFWJ?RCB (FWNCPKCRFWJ?RCB) %/" PCEGMLQ UGRF BC?ACRWJ?RCB FGQRMLCQ ?PC RGEFRJW AMGJCB ?LB RP?LQAPGNRGML?JJW GL?ARGTC.
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!@>LI< 16.8 (FPQLKB MOLQBFKP >KA $.! KR@IBLQFABP @>K ?B JLAFCFBA @EBJF@>IIV. -LAFCF@>QFLKP >CCB@Q KR@IBLPLJB PM>@FKD >KA DBKB BUMOBPPFLK. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .)()
5FGQ RWNC MD ECLC PCESJ?RGML GQ A?JJCB CNGECLCRGA PCESJ?RGML. &NGECLCRGA KC?LQ t?PMSLB ECLCRGAQ.u 5FC AF?LECQ RF?R MAASP RM RFC FGQRMLC NPMRCGLQ ?LB %/" BM LMR ?JRCP RFC LSAJCMRGBC QCOSCLAC ?LB ?PC LMR NCPK?LCLR. *LQRC?B, RFCQC AF?LECQ ?PC RCKNMP?PW (?JRFMSEF RFCW MDRCL NCPQGQR RFPMSEF KSJRGNJC PMSLBQ MD ACJJ BGTGQGML) ?LB ?JRCP RFC AFPMKMQMK?J QRPSARSPC (MNCL MP AJMQCB) ?Q LCCBCB. " ECLC A?L @C RSPLCB ML MP MDD BCNCLBGLE SNML RFC JMA?RGML ?LB KMBGDGA?RGMLQ RM RFC FGQRMLC NPMRCGLQ ?LB %/". *D ? ECLC GQ RM @C RP?LQAPG@CB, RFC FGQRMLC NPMRCGLQ ?LB %/" ?PC KMBGDGCB QSPPMSLBGLE RFC AFPMKMQMK?J PCEGML CLAMBGLE RF?R ECLC. 5FGQ MNCLQ RFC AFPMKMQMK?J PCEGML RM ?JJMU ?AACQQ DMP 3/" NMJWKCP?QC ?LB MRFCP NPMRCGLQ, A?JJCB transcription factors, RM @GLB RM RFC NPMKMRCP PCEGML, JMA?RCB HSQR SNQRPC?K MD RFC ECLC, ?LB GLGRG?RC RP?LQAPGNRGML. *D ? ECLC GQ RM PCK?GL RSPLCB MDD, MP QGJCLACB, RFC FGQRMLC NPMRCGLQ ?LB %/" F?TC BGDDCPCLR KMBGDGA?RGMLQ RF?R QGEL?J ? AJMQCB AFPMKMQMK?J AMLDGESP?RGML. *L RFGQ AJMQCB AMLDGESP?RGML, RFC 3/" NMJWKCP?QC ?LB RP?LQAPGNRGML D?ARMPQ BM LMR F?TC ?AACQQ RM RFC %/" ?LB RP?LQAPGNRGML A?LLMR MAASP (Figure 16.7).
7GCU this video (http://openstaxcollege.org/l/epigenetic_reg) RF?R BCQAPG@CQ FMU CNGECLCRGA PCESJ?RGML AMLRPMJQ ECLC CVNPCQQGML.
16.4 R LB8IPFK@: /I8EJ:I@GK@FE "<E< -<>LC8K@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %GQASQQ RFC PMJC MD RP?LQAPGNRGML D?ARMPQ GL ECLC PCESJ?RGML z &VNJ?GL FMU CLF?LACPQ ?LB PCNPCQQMPQ PCESJ?RC ECLC CVNPCQQGML
#(!04%2 16 X '%.% %802%33)/.
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-GIC NPMI?PWMRGA ACJJQ, RFC RP?LQAPGNRGML MD ECLCQ GL CSI?PWMRCQ PCOSGPCQ RFC ?ARGMLQ MD ?L 3/" NMJWKCP?QC RM @GLB RM ? QCOSCLAC SNQRPC?K MD ? ECLC RM GLGRG?RC RP?LQAPGNRGML. )MUCTCP, SLJGIC NPMI?PWMRGA ACJJQ, RFC CSI?PWMRGA 3/" NMJWKCP?QC PCOSGPCQ MRFCP NPMRCGLQ, MP RP?LQAPGNRGML D?ARMPQ, RM D?AGJGR?RC RP?LQAPGNRGML GLGRG?RGML. 5P?LQAPGNRGML D?ARMPQ ?PC NPMRCGLQ RF?R @GLB RM RFC NPMKMRCP QCOSCLAC ?LB MRFCP PCESJ?RMPW QCOSCLACQ RM AMLRPMJ RFC RP?LQAPGNRGML MD RFC R?PECR ECLC. 3/" NMJWKCP?QC @W GRQCJD A?LLMR GLGRG?RC RP?LQAPGNRGML GL CSI?PWMRGA ACJJQ. 5P?LQAPGNRGML D?ARMPQ KSQR @GLB RM RFC NPMKMRCP PCEGML DGPQR ?LB PCAPSGR 3/" NMJWKCP?QC RM RFC QGRC DMP RP?LQAPGNRGML RM @C CQR?@JGQFCB.
7GCU RFC NPMACQQ MD RP?LQAPGNRGMLqRFC K?IGLE MD 3/" DPMK ? %/" RCKNJ?RCq?R this site (http://openstaxcollege.org/l/transcript_RNA) .
/?< +IFDFK<I 8E; K?< /I8EJ:I@GK@FE (8:?@E<IP

(CLCQ ?PC MPE?LGXCB RM K?IC RFC AMLRPMJ MD ECLC CVNPCQQGML C?QGCP. 5FC NPMKMRCP PCEGML GQ GKKCBG?RCJW SNQRPC?K MD RFC AMBGLE QCOSCLAC. 5FGQ PCEGML A?L @C QFMPR (MLJW ? DCU LSAJCMRGBCQ GL JCLERF) MP OSGRC JMLE (FSLBPCBQ MD LSAJCMRGBCQ JMLE). 5FC JMLECP RFC NPMKMRCP, RFC KMPC ?T?GJ?@JC QN?AC DMP NPMRCGLQ RM @GLB. 5FGQ ?JQM ?BBQ KMPC AMLRPMJ RM RFC RP?LQAPGNRGML NPMACQQ. 5FC JCLERF MD RFC NPMKMRCP GQ ECLC-QNCAGDGA ?LB A?L BGDDCP BP?K?RGA?JJW @CRUCCL ECLCQ. $MLQCOSCLRJW, RFC JCTCJ MD AMLRPMJ MD ECLC CVNPCQQGML A?L ?JQM BGDDCP OSGRC BP?K?RGA?JJW @CRUCCL ECLCQ. 5FC NSPNMQC MD RFC NPMKMRCP GQ RM @GLB RP?LQAPGNRGML D?ARMPQ RF?R AMLRPMJ RFC GLGRG?RGML MD RP?LQAPGNRGML. 8GRFGL RFC NPMKMRCP PCEGML, HSQR SNQRPC?K MD RFC RP?LQAPGNRGML?J QR?PR QGRC, PCQGBCQ RFC 5"5" @MV. 5FGQ @MV GQ QGKNJW ? PCNC?R MD RFWKGLC ?LB ?BCLGLC BGLSAJCMRGBCQ (JGRCP?JJW, 5"5" PCNC?RQ). 3/" NMJWKCP?QC @GLBQ RM RFC RP?LQAPGNRGML GLGRG?RGML AMKNJCV, ?JJMUGLE RP?LQAPGNRGML RM MAASP. 5M GLGRG?RC RP?LQAPGNRGML, ? RP?LQAPGNRGML D?ARMP (5'**%) GQ RFC DGPQR RM @GLB RM RFC 5"5" @MV. #GLBGLE MD 5'**% PCAPSGRQ MRFCP RP?LQAPGNRGML D?ARMPQ, GLAJSBGLE 5'**#, 5'**&, 5'**', ?LB 5'**) RM RFC 5"5" @MV. 0LAC RFGQ AMKNJCV GQ ?QQCK@JCB, 3/" NMJWKCP?QC A?L @GLB RM GRQ SNQRPC?K QCOSCLAC. 8FCL @MSLB ?JMLE UGRF RFC RP?LQAPGNRGML D?ARMPQ, 3/" NMJWKCP?QC GQ NFMQNFMPWJ?RCB. 5FGQ PCJC?QCQ N?PR MD RFC NPMRCGL DPMK RFC %/" RM ?ARGT?RC RFC RP?LQAPGNRGML GLGRG?RGML AMKNJCV ?LB NJ?ACQ 3/" NMJWKCP?QC GL RFC AMPPCAR MPGCLR?RGML RM @CEGL RP?LQAPGNRGML; %/"-@CLBGLE NPMRCGL @PGLEQ RFC CLF?LACP, UFGAF A?L @C OSGRC ? BGQR?LAC DPMK RFC ECLC, GL AMLR?AR UGRF RP?LQAPGNRGML D?ARMPQ ?LB KCBG?RMP NPMRCGLQ (Figure 16.9).
434
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!@>LI< 16.9 !K BKE>K@BO FP > $.! PBNRBK@B QE>Q MOLJLQBP QO>KP@OFMQFLK. %>@E BKE>K@BO FP J>AB RM LC PELOQ $.! PBNRBK@BP @>IIBA AFPQ>I @LKQOLI BIBJBKQP. !@QFS>QLOP ?LRKA QL QEB AFPQ>I @LKQOLI BIBJBKQP FKQBO>@Q TFQE JBAF>QLO MOLQBFKP >KA QO>KP@OFMQFLK C>@QLOP. 4TL AFCCBOBKQ DBKBP J>V E>SB QEB P>JB MOLJLQBO ?RQ AFCCBOBKQ AFPQ>I @LKQOLI BIBJBKQP, BK>?IFKD AFCCBOBKQF>I DBKB BUMOBPPFLK.
*L ?BBGRGML RM RFC ECLCP?J RP?LQAPGNRGML D?ARMPQ, MRFCP RP?LQAPGNRGML D?ARMPQ A?L @GLB RM RFC NPMKMRCP RM PCESJ?RC ECLC RP?LQAPGNRGML. 5FCQC RP?LQAPGNRGML D?ARMPQ @GLB RM RFC NPMKMRCPQ MD ? QNCAGDGA QCR MD ECLCQ. 5FCW ?PC LMR ECLCP?J RP?LQAPGNRGML D?ARMPQ RF?R @GLB RM CTCPW NPMKMRCP AMKNJCV, @SR ?PC PCAPSGRCB RM ? QNCAGDGA QCOSCLAC ML RFC NPMKMRCP MD ? QNCAGDGA ECLC. 5FCPC ?PC FSLBPCBQ MD RP?LQAPGNRGML D?ARMPQ GL ? ACJJ RF?R C?AF @GLB QNCAGDGA?JJW RM ? N?PRGASJ?P %/" QCOSCLAC KMRGD. 8FCL RP?LQAPGNRGML D?ARMPQ @GLB RM RFC NPMKMRCP HSQR SNQRPC?K MD RFC CLAMBCB ECLC, GR GQ PCDCPPCB RM ?Q ? cis-acting element, @CA?SQC GR GQ ML RFC Q?KC AFPMKMQMKC HSQR LCVR RM RFC ECLC. 5FC PCEGML RF?R ? N?PRGASJ?P RP?LQAPGNRGML D?ARMP @GLBQ RM GQ A?JJCB RFC transcription factor binding site. 5P?LQAPGNRGML D?ARMPQ PCQNMLB RM CLTGPMLKCLR?J QRGKSJG RF?R A?SQC RFC NPMRCGLQ RM DGLB RFCGP @GLBGLE QGRCQ ?LB GLGRG?RC RP?LQAPGNRGML MD RFC ECLC RF?R GQ LCCBCB.
E?8E:<IJ 8E; /I8EJ:I@GK@FE

*L QMKC CSI?PWMRGA ECLCQ, RFCPC ?PC PCEGMLQ RF?R FCJN GLAPC?QC MP CLF?LAC RP?LQAPGNRGML. 5FCQC PCEGMLQ, A?JJCB enhancers, ?PC LMR LCACQQ?PGJW AJMQC RM RFC ECLCQ RFCW CLF?LAC. 5FCW A?L @C JMA?RCB SNQRPC?K MD ? ECLC, UGRFGL RFC AMBGLE PCEGML MD RFC ECLC, BMULQRPC?K MD ? ECLC, MP K?W @C RFMSQ?LBQ MD LSAJCMRGBCQ ?U?W. &LF?LACP PCEGMLQ ?PC @GLBGLE QCOSCLACQ, MP QGRCQ, DMP RP?LQAPGNRGML D?ARMPQ. 8FCL ? %/"-@CLBGLE NPMRCGL @GLBQ, RFC QF?NC MD RFC %/" AF?LECQ (Figure 16.9). 5FGQ QF?NC AF?LEC ?JJMUQ DMP RFC GLRCP?ARGML MD RFC ?ARGT?RMPQ @MSLB RM RFC CLF?LACPQ UGRF RFC RP?LQAPGNRGML D?ARMPQ @MSLB RM RFC NPMKMRCP PCEGML ?LB RFC 3/" NMJWKCP?QC. 8FCPC?Q %/" GQ ECLCP?JJW BCNGARCB ?Q ? QRP?GEFR JGLC GL RUM BGKCLQGMLQ, GR GQ ?ARS?JJW ? RFPCC-BGKCLQGML?J M@HCAR. 5FCPCDMPC, ? LSAJCMRGBC QCOSCLAC RFMSQ?LBQ MD LSAJCMRGBCQ ?U?W A?L DMJB MTCP ?LB GLRCP?AR UGRF ? QNCAGDGA NPMKMRCP.
/LIE@E> "<E<J *==: /I8EJ:I@GK@FE8C -<GI<JJFIJ

-GIC NPMI?PWMRGA ACJJQ, CSI?PWMRGA ACJJQ ?JQM F?TC KCAF?LGQKQ RM NPCTCLR RP?LQAPGNRGML. 5P?LQAPGNRGML?J PCNPCQQMPQ A?L @GLB RM NPMKMRCP MP CLF?LACP PCEGMLQ ?LB @JMAI RP?LQAPGNRGML. -GIC RFC RP?LQAPGNRGML?J ?ARGT?RMPQ, PCNPCQQMPQ PCQNMLB RM CVRCPL?J QRGKSJG RM NPCTCLR RFC @GLBGLE MD ?ARGT?RGLE RP?LQAPGNRGML D?ARMPQ.
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16.5 R LB8IPFK@: +FJK-KI8EJ:I@GK@FE8C "<E< -<>LC8K@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z 6LBCPQR?LB 3/" QNJGAGLE ?LB CVNJ?GL GRQ PMJC GL PCESJ?RGLE ECLC CVNPCQQGML z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
-)A JGC@:@E>, K?< =@IJK JK8>< F= GFJK-KI8EJ:I@GK@FE8C :FEKIFC

*L CSI?PWMRGA ACJJQ, RFC 3/" RP?LQAPGNR MDRCL AMLR?GLQ PCEGMLQ, A?JJCB GLRPMLQ, RF?R ?PC PCKMTCB NPGMP RM RP?LQJ?RGML. 5FC PCEGMLQ MD 3/" RF?R AMBC DMP NPMRCGL ?PC A?JJCB CVMLQ (Figure 16.10). "DRCP ?L 3/" KMJCASJC F?Q @CCL RP?LQAPG@CB, @SR NPGMP RM GRQ BCN?PRSPC DPMK RFC LSAJCSQ RM @C RP?LQJ?RCB, RFC 3/" GQ NPMACQQCB ?LB RFC GLRPMLQ ?PC PCKMTCB @W QNJGAGLE.
!@>LI< 16.10 0OB-J2.! @>K ?B >IQBOK>QFSBIV PMIF@BA QL @OB>QB AFCCBOBKQ MOLQBFKP.
436
#(!04%2 16 X '%.% %802%33)/.
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!@>LI< 16.11 4EBOB >OB CFSB ?>PF@ JLABP LC >IQBOK>QFSB PMIF@FKD.
(LT @LRIA >IQBOK>QFSB PMIF@FKD BSLISB )KQOLKP E>SB > ?BDFKKFKD >KA BKAFKD OB@LDKFQFLK PBNRBK@B; FQ FP B>PV QL FJ>DFKB QEB C>FIROB LC QEB PMIF@FKD JB@E>KFPJ QL FABKQFCV QEB BKA LC >K FKQOLK >KA FKPQB>A CFKA QEB BKA LC QEB KBUQ FKQOLK, QERP OBJLSFKD QTL FKQOLKP >KA QEB FKQBOSBKFKD BULK. )K C>@Q, QEBOB >OB JB@E>KFPJP FK MI>@B QL MOBSBKQ PR@E FKQOLK PHFMMFKD, ?RQ JRQ>QFLKP >OB IFHBIV QL IB>A QL QEBFO C>FIROB. 3R@E xJFPQ>HBPy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437
7GQS?JGXC FMU K3/" QNJGAGLE F?NNCLQ @W U?RAFGLE RFC NPMACQQ GL ?ARGML GL this video (http://openstaxcollege.org/l/mRNA_splicing) .
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#CDMPC RFC K3/" JC?TCQ RFC LSAJCSQ, GR GQ EGTCL RUM NPMRCARGTC "A?NQ" RF?R NPCTCLR RFC CLB MD RFC QRP?LB DPMK BCEP?BGLE BSPGLE GRQ HMSPLCW. 5FC 5' cap, UFGAF GQ NJ?ACB ML RFC 5' CLB MD RFC K3/", GQ SQS?JJW AMKNMQCB MD ? KCRFWJ?RCB ES?LMQGLC RPGNFMQNF?RC KMJCASJC ((51). 5FC poly-A tail, UFGAF GQ ?RR?AFCB RM RFC 3' CLB, GQ SQS?JJW AMKNMQCB MD ? QCPGCQ MD ?BCLGLC LSAJCMRGBCQ. 0LAC RFC 3/" GQ RP?LQNMPRCB RM RFC AWRMNJ?QK, RFC JCLERF MD RGKC RF?R RFC 3/" PCQGBCQ RFCPC A?L @C AMLRPMJJCB. &?AF 3/" KMJCASJC F?Q ? BCDGLCB JGDCQN?L ?LB BCA?WQ ?R ? QNCAGDGA P?RC. 5FGQ P?RC MD BCA?W A?L GLDJSCLAC FMU KSAF NPMRCGL GQ GL RFC ACJJ. *D RFC BCA?W P?RC GQ GLAPC?QCB, RFC 3/" UGJJ LMR CVGQR GL RFC AWRMNJ?QK ?Q JMLE, QFMPRCLGLE RFC RGKC DMP RP?LQJ?RGML RM MAASP. $MLTCPQCJW, GD RFC P?RC MD BCA?W GQ BCAPC?QCB, RFC 3/" KMJCASJC UGJJ PCQGBC GL RFC AWRMNJ?QK JMLECP ?LB KMPC NPMRCGL A?L @C RP?LQJ?RCB. 5FGQ P?RC MD BCA?W GQ PCDCPPCB RM ?Q RFC 3/" QR?@GJGRW. *D RFC 3/" GQ QR?@JC, GR UGJJ @C BCRCARCB DMP JMLECP NCPGMBQ MD RGKC GL RFC AWRMNJ?QK. #GLBGLE MD NPMRCGLQ RM RFC 3/" A?L GLDJSCLAC GRQ QR?@GJGRW. 1PMRCGLQ, A?JJCB RNA-binding proteins, MP 3#1Q, A?L @GLB RM RFC PCEGMLQ MD RFC 3/" HSQR SNQRPC?K MP BMULQRPC?K MD RFC NPMRCGL-AMBGLE PCEGML. 5FCQC PCEGMLQ GL RFC 3/" RF?R ?PC LMR RP?LQJ?RCB GLRM NPMRCGL ?PC A?JJCB RFC untranslated regions, MP 653Q. 5FCW ?PC LMR GLRPMLQ (RFMQC F?TC @CCL PCKMTCB GL RFC LSAJCSQ). 3?RFCP, RFCQC ?PC PCEGMLQ RF?R PCESJ?RC K3/" JMA?JGX?RGML, QR?@GJGRW, ?LB NPMRCGL RP?LQJ?RGML. 5FC PCEGML HSQR @CDMPC RFC NPMRCGL-AMBGLE PCEGML GQ A?JJCB RFC 5' UTR, UFCPC?Q RFC PCEGML ?DRCP RFC AMBGLE PCEGML GQ A?JJCB RFC 3' UTR (Figure 16.12). 5FC @GLBGLE MD 3#1Q RM RFCQC PCEGMLQ A?L GLAPC?QC MP BCAPC?QC RFC QR?@GJGRW MD ?L 3/" KMJCASJC, BCNCLBGLE ML RFC QNCAGDGA 3#1 RF?R @GLBQ.
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-GIC RP?LQAPGNRGML, RP?LQJ?RGML GQ AMLRPMJJCB @W NPMRCGLQ RF?R @GLB ?LB GLGRG?RC RFC NPMACQQ. *L RP?LQJ?RGML, RFC AMKNJCV RF?R ?QQCK@JCQ RM QR?PR RFC NPMACQQ GQ PCDCPPCB RM ?Q RFC initiation complex. 5FC DGPQR NPMRCGL RM @GLB RM RFC 3/" RM GLGRG?RC RP?LQJ?RGML GQ RFC eukaryotic initiation factor-2 (eIF-2). 5FC C*'-2 NPMRCGL GQ ?ARGTC UFCL GR @GLBQ RM RFC FGEF-CLCPEW KMJCASJC guanosine triphosphate (GTP). (51 NPMTGBCQ RFC CLCPEW RM QR?PR RFC PC?ARGML @W EGTGLE SN ? NFMQNF?RC ?LB @CAMKGLE guanosine diphosphate (GDP). 5FC C*'-2 NPMRCGL @MSLB RM (51 @GLBQ RM RFC QK?JJ 40S ribosomal subunit. 8FCL @MSLB, RFC KCRFGMLGLC GLGRG?RMP R3/" ?QQMAG?RCQ UGRF RFC C*'-2/404 PG@MQMKC AMKNJCV, @PGLEGLE ?JMLE UGRF GR RFC K3/" RM @C RP?LQJ?RCB. "R RFGQ NMGLR, UFCL RFC GLGRG?RMP AMKNJCV GQ ?QQCK@JCB, RFC (51 GQ AMLTCPRCB GLRM (%1 ?LB CLCPEW GQ PCJC?QCB. 5FC NFMQNF?RC ?LB RFC C*'-2 NPMRCGL ?PC PCJC?QCB DPMK RFC AMKNJCV ?LB RFC J?PEC 60S ribosomal subunit @GLBQ RM RP?LQJ?RC RFC 3/". 5FC @GLBGLE MD C*'-2 RM RFC 3/" GQ AMLRPMJJCB @W NFMQNFMPWJ?RGML. *D C*'-2 GQ NFMQNFMPWJ?RCB, GR SLBCPEMCQ ? AMLDMPK?RGML?J AF?LEC ?LB A?LLMR @GLB RM (51. 5FCPCDMPC, RFC GLGRG?RGML AMKNJCV A?LLMR DMPK NPMNCPJW ?LB RP?LQJ?RGML GQ GKNCBCB (Figure 16.13). 8FCL C*'-2 PCK?GLQ SLNFMQNFMPWJ?RCB, GR @GLBQ RFC 3/" ?LB ?ARGTCJW RP?LQJ?RCQ RFC NPMRCGL.
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439
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C8E:<I 8E; +FJK-KI8EJ:I@GK@FE8C CFEKIFC

$F?LECQ GL RFC NMQR-RP?LQAPGNRGML?J AMLRPMJ MD ? ECLC A?L ?JQM PCQSJR GL A?LACP. 3CACLRJW, QCTCP?J EPMSNQ MD PCQC?PAFCPQ F?TC QFMUL RF?R QNCAGDGA A?LACPQ F?TC ?JRCPCB CVNPCQQGML MD KG3/"Q. #CA?SQC KG3/"Q @GLB RM RFC 3' 653 MD 3/" KMJCASJCQ RM BCEP?BC RFCK, MTCPCVNPCQQGML MD RFCQC KG3/"Q AMSJB @C BCRPGKCLR?J RM LMPK?J ACJJSJ?P ?ARGTGRW. 5MM K?LW KG3/"Q AMSJB BP?K?RGA?JJW BCAPC?QC RFC 3/" NMNSJ?RGML JC?BGLE RM ? BCAPC?QC GL NPMRCGL CVNPCQQGML. 4CTCP?J QRSBGCQ F?TC BCKMLQRP?RCB ? AF?LEC GL RFC KG3/" NMNSJ?RGML GL QNCAGDGA A?LACP RWNCQ. *R ?NNC?PQ RF?R RFC QS@QCR MD KG3/"Q CVNPCQQCB GL @PC?QR A?LACP ACJJQ GQ OSGRC BGDDCPCLR DPMK RFC QS@QCR CVNPCQQCB GL JSLE A?LACP ACJJQ MP CTCL DPMK LMPK?J @PC?QR ACJJQ. 5FGQ QSEECQRQ RF?R ?JRCP?RGMLQ GL KG3/" ?ARGTGRW A?L AMLRPG@SRC RM RFC EPMURF MD @PC?QR A?LACP ACJJQ. 5FCQC RWNCQ MD QRSBGCQ ?JQM QSEECQR RF?R GD QMKC KG3/"Q ?PC QNCAGDGA?JJW CVNPCQQCB MLJW GL A?LACP ACJJQ, RFCW AMSJB @C NMRCLRG?J BPSE R?PECRQ. *R UMSJB, RFCPCDMPC, @C AMLACGT?@JC RF?R LCU BPSEQ RF?R RSPL MDD KG3/" CVNPCQQGML GL A?LACP AMSJB @C ?L CDDCARGTC KCRFMB RM RPC?R A?LACP.
C8E:<I 8E; /I8EJC8K@FE8C/+FJK-KI8EJC8K@FE8C CFEKIFC

5FCPC ?PC K?LW CV?KNJCQ MD FMU RP?LQJ?RGML?J MP NMQR-RP?LQJ?RGML?J KMBGDGA?RGMLQ MD NPMRCGLQ ?PGQC GL A?LACP. .MBGDGA?RGMLQ ?PC DMSLB GL A?LACP ACJJQ DPMK RFC GLAPC?QCB RP?LQJ?RGML MD ? NPMRCGL RM AF?LECQ GL NPMRCGL NFMQNFMPWJ?RGML RM ?JRCPL?RGTC QNJGAC T?PG?LRQ MD ? NPMRCGL. "L CV?KNJC MD FMU RFC CVNPCQQGML MD ?L ?JRCPL?RGTC DMPK MD ? NPMRCGL A?L F?TC BP?K?RGA?JJW BGDDCPCLR MSRAMKCQ GQ QCCL GL AMJML A?LACP ACJJQ. 5FC A-'JGN NPMRCGL, ? NPMRCGL GLTMJTCB GL KCBG?RGLE RFC ACJJ BC?RF N?RFU?W, AMKCQ GL RUM DMPKQ: JMLE (A-'-*1-) ?LB QFMPR (A-'-*14). #MRF DMPKQ ?NNC?P RM @C GLTMJTCB GL GLGRG?RGLE AMLRPMJJCB ACJJ BC?RF KCAF?LGQKQ GL LMPK?J ACJJQ. )MUCTCP, GL AMJML A?LACP ACJJQ, CVNPCQQGML MD RFC JMLE DMPK PCQSJRQ GL GLAPC?QCB ACJJ EPMURF GLQRC?B MD ACJJ BC?RF. $JC?PJW, RFC CVNPCQQGML MD RFC UPMLE NPMRCGL BP?K?RGA?JJW ?JRCPQ ACJJ DSLARGML ?LB AMLRPG@SRCQ RM RFC BCTCJMNKCLR MD A?LACP.
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
CC@E@:8C /I@8C CFFI;@E8KFI

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442
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)/8-8:K@E> <C<D<EK RP?LQAPGNRGML D?ARMP @GLBGLE QGRCQ UGRFGL RFC NPMKMRCP RF?R PCESJ?RC RFC RP?LQAPGNRGML MD ? ECLC ?BH?ACLR RM GR 97'38-8:K@E> <C<D<EK RP?LQAPGNRGML D?ARMP @GLBGLE QGRC DMSLB MSRQGBC RFC NPMKMRCP MP ML ?LMRFCP AFPMKMQMKC RF?R GLDJSCLACQ RFC RP?LQAPGNRGML MD ? N?PRGASJ?P ECLC 3' 0/- 3' SLRP?LQJ?RCB PCEGML; PCEGML HSQR BMULQRPC?K MD RFC NPMRCGL-AMBGLE PCEGML GL ?L 3/" KMJCASJC RF?R GQ LMR RP?LQJ?RCB 5' 0/- 5' SLRP?LQJ?RCB PCEGML; PCEGML HSQR SNQRPC?K MD RFC NPMRCGL-AMBGLE PCEGML GL ?L 3/" KMJCASJC RF?R GQ LMR RP?LQJ?RCB 5' :8G ? KCRFWJ?RCB ES?LMQGLC RPGNFMQNF?RC ((51) KMJCASJC RF?R GQ ?RR?AFCB RM RFC 5' CLB MD ? KCQQCLECP 3/" RM NPMRCAR RFC CLB DPMK BCEP?B?RGML 8:K@M8KFI NPMRCGL RF?R @GLBQ RM NPMI?PWMRGA MNCP?RMPQ RM GLAPC?QC RP?LQAPGNRGML :8K89FC@K< 8:K@M8KFI GIFK<@E (CA+) NPMRCGL RF?R AMKNJCVCQ UGRF A".1 RM @GLB RM RFC NPMKMRCP QCOSCLACQ MD MNCPMLQ RF?R AMLRPMJ QSE?P NPMACQQGLE UFCL EJSAMQC GQ LMR ?T?GJ?@JC D)A D<K?PC8K@FE CNGECLCRGA KMBGDGA?RGML RF?R JC?BQ RM ECLC QGJCLAGLE; AMKKMLJW DMSLB GL A?LACP ACJJQ ;@:<I CLXWKC RF?R AFMNQ RFC NPC-KG3/" GLRM RFC K?RSPC DMPK MD RFC KG3/" <E?8E:<E<K@: FCPGR?@JC AF?LECQ RF?R BM LMR GLTMJTC AF?LECQ GL RFC %/" QCOSCLAC <LB8IPFK@: @E@K@8K@FE =8:KFI-2 (<$!-2) NPMRCGL RF?R @GLBQ DGPQR RM ?L K3/" RM GLGRG?RC RP?LQJ?RGML ><E< <OGI<JJ@FE NPMACQQCQ RF?R AMLRPMJ RFC RSPLGLE ML MP RSPLGLE MDD MD ? ECLC >L8E@E< ;@G?FJG?8K< ("D+) KMJCASJC RF?R GQ JCDR ?DRCP RFC CLCPEW GQ SQCB RM QR?PR RP?LQJ?RGML >L8E@E< KI@G?FJG?8K< ("/+) CLCPEW-NPMTGBGLE KMJCASJC RF?R @GLBQ RM C*'-2 ?LB GQ LCCBCB DMP RP?LQJ?RGML ?@JKFE< 8:<KPC8K@FE CNGECLCRGA KMBGDGA?RGML RF?R JC?BQ RM ECLC QGJCLAGLE; AMKKMLJW DMSLB GL A?LACP ACJJQ DMSLB GL A?LACP ACJJQ @E;L:@9C< FG<IFE MNCPML RF?R A?L @C ?ARGT?RCB MP PCNPCQQCB BCNCLBGLE ML ACJJSJ?P LCCBQ ?LB RFC QSPPMSLBGLE CLTGPMLKCLR @E@K@8K@FE :FDGC<O NPMRCGL AMKNJCV AMLR?GLGLE C*'2-2 RF?R QR?PRQ RP?LQJ?RGML C8: FG<IFE MNCPML GL NPMI?PWMRGA ACJJQ RF?R CLAMBCQ ECLCQ PCOSGPCB DMP NPMACQQGLE ?LB GLR?IC MD J?ARMQC C8I>< 60. I@9FJFD8C JL9LE@K QCAMLB, J?PECP PG@MQMK?J QS@SLGR RF?R @GLBQ RM RFC 3/" RM RP?LQJ?RC GR GLRM NPMRCGL D@:IF-)A (D@-)A) QK?JJ 3/" KMJCASJCQ (?NNPMVGK?RCJW 21 LSAJCMRGBCQ GL JCLERF) RF?R @GLB RM 3/" KMJCASJCQ RM BCEP?BC RFCK DP: MLAMECLC RF?R A?SQCQ A?LACP GL K?LW A?LACP ACJJQ E<>8K@M< I<>LC8KFI NPMRCGL RF?R NPCTCLRQ RP?LQAPGNRGML FG<I8KFI PCEGML MD %/" MSRQGBC MD RFC NPMKMRCP PCEGML RF?R @GLBQ ?ARGT?RMPQ MP PCNPCQQMPQ RF?R AMLRPMJ ECLC CVNPCQQGML GL NPMI?PWMRGA ACJJQ
#(!04%2 16 X '%.% %802%33)/.
443
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
C#A+/ - .0((A-4
16.1 -<>LC8K@FE F= "<E< OGI<JJ@FE
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
16.2 +IFB8IPFK@: "<E< -<>LC8K@FE

5FC PCESJ?RGML MD ECLC CVNPCQQGML GL NPMI?PWMRGA ACJJQ MAASPQ ?R RFC RP?LQAPGNRGML?J JCTCJ. 5FCPC ?PC RFPCC U?WQ RM AMLRPMJ RFC RP?LQAPGNRGML MD ?L MNCPML: PCNPCQQGTC AMLRPMJ, ?ARGT?RMP AMLRPMJ, ?LB GLBSAG@JC AMLRPMJ. 3CNPCQQGTC AMLRPMJ, RWNGDGCB @W RFC ;97 MNCPML, SQCQ NPMRCGLQ @MSLB RM RFC MNCP?RMP QCOSCLAC RM NFWQGA?JJW NPCTCLR RFC @GLBGLE MD 3/" NMJWKCP?QC ?LB RFC ?ARGT?RGML MD RP?LQAPGNRGML. 5FCPCDMPC, GD RPWNRMNF?L GQ LMR LCCBCB, RFC PCNPCQQMP GQ @MSLB RM RFC MNCP?RMP ?LB RP?LQAPGNRGML PCK?GLQ MDD. "ARGT?RMP AMLRPMJ, RWNGDGCB @W RFC ?ARGML MD $"1, GLAPC?QCQ RFC @GLBGLE ?@GJGRW MD 3/" NMJWKCP?QC RM RFC NPMKMRCP UFCL $"1 GQ @MSLB. *L RFGQ A?QC, JMU JCTCJQ MD EJSAMQC PCQSJR GL RFC @GLBGLE MD A".1 RM $"1. $"1 RFCL @GLBQ RFC NPMKMRCP, UFGAF ?JJMUQ 3/" NMJWKCP?QC RM @GLB RM RFC NPMKMRCP @CRRCP. *L RFC J?QR
444
#(!04%2 16 X '%.% %802%33)/.
CV?KNJCqRFC 3(* MNCPMLqRUM AMLBGRGMLQ KSQR @C KCR RM GLGRG?RC RP?LQAPGNRGML. (JSAMQC KSQR LMR @C NPCQCLR, ?LB J?ARMQC KSQR @C ?T?GJ?@JC DMP RFC 3(* MNCPML RM @C RP?LQAPG@CB. *D EJSAMQC GQ ?@QCLR, $"1 @GLBQ RM RFC MNCP?RMP. *D J?ARMQC GQ NPCQCLR, RFC PCNPCQQMP NPMRCGL BMCQ LMR @GLB RM GRQ MNCP?RMP. 0LJW UFCL @MRF AMLBGRGMLQ ?PC KCR UGJJ 3/" NMJWKCP?QC @GLB RM RFC NPMKMRCP RM GLBSAC RP?LQAPGNRGML.
16.3 LB8IPFK@: G@><E<K@: "<E< -<>LC8K@FE

*L CSI?PWMRGA ACJJQ, RFC DGPQR QR?EC MD ECLC CVNPCQQGML AMLRPMJ MAASPQ ?R RFC CNGECLCRGA JCTCJ. &NGECLCRGA KCAF?LGQKQ AMLRPMJ ?AACQQ RM RFC AFPMKMQMK?J PCEGML RM ?JJMU ECLCQ RM @C RSPLCB ML MP MDD. 5FCQC KCAF?LGQKQ AMLRPMJ FMU %/" GQ N?AICB GLRM RFC LSAJCSQ @W PCESJ?RGLE FMU RGEFRJW RFC %/" GQ UMSLB ?PMSLB FGQRMLC NPMRCGLQ. 5FC ?BBGRGML MP PCKMT?J MD AFCKGA?J KMBGDGA?RGMLQ (MP DJ?EQ) RM FGQRMLC NPMRCGLQ MP %/" QGEL?JQ RM RFC ACJJ RM MNCL MP AJMQC ? AFPMKMQMK?J PCEGML. 5FCPCDMPC, CSI?PWMRGA ACJJQ A?L AMLRPMJ UFCRFCP ? ECLC GQ CVNPCQQCB @W AMLRPMJJGLE ?AACQQG@GJGRW RM RP?LQAPGNRGML D?ARMPQ ?LB RFC @GLBGLE MD 3/" NMJWKCP?QC RM GLGRG?RC RP?LQAPGNRGML.
16.4 LB8IPFK@: /I8EJ:I@GK@FE "<E< -<>LC8K@FE

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
16.5 LB8IPFK@: +FJK-KI8EJ:I@GK@FE8C "<E< -<>LC8K@FE

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
16.6 LB8IPFK@: /I8EJC8K@FE8C 8E; +FJK-KI8EJC8K@FE8C "<E< -<>LC8K@FE

$F?LEGLE RFC QR?RSQ MD RFC 3/" MP RFC NPMRCGL GRQCJD A?L ?DDCAR RFC ?KMSLR MD NPMRCGL, RFC DSLARGML MD RFC NPMRCGL, MP FMU JMLE GR GQ DMSLB GL RFC ACJJ. 5M RP?LQJ?RC RFC NPMRCGL, ? NPMRCGL GLGRG?RMP AMKNJCV KSQR ?QQCK@JC ML RFC 3/". .MBGDGA?RGMLQ (QSAF ?Q NFMQNFMPWJ?RGML) MD NPMRCGLQ GL RFGQ AMKNJCV A?L NPCTCLR NPMNCP RP?LQJ?RGML DPMK MAASPPGLE. 0LAC ? NPMRCGL F?Q @CCL QWLRFCQGXCB, GR A?L @C KMBGDGCB (NFMQNFMPWJ?RCB, ?ACRWJ?RCB, KCRFWJ?RCB, MP S@GOSGRGL?RCB). 5FCQC NMQR-RP?LQJ?RGML?J KMBGDGA?RGMLQ A?L EPC?RJW GKN?AR RFC QR?@GJGRW, BCEP?B?RGML, MP DSLARGML MD RFC NPMRCGL.
16.7 C8E:LC8K@FE

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1. Figure 16.5 *L E. *630, RFC ;97 MNCPML GQ ML @W BCD?SJR, UFGJC RFC 3(* MNCPML GQ MDD. 8FW BM WMS RFGLI RF?R RFGQ GQ RFC A?QC 2. Figure 16.7 *L DCK?JCQ, MLC MD RFC RUM 9 AFPMKMQMKCQ GQ GL?ARGT?RCB BSPGLE CK@PWMLGA BCTCJMNKCLR @CA?SQC MD CNGECLCRGA AF?LECQ RM RFC
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AFPMK?RGL. 8F?R GKN?AR BM WMS RFGLI RFCQC AF?LECQ UMSJB F?TC ML LSAJCMQMKC N?AIGLE 3. Figure 16.13 "L GLAPC?QC GL NFMQNFMPWJ?RGML JCTCJQ MD C*'-2 F?Q @CCL M@QCPTCB GL N?RGCLRQ UGRF
LCSPMBCECLCP?RGTC BGQC?QCQ QSAF ?Q "JXFCGKCPsQ, 1?PIGLQMLsQ, ?LB )SLRGLERMLsQ. 8F?R GKN?AR BM WMS RFGLI RFGQ KGEFR F?TC ML NPMRCGL QWLRFCQGQ
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4. $MLRPMJ MD ECLC CVNPCQQGML GL CSI?PWMRGA ACJJQ MAASPQ ?R UFGAF JCTCJ(Q) >. MLJW RFC RP?LQAPGNRGML?J JCTCJ ?. CNGECLCRGA ?LB RP?LQAPGNRGML?J JCTCJQ @. CNGECLCRGA, RP?LQAPGNRGML?J, ?LB RP?LQJ?RGML?J JCTCJQ A. CNGECLCRGA, RP?LQAPGNRGML?J, NMQRRP?LQAPGNRGML?J, RP?LQJ?RGML?J, ?LB NMQRRP?LQJ?RGML?J JCTCJQ 5. 1MQR-RP?LQJ?RGML?J AMLRPMJ PCDCPQ RM: >. PCESJ?RGML MD ECLC CVNPCQQGML ?DRCP RP?LQAPGNRGML ?. PCESJ?RGML MD ECLC CVNPCQQGML ?DRCP RP?LQJ?RGML @. AMLRPMJ MD CNGECLCRGA ?ARGT?RGML A. NCPGMB @CRUCCL RP?LQAPGNRGML ?LB RP?LQJ?RGML 6. *D EJSAMQC GQ ?@QCLR, @SR QM GQ J?ARMQC, RFC 3(* MNCPML UGJJ @C >>>>>>>>. >. ?ARGT?RCB ?. PCNPCQQCB @. ?ARGT?RCB, @SR MLJW N?PRG?JJW A. KSR?RCB 7. 1PMI?PWMRGA ACJJQ J?AI ? LSAJCSQ. 5FCPCDMPC, RFC ECLCQ GL NPMI?PWMRGA ACJJQ ?PC: >. ?JJ CVNPCQQCB, ?JJ MD RFC RGKC ?. RP?LQAPG@CB ?LB RP?LQJ?RCB ?JKMQR QGKSJR?LCMSQJW @. RP?LQAPGNRGML?JJW AMLRPMJJCB @CA?SQC RP?LQJ?RGML @CEGLQ @CDMPC RP?LQAPGNRGML CLBQ A. @ ?LB A ?PC @MRF RPSC 8. 8F?R ?PC CNGECLCRGA KMBGDGA?RGMLQ >. RFC ?BBGRGML MD PCTCPQG@JC AF?LECQ RM FGQRMLC NPMRCGLQ ?LB %/" ?. RFC PCKMT?J MD LSAJCMQMKCQ DPMK RFC %/" @. RFC ?BBGRGML MD KMPC LSAJCMQMKCQ RM RFC %/" A. KSR?RGML MD RFC %/" QCOSCLAC 9. 8FGAF MD RFC DMJJMUGLE ?PC RPSC MD CNGECLCRGA AF?LECQ >. ?JJMU %/" RM @C RP?LQAPG@CB ?. KMTC FGQRMLCQ RM MNCL MP AJMQC ? AFPMKMQMK?J PCEGML @. ?PC RCKNMP?PW A. ?JJ MD RFC ?@MTC 10. 5FC @GLBGLE MD >>>>>>>> GQ PCOSGPCB DMP RP?LQAPGNRGML RM QR?PR. >. ? NPMRCGL ?. %/" NMJWKCP?QC @. 3/" NMJWKCP?QC A. ? RP?LQAPGNRGML D?ARMP 11. 8F?R UGJJ PCQSJR DPMK RFC @GLBGLE MD ? RP?LQAPGNRGML D?ARMP RM ?L CLF?LACP PCEGML >. BCAPC?QCB RP?LQAPGNRGML MD ?L ?BH?ACLR ECLC ?. GLAPC?QCB RP?LQAPGNRGML MD ? BGQR?LR ECLC @. ?JRCP?RGML MD RFC RP?LQJ?RGML MD ?L ?BH?ACLR ECLC A. GLGRG?RGML MD RFC PCAPSGRKCLR MD 3/" NMJWKCP?QC 12. 8FGAF MD RFC DMJJMUGLE ?PC GLTMJTCB GL NMQRRP?LQAPGNRGML?J AMLRPMJ >. AMLRPMJ MD 3/" QNJGAGLE ?. AMLRPMJ MD 3/" QFSRRJGLE @. AMLRPMJ MD 3/" QR?@GJGRW A. ?JJ MD RFC ?@MTC 13. #GLBGLE MD ?L 3/" @GLBGLE NPMRCGL UGJJ >>>>>>>> RFC QR?@GJGRW MD RFC 3/" KMJCASJC. >. ?. @. A. GLAPC?QC BCAPC?QC LCGRFCP GLAPC?QC LMP BCAPC?QC CGRFCP GLAPC?QC MP BCAPC?QC
14. 1MQR-RP?LQJ?RGML?J KMBGDGA?RGMLQ MD NPMRCGLQ A?L ?DDCAR UFGAF MD RFC DMJJMUGLE >. NPMRCGL DSLARGML ?. RP?LQAPGNRGML?J PCESJ?RGML @. AFPMK?RGL KMBGDGA?RGML A. ?JJ MD RFC ?@MTC 15. $?LACP A?SQGLE ECLCQ ?PC A?JJCB >>>>>>>>. >. ?. @. A. RP?LQDMPK?RGML ECLCQ RSKMP QSNNPCQQMP ECLCQ MLAMECLCQ KSR?RCB ECLCQ
16. 5?PECRCB RFCP?NGCQ ?PC SQCB GL N?RGCLRQ UGRF ? QCR ECLC CVNPCQQGML N?RRCPL. " R?PECRCB RFCP?NW RF?R NPCTCLRQ RFC ?ARGT?RGML MD RFC CQRPMECL PCACNRMP GL @PC?QR A?LACP UMSJB @C @CLCDGAG?J RM UFGAF RWNC MD N?RGCLR >. N?RGCLRQ UFM CVNPCQQ RFC &('3 PCACNRMP GL LMPK?J ACJJQ ?. N?RGCLRQ UGRF ? KSR?RGML RF?R GL?ARGT?RCQ RFC CQRPMECL PCACNRMP @. N?RGCLRQ UGRF JMRQ MD RFC CQRPMECL PCACNRMP CVNPCQQCB GL RFCGP RSKMP A. N?RGCLRQ RF?R F?TC LM CQRPMECL PCACNRMP CVNPCQQCB GL RFCGP RSKMP
446
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17. /?KC RUM BGDDCPCLACQ @CRUCCL NPMI?PWMRGA ?LB CSI?PWMRGA ACJJQ ?LB FMU RFCQC BGDDCPCLACQ @CLCDGR KSJRGACJJSJ?P MPE?LGQKQ. 18. %CQAPG@C FMU AMLRPMJJGLE ECLC CVNPCQQGML UGJJ ?JRCP RFC MTCP?JJ NPMRCGL JCTCJQ GL RFC ACJJ. 19. %CQAPG@C FMU RP?LQAPGNRGML GL NPMI?PWMRGA ACJJQ A?L @C ?JRCPCB @W CVRCPL?J QRGKSJ?RGML QSAF ?Q CVACQQ J?ARMQC GL RFC CLTGPMLKCLR. 20. 8F?R GQ RFC BGDDCPCLAC @CRUCCL ? PCNPCQQG@JC ?LB ?L GLBSAG@JC MNCPML 21. *L A?LACP ACJJQ, ?JRCP?RGML RM CNGECLCRGA KMBGDGA?RGMLQ RSPLQ MDD ECLCQ RF?R ?PC LMPK?JJW CVNPCQQCB. )WNMRFCRGA?JJW, FMU AMSJB WMS PCTCPQC RFGQ NPMACQQ RM RSPL RFCQC ECLCQ @?AI ML 22. " KSR?RGML UGRFGL RFC NPMKMRCP PCEGML A?L ?JRCP RP?LQAPGNRGML MD ? ECLC. %CQAPG@C FMU RFGQ A?L F?NNCL. 23. 8F?R AMSJB F?NNCL GD ? ACJJ F?B RMM KSAF MD ?L ?ARGT?RGLE RP?LQAPGNRGML D?ARMP NPCQCLR 24. %CQAPG@C FMU 3#1Q A?L NPCTCLR KG3/"Q DPMK BCEP?BGLE ?L 3/" KMJCASJC. 25. )MU A?L CVRCPL?J QRGKSJG ?JRCP NMQRRP?LQAPGNRGML?J AMLRPMJ MD ECLC CVNPCQQGML 26. 1PMRCGL KMBGDGA?RGML A?L ?JRCP ECLC CVNPCQQGML GL K?LW U?WQ. %CQAPG@C FMU NFMQNFMPWJ?RGML MD NPMRCGLQ A?L ?JRCP ECLC CVNPCQQGML. 27. "JRCPL?RGTC DMPKQ MD ? NPMRCGL A?L @C @CLCDGAG?J MP F?PKDSJ RM ? ACJJ. 8F?R BM WMS RFGLI UMSJB F?NNCL GD RMM KSAF MD ?L ?JRCPL?RGTC NPMRCGL @MSLB RM RFC 3' 653 MD ?L 3/" ?LB A?SQCB GR RM BCEP?BC 28. $F?LECQ GL CNGECLCRGA KMBGDGA?RGMLQ ?JRCP RFC ?AACQQG@GJGRW ?LB RP?LQAPGNRGML MD %/". %CQAPG@C FMU CLTGPMLKCLR?J QRGKSJG, QSAF ?Q SJRP?TGMJCR JGEFR CVNMQSPC, AMSJB KMBGDW ECLC CVNPCQQGML. 29. /CU BPSEQ ?PC @CGLE BCTCJMNCB RF?R BCAPC?QC %/" KCRFWJ?RGML ?LB NPCTCLR RFC PCKMT?J MD ?ACRWJ EPMSNQ DPMK FGQRMLC NPMRCGLQ. &VNJ?GL FMU RFCQC BPSEQ AMSJB ?DDCAR ECLC CVNPCQQGML RM FCJN IGJJ RSKMP ACJJQ. 30. )MU A?L SLBCPQR?LBGLE RFC ECLC CVNPCQQGML N?RRCPL GL ? A?LACP ACJJ RCJJ WMS QMKCRFGLE ?@MSR RF?R QNCAGDGA DMPK MD A?LACP
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17.1 R B@FK<:?EFCF>P
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448
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472
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16. 4COSCLAGLE ?L GLBGTGBS?J NCPQMLsQ ECLMKC >. GQ ASPPCLRJW NMQQG@JC ?. AMSJB JC?B RM JCE?J GQQSCQ PCE?PBGLE BGQAPGKGL?RGML ?LB NPGT?AW @. AMSJB FCJN K?IC GLDMPKCB AFMGACQ ?@MSR KCBGA?J RPC?RKCLR A. ?JJ MD RFC ?@MTC 17. 8F?R GQ RFC KMQR AF?JJCLEGLE GQQSC D?AGLE ECLMKC QCOSCLAGLE >. RFC GL?@GJGRW RM BCTCJMN D?QR ?LB ?AASP?RC QCOSCLAGLE RCAFLGOSCQ ?. RFC CRFGAQ MD SQGLE GLDMPK?RGML DPMK ECLMKCQ ?R RFC GLBGTGBS?J JCTCJ @. RFC ?T?GJ?@GJGRW ?LB QR?@GJGRW MD %/" A. ?JJ MD RFC ?@MTC 18. (CLMKGAQ A?L @C SQCB GL ?EPGASJRSPC RM: >. ?. @. A. ECLCP?RC LCU FW@PGB QRP?GLQ GKNPMTC BGQC?QC PCQGQR?LAC GKNPMTC WGCJB ?JJ MD RFC ?@MTC
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30. )MU F?Q NPMRCMKGAQ @CCL SQCB GL A?LACP BCRCARGML ?LB RPC?RKCLR
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8?JJ?AC ?LB %?PUGL @MRF M@QCPTCB QGKGJ?P N?RRCPLQ GL MRFCP MPE?LGQKQ ?LB RFCW GLBCNCLBCLRJW BCTCJMNCB RFC Q?KC CVNJ?L?RGML DMP FMU ?LB UFW QSAF AF?LECQ AMSJB R?IC NJ?AC. %?PUGL A?JJCB RFGQ KCAF?LGQK L?RSP?J QCJCARGML. Natural selection, ?JQM ILMUL ?Q tQSPTGT?J MD RFC DGRRCQR,u GQ RFC KMPC NPMJGDGA PCNPMBSARGML MD GLBGTGBS?JQ UGRF D?TMP?@JC RP?GRQ RF?R QSPTGTC CLTGPMLKCLR?J AF?LEC @CA?SQC MD RFMQC RP?GRQ; RFGQ JC?BQ RM CTMJSRGML?PW AF?LEC. 'MP CV?KNJC, ? NMNSJ?RGML MD EG?LR RMPRMGQCQ DMSLB GL RFC (?J?N?EMQ "PAFGNCJ?EM U?Q M@QCPTCB @W %?PUGL RM F?TC JMLECP LCAIQ RF?L RFMQC RF?R JGTCB ML MRFCP GQJ?LBQ UGRF BPW JMUJ?LBQ. 5FCQC RMPRMGQCQ UCPC tQCJCARCBu @CA?SQC RFCW AMSJB PC?AF KMPC JC?TCQ ?LB ?AACQQ KMPC DMMB RF?L RFMQC UGRF QFMPR LCAIQ. *L RGKCQ MD BPMSEFR UFCL DCUCP JC?TCQ UMSJB @C ?T?GJ?@JC, RFMQC RF?R AMSJB PC?AF KMPC JC?TCQ F?B ? @CRRCP AF?LAC RM C?R ?LB QSPTGTC RF?L RFMQC RF?R AMSJBLs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s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igure 18.3) NPCQCLRGLE RFC GBC? MD L?RSP?J QCJCARGML UCPC PC?B RMECRFCP GL 1858 @CDMPC RFC -GLLC?L 4MAGCRW GL -MLBML. 5FC DMJJMUGLE WC?P %?PUGLsQ @MMI, O5 ;/, O90.05 6- 7,*0,:, U?Q NS@JGQFCB. )GQ @MMI MSRJGLCB GL AMLQGBCP?@JC BCR?GJ FGQ ?PESKCLRQ DMP CTMJSRGML @W L?RSP?J QCJCARGML.
480
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%CKMLQRP?RGMLQ MD CTMJSRGML @W L?RSP?J QCJCARGML ?PC RGKC AMLQSKGLE ?LB BGDDGASJR RM M@R?GL. 0LC MD RFC @CQR CV?KNJCQ F?Q @CCL BCKMLQRP?RCB GL RFC TCPW @GPBQ RF?R FCJNCB RM GLQNGPC %?PUGLsQ RFCMPW: RFC (?JaN?EMQ DGLAFCQ. 1CRCP ?LB 3MQCK?PW (P?LR ?LB RFCGP AMJJC?ESCQ F?TC QRSBGCB (?JaN?EMQ DGLAF NMNSJ?RGMLQ CTCPW WC?P QGLAC 1976 ?LB F?TC NPMTGBCB GKNMPR?LR BCKMLQRP?RGMLQ MD L?RSP?J QCJCARGML. 5FC (P?LRQ DMSLB AF?LECQ DPMK MLC ECLCP?RGML RM RFC LCVR GL RFC BGQRPG@SRGML MD @C?I QF?NCQ UGRF RFC KCBGSK EPMSLB DGLAF ML RFC (?JaN?EMQ GQJ?LB MD %?NFLC .?HMP. 5FC @GPBQ F?TC GLFCPGRCB T?PG?RGML GL RFC @GJJ QF?NC UGRF QMKC @GPBQ F?TGLE UGBC BCCN @GJJQ ?LB MRFCPQ F?TGLE RFGLLCP @GJJQ. %SPGLE ? NCPGMB GL UFGAF P?GLD?JJ U?Q FGEFCP RF?L LMPK?J @CA?SQC MD ?L &J /GcM, RFC J?PEC F?PB QCCBQ RF?R J?PEC-@GJJCB @GPBQ ?RC UCPC PCBSACB GL LSK@CP; FMUCTCP, RFCPC U?Q ?L ?@SLB?LAC MD RFC QK?JJ QMDR QCCBQ UFGAF RFC QK?JJ-@GJJCB @GPBQ ?RC. 5FCPCDMPC, QSPTGT?J ?LB PCNPMBSARGML UCPC KSAF @CRRCP GL RFC DMJJMUGLE WC?PQ DMP RFC QK?JJ@GJJCB @GPBQ. *L RFC WC?PQ DMJJMUGLE RFGQ &J /GcM, RFC (P?LRQ KC?QSPCB @C?I QGXCQ GL RFC NMNSJ?RGML ?LB DMSLB RF?R RFC ?TCP?EC @GJJ QGXC U?Q QK?JJCP. 4GLAC @GJJ QGXC GQ ?L GLFCPGRCB RP?GR, N?PCLRQ UGRF QK?JJCP @GJJQ F?B KMPC MDDQNPGLE ?LB RFC QGXC MD @GJJQ F?B CTMJTCB RM @C QK?JJCP. "Q AMLBGRGMLQ GKNPMTCB GL 1987 ?LB J?PECP QCCBQ @CA?KC KMPC ?T?GJ?@JC, RFC RPCLB RMU?PB QK?JJCP ?TCP?EC @GJJ QGXC AC?QCB.
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!@>LI< 18.4 ! CFBIA ?FLILDFPQ QO>KNRFIFWBP > MLI>O ?B>O CLO PQRAV. (@OBAFQ: +>OBK 2ELAB)
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
+IF:<JJ<J 8E; +8KK<IEJ F= MFCLK@FE

/?RSP?J QCJCARGML A?L MLJW R?IC NJ?AC GD RFCPC GQ variation, MP BGDDCPCLACQ, ?KMLE GLBGTGBS?JQ GL ? NMNSJ?RGML. *KNMPR?LRJW, RFCQC BGDDCPCLACQ KSQR F?TC QMKC ECLCRGA @?QGQ; MRFCPUGQC, RFC QCJCARGML UGJJ LMR JC?B RM AF?LEC GL RFC LCVR ECLCP?RGML. 5FGQ GQ APGRGA?J @CA?SQC T?PG?RGML ?KMLE GLBGTGBS?JQ A?L @C A?SQCB @W LML-ECLCRGA PC?QMLQ QSAF ?Q ?L GLBGTGBS?J @CGLE R?JJCP @CA?SQC MD @CRRCP LSRPGRGML P?RFCP RF?L BGDDCPCLR ECLCQ. (CLCRGA BGTCPQGRW GL ? NMNSJ?RGML AMKCQ DPMK RUM K?GL KCAF?LGQKQ: KSR?RGML ?LB QCVS?J PCNPMBSARGML. .SR?RGML, ? AF?LEC GL %/", GQ RFC SJRGK?RC QMSPAC MD LCU ?JJCJCQ, MP LCU ECLCRGA T?PG?RGML GL ?LW NMNSJ?RGML. 5FC ECLCRGA AF?LECQ A?SQCB @W KSR?RGML A?L F?TC MLC MD RFPCC MSRAMKCQ ML RFC NFCLMRWNC. " KSR?RGML ?DDCARQ RFC NFCLMRWNC MD RFC MPE?LGQK GL ? U?W RF?R EGTCQ GR PCBSACB DGRLCQQqJMUCP JGICJGFMMB MD QSPTGT?J MP DCUCP MDDQNPGLE. " KSR?RGML K?W NPMBSAC ? NFCLMRWNC UGRF ? @CLCDGAG?J CDDCAR ML DGRLCQQ. "LB, K?LW KSR?RGMLQ UGJJ ?JQM F?TC LM CDDCAR ML RFC DGRLCQQ MD RFC NFCLMRWNC; RFCQC ?PC A?JJCB LCSRP?J KSR?RGMLQ. .SR?RGMLQ K?W ?JQM F?TC ? UFMJC P?LEC MD CDDCAR QGXCQ ML RFC DGRLCQQ MD RFC MPE?LGQK RF?R CVNPCQQCQ RFCK GL RFCGP NFCLMRWNC, DPMK ? QK?JJ CDDCAR RM ? EPC?R CDDCAR. 4CVS?J PCNPMBSARGML ?JQM JC?BQ RM ECLCRGA BGTCPQGRW: UFCL RUM N?PCLRQ PCNPMBSAC, SLGOSC AMK@GL?RGMLQ MD ?JJCJCQ ?QQCK@JC RM NPMBSAC RFC SLGOSC ECLMRWNCQ ?LB RFSQ NFCLMRWNCQ GL C?AF MD RFC MDDQNPGLE. " FCPGR?@JC RP?GR RF?R FCJNQ RFC QSPTGT?J ?LB PCNPMBSARGML MD ?L MPE?LGQK GL GRQ NPCQCLR CLTGPMLKCLR GQ A?JJCB ?L adaptation. 4AGCLRGQRQ BCQAPG@C EPMSNQ MD MPE?LGQKQ @CAMKGLE ?B?NRCB RM RFCGP CLTGPMLKCLR UFCL ? AF?LEC GL RFC P?LEC MD ECLCRGA T?PG?RGML MAASPQ MTCP RGKC RF?R GLAPC?QCQ MP K?GLR?GLQ RFC tDGRu MD RFC NMNSJ?RGML RM GRQ CLTGPMLKCLR. 5FC UC@@CB DCCR MD NJ?RWNSQCQ ?PC ?L ?B?NR?RGML DMP QUGKKGLE. 5FC QLMU JCMN?PBQs RFGAI DSP GQ ?L ?B?NR?RGML DMP JGTGLE GL RFC AMJB. 5FC AFCCR?FQs D?QR QNCCB GQ ?L ?B?NR?RGML DMP A?RAFGLE NPCW.
482
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8FCRFCP MP LMR ? RP?GR GQ D?TMP?@JC BCNCLBQ ML RFC CLTGPMLKCLR?J AMLBGRGMLQ ?R RFC RGKC. 5FC Q?KC RP?GRQ ?PC LMR ?JU?WQ QCJCARCB @CA?SQC CLTGPMLKCLR?J AMLBGRGMLQ A?L AF?LEC. 'MP CV?KNJC, AMLQGBCP ? QNCAGCQ MD NJ?LR RF?R EPCU GL ? KMGQR AJGK?RC ?LB BGB LMR LCCB RM AMLQCPTC U?RCP. -?PEC JC?TCQ UCPC QCJCARCB @CA?SQC RFCW ?JJMUCB RFC NJ?LR RM M@R?GL KMPC CLCPEW DPMK RFC QSL. -?PEC JC?TCQ PCOSGPC KMPC U?RCP RM K?GLR?GL RF?L QK?JJ JC?TCQ, ?LB RFC KMGQR CLTGPMLKCLR NPMTGBCB D?TMP?@JC AMLBGRGMLQ RM QSNNMPR J?PEC JC?TCQ. "DRCP RFMSQ?LBQ MD WC?PQ, RFC AJGK?RC AF?LECB, ?LB RFC ?PC? LM JMLECP F?B CVACQQ U?RCP. 5FC BGPCARGML MD L?RSP?J QCJCARGML QFGDRCB QM RF?R NJ?LRQ UGRF QK?JJ JC?TCQ UCPC QCJCARCB @CA?SQC RFMQC NMNSJ?RGMLQ UCPC ?@JC RM AMLQCPTC U?RCP RM QSPTGTC RFC LCU CLTGPMLKCLR?J AMLBGRGMLQ. 5FC CTMJSRGML MD QNCAGCQ F?Q PCQSJRCB GL CLMPKMSQ T?PG?RGML GL DMPK ?LB DSLARGML. 4MKCRGKCQ, CTMJSRGML EGTCQ PGQC RM EPMSNQ MD MPE?LGQKQ RF?R @CAMKC RPCKCLBMSQJW BGDDCPCLR DPMK C?AF MRFCP. 8FCL RUM QNCAGCQ CTMJTC GL BGTCPQC BGPCARGMLQ DPMK ? AMKKML NMGLR, GR GQ A?JJCB divergent evolution. 4SAF BGTCPECLR CTMJSRGML A?L @C QCCL GL RFC DMPKQ MD RFC PCNPMBSARGTC MPE?LQ MD DJMUCPGLE NJ?LRQ UFGAF QF?PC RFC Q?KC @?QGA ?L?RMKGCQ; FMUCTCP, RFCW A?L JMMI TCPW BGDDCPCLR ?Q ? PCQSJR MD QCJCARGML GL BGDDCPCLR NFWQGA?J CLTGPMLKCLRQ ?LB ?B?NR?RGML RM BGDDCPCLR IGLBQ MD NMJJGL?RMPQ (Figure 18.5).
!@>LI< 18.5 &ILTBOFKD MI>KQP BSLISBA COLJ > @LJJLK >K@BPQLO. .LQF@B QE>Q QEB (>) ABKPB ?I>WFKD PQ>O (L0(;9<: :70*(;() >KA QEB (?) MROMIB @LKBCILTBO (E*/05(*,( 7<97<9,() S>OV FK >MMB>O>K@B, VBQ ?LQE PE>OB > PFJFI>O ?>PF@ JLOMELILDV. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V $OBT !SBOV; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V #LOV :>KHBO)
*L MRFCP A?QCQ, QGKGJ?P NFCLMRWNCQ CTMJTC GLBCNCLBCLRJW GL BGQR?LRJW PCJ?RCB QNCAGCQ. 'MP CV?KNJC, DJGEFR F?Q CTMJTCB GL @MRF @?RQ ?LB GLQCARQ, ?LB RFCW @MRF F?TC QRPSARSPCQ UC PCDCP RM ?Q UGLEQ, UFGAF ?PC ?B?NR?RGMLQ RM DJGEFR. )MUCTCP, RFC UGLEQ MD @?RQ ?LB GLQCARQ F?TC CTMJTCB DPMK TCPW BGDDCPCLR MPGEGL?J QRPSARSPCQ. 5FGQ NFCLMKCLML GQ A?JJCB convergent evolution, UFCPC QGKGJ?P RP?GRQ CTMJTC GLBCNCLBCLRJW GL QNCAGCQ RF?R BM LMR QF?PC ? PCACLR AMKKML ?LACQRPW. 5FC RUM QNCAGCQ A?KC RM RFC Q?KC DSLARGML, DJWGLE, @SR BGB QM QCN?P?RCJW DPMK C?AF MRFCP. 5FCQC NFWQGA?J AF?LECQ MAASP MTCP CLMPKMSQ QN?LQ MD RGKC ?LB FCJN CVNJ?GL FMU CTMJSRGML MAASPQ. /?RSP?J QCJCARGML ?ARQ ML GLBGTGBS?J MPE?LGQKQ, UFGAF GL RSPL A?L QF?NC ?L CLRGPC QNCAGCQ. "JRFMSEF L?RSP?J QCJCARGML K?W UMPI GL ? QGLEJC ECLCP?RGML ML ?L GLBGTGBS?J, GR A?L R?IC RFMSQ?LBQ MP CTCL KGJJGMLQ MD WC?PQ DMP RFC ECLMRWNC MD ?L CLRGPC QNCAGCQ RM CTMJTC. *R GQ MTCP RFCQC J?PEC RGKC QN?LQ RF?R JGDC ML C?PRF F?Q AF?LECB ?LB AMLRGLSCQ RM AF?LEC.
M@;<E:< F= MFCLK@FE
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igure 18.6). 'MP CV?KNJC, QAGCLRGQRQ F?TC PCAMTCPCB FGEFJW BCR?GJCB PCAMPBQ QFMUGLE RFC CTMJSRGML MD FSK?LQ ?LB FMPQCQ (Figure 18.6). 5FC UF?JC DJGNNCP QF?PCQ ? QGKGJ?P KMPNFMJMEW RM ?NNCLB?ECQ MD @GPBQ ?LB K?KK?JQ (Figure 18.7) GLBGA?RGLE RF?R RFCQC QNCAGCQ QF?PC ? AMKKML ?LACQRMP.
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483
!@>LI< 18.6 )K QEFP (>) AFPMI>V, CLPPFI ELJFKFAP >OB >OO>KDBA COLJ LIABPQ (?LQQLJ) QL KBTBPQ (QLM). !P ELJFKFAP BSLISBA, QEB PE>MB LC QEB PHRII @E>KDBA. !K >OQFPQwP OBKAFQFLK LC (?) BUQFK@Q PMB@FBP LC QEB DBKRP E8<<: OBSB>IP QE>Q QEBPB >K@FBKQ PMB@FBP OBPBJ?IBA QEB JLABOK ELOPB (E8<<: -,9<:) ?RQ S>OFBA FK PFWB.
A3'942> '3* E2(7>414-> "LMRFCP RWNC MD CTGBCLAC DMP CTMJSRGML GQ RFC NPCQCLAC MD QRPSARSPCQ GL MPE?LGQKQ RF?R QF?PC RFC Q?KC @?QGA DMPK. 'MP CV?KNJC, RFC @MLCQ GL RFC ?NNCLB?ECQ MD ? FSK?L, BME, @GPB, ?LB UF?JC ?JJ QF?PC RFC Q?KC MTCP?JJ AMLQRPSARGML (Figure 18.7) PCQSJRGLE DPMK RFCGP MPGEGL GL RFC ?NNCLB?ECQ MD ? AMKKML ?LACQRMP. 0TCP RGKC, CTMJSRGML JCB RM AF?LECQ GL RFC QF?NCQ ?LB QGXCQ MD RFCQC @MLCQ GL BGDDCPCLR QNCAGCQ, @SR RFCW F?TC K?GLR?GLCB RFC Q?KC MTCP?JJ J?WMSR. 4AGCLRGQRQ A?JJ RFCQC QWLMLWKMSQ N?PRQ homologous structures.
!@>LI< 18.7 4EB PFJFI>O @LKPQOR@QFLK LC QEBPB >MMBKA>DBP FKAF@>QBP QE>Q QEBPB LOD>KFPJP PE>OB > @LJJLK >K@BPQLO.
4MKC QRPSARSPCQ CVGQR GL MPE?LGQKQ RF?R F?TC LM ?NN?PCLR DSLARGML ?R ?JJ, ?LB ?NNC?P RM @C PCQGBS?J N?PRQ DPMK ? N?QR AMKKML ?LACQRMP. 5FCQC SLSQCB QRPSARSPCQ UGRFMSR DSLARGML ?PC A?JJCB vestigial structures. 0RFCP CV?KNJCQ MD TCQRGEG?J QRPSARSPCQ ?PC UGLEQ ML DJGEFRJCQQ @GPBQ, JC?TCQ ML QMKC A?ARG, ?LB FGLB JCE @MLCQ GL UF?JCQ.
7GQGR RFGQ interactive site (http://openstaxcollege.org/l/bone_structures) RM ESCQQ UFGAF @MLCQ QRPSARSPCQ ?PC FMKMJMEMSQ ?LB UFGAF ?PC ?L?JMEMSQ, ?LB QCC CV?KNJCQ MD CTMJSRGML?PW ?B?NR?RGMLQ RM GJJSQRP?RC RFCQC AMLACNRQ. "LMRFCP CTGBCLAC MD CTMJSRGML GQ RFC AMLTCPECLAC MD DMPK GL MPE?LGQKQ RF?R QF?PC QGKGJ?P CLTGPMLKCLRQ. 'MP CV?KNJC, QNCAGCQ MD SLPCJ?RCB ?LGK?JQ, QSAF ?Q RFC ?PARGA DMV ?LB NR?PKGE?L, JGTGLE
484
#(!04%2 18 X %6/,54)/. !.$ 4(% /2)'). /& 30%#)%3
GL RFC ?PARGA PCEGML F?TC @CCL QCJCARCB DMP QC?QML?J UFGRC NFCLMRWNCQ BSPGLE UGLRCP RM @JCLB UGRF RFC QLMU ?LB GAC (Figure 18.8ab). 5FCQC QGKGJ?PGRGCQ MAASP LMR @CA?SQC MD AMKKML ?LACQRPW, @SR @CA?SQC MD QGKGJ?P QCJCARGML NPCQQSPCQqRFC @CLCDGRQ MD LMR @CGLE QCCL @W NPCB?RMPQ.
!@>LI< 18.8 4EB TEFQB TFKQBO @L>Q LC QEB (>) >O@QF@ CLU >KA QEB (?) MQ>OJFD>KwP MIRJ>DB >OB >A>MQ>QFLKP QL QEBFO BKSFOLKJBKQP. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V +BFQE -LOBELRPB)
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"JRFMSEF RFC RFCMPW MD CTMJSRGML ECLCP?RCB QMKC AMLRPMTCPQW UFCL GR U?Q DGPQR NPMNMQCB, GR U?Q ?JKMQR SLGTCPQ?JJW ?AACNRCB @W @GMJMEGQRQ, N?PRGASJ?PJW WMSLECP @GMJMEGQRQ, UGRFGL 20 WC?PQ ?DRCP NS@JGA?RGML MD O5 ;/, O90.05 6- 7,*0,:. /CTCPRFCJCQQ, RFC RFCMPW MD CTMJSRGML GQ ? BGDDGASJR AMLACNR ?LB KGQAMLACNRGMLQ ?@MSR FMU GR UMPIQ ?@MSLB.
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1MNSJ?RGMLQ MD QNCAGCQ QF?PC ? ECLC NMMJ: ? AMJJCARGML MD ?JJ RFC T?PG?LRQ MD ECLCQ GL RFC QNCAGCQ. "E?GL, RFC @?QGQ RM ?LW AF?LECQ GL ? EPMSN MP NMNSJ?RGML MD MPE?LGQKQ KSQR @C ECLCRGA DMP RFGQ GQ RFC MLJW U?W RM QF?PC ?LB N?QQ ML RP?GRQ. 8FCL T?PG?RGMLQ MAASP UGRFGL ? QNCAGCQ, RFCW A?L MLJW @C N?QQCB RM RFC LCVR ECLCP?RGML ?JMLE RUM K?GL N?RFU?WQ: ?QCVS?J PCNPMBSARGML MP QCVS?J PCNPMBSARGML. 5FC AF?LEC UGJJ @C N?QQCB ML ?QCVS?JJW QGKNJW GD RFC PCNPMBSAGLE ACJJ NMQQCQQCQ RFC AF?LECB RP?GR. 'MP RFC AF?LECB RP?GR RM @C N?QQCB ML @W QCVS?J PCNPMBSARGML, ? E?KCRC, QSAF ?Q ? QNCPK MP CEE ACJJ, KSQR NMQQCQQ RFC AF?LECB RP?GR. *L MRFCP UMPBQ, QCVS?JJW-PCNPMBSAGLE MPE?LGQKQ A?L CVNCPGCLAC QCTCP?J ECLCRGA AF?LECQ GL RFCGP @MBW ACJJQ, @SR GD RFCQC AF?LECQ BM LMR MAASP GL ? QNCPK MP CEE ACJJ, RFC AF?LECB RP?GR UGJJ LCTCP PC?AF RFC LCVR ECLCP?RGML. 0LJW FCPGR?@JC RP?GRQ A?L CTMJTC. 5FCPCDMPC, PCNPMBSARGML NJ?WQ ? N?P?KMSLR PMJC DMP ECLCRGA AF?LEC RM R?IC PMMR GL ? NMNSJ?RGML MP QNCAGCQ. *L QFMPR, MPE?LGQKQ KSQR @C ?@JC RM PCNPMBSAC UGRF C?AF MRFCP RM N?QQ LCU RP?GRQ RM MDDQNPGLE.
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speciation: RFC DMPK?RGML MD RUM QNCAGCQ DPMK MLC MPGEGL?J QNCAGCQ. %?PUGL CLTGQGMLCB RFGQ NPMACQQ ?Q ? @P?LAFGLE CTCLR ?LB BG?EP?KKCB RFC NPMACQQ GL RFC MLJW GJJSQRP?RGML DMSLB GL O5 ;/, O90.05 6- 7,*0,: (Figure 18.11a). $MKN?PC RFGQ GJJSQRP?RGML RM RFC BG?EP?K MD CJCNF?LR CTMJSRGML (Figure 18.11b), UFGAF QFMUQ RF?R ?Q MLC
488
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'MP QNCAG?RGML RM MAASP, RUM LCU NMNSJ?RGMLQ KSQR @C DMPKCB DPMK MLC MPGEGL?J NMNSJ?RGML ?LB RFCW KSQR CTMJTC GL QSAF ? U?W RF?R GR @CAMKCQ GKNMQQG@JC DMP GLBGTGBS?JQ DPMK RFC RUM LCU NMNSJ?RGMLQ RM GLRCP@PCCB. #GMJMEGQRQ F?TC NPMNMQCB KCAF?LGQKQ @W UFGAF RFGQ AMSJB MAASP RF?R D?JJ GLRM RUM @PM?B A?RCEMPGCQ. Allopatric speciation (?JJM- = "MRFCP"; -N?RPGA = "FMKCJ?LB") GLTMJTCQ ECMEP?NFGA QCN?P?RGML MD NMNSJ?RGMLQ DPMK ? N?PCLR QNCAGCQ ?LB QS@QCOSCLR CTMJSRGML. Sympatric speciation (QWK- = "Q?KC"; -N?RPGA = "FMKCJ?LB") GLTMJTCQ QNCAG?RGML MAASPPGLE UGRFGL ? N?PCLR QNCAGCQ PCK?GLGLE GL MLC JMA?RGML. #GMJMEGQRQ RFGLI MD QNCAG?RGML CTCLRQ ?Q RFC QNJGRRGLE MD MLC ?LACQRP?J QNCAGCQ GLRM RUM BCQACLB?LR QNCAGCQ. 5FCPC GQ LM PC?QML UFW RFCPC KGEFR LMR @C KMPC RF?L RUM QNCAGCQ DMPKCB ?R MLC RGKC CVACNR RF?R GR GQ JCQQ JGICJW ?LB KSJRGNJC CTCLRQ A?L @C AMLACNRS?JGXCB ?Q QGLEJC QNJGRQ MAASPPGLE AJMQC GL RGKC.
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" ECMEP?NFGA?JJW AMLRGLSMSQ NMNSJ?RGML F?Q ? ECLC NMMJ RF?R GQ PCJ?RGTCJW FMKMECLCMSQ. (CLC DJMU, RFC KMTCKCLR MD ?JJCJCQ ?APMQQ RFC P?LEC MD RFC QNCAGCQ, GQ PCJ?RGTCJW DPCC @CA?SQC GLBGTGBS?JQ A?L KMTC ?LB RFCL K?RC UGRF GLBGTGBS?JQ GL RFCGP LCU JMA?RGML. 5FSQ, RFC DPCOSCLAW MD ?L ?JJCJC ?R MLC CLB MD ? BGQRPG@SRGML UGJJ @C QGKGJ?P RM RFC DPCOSCLAW MD RFC ?JJCJC ?R RFC MRFCP CLB. 8FCL NMNSJ?RGMLQ @CAMKC ECMEP?NFGA?JJW BGQAMLRGLSMSQ, RF?R DPCC-DJMU MD ?JJCJCQ GQ NPCTCLRCB. 8FCL RF?R QCN?P?RGML J?QRQ DMP ? NCPGMB MD RGKC, RFC RUM NMNSJ?RGMLQ ?PC ?@JC RM CTMJTC ?JMLE BGDDCPCLR RP?HCARMPGCQ. 5FSQ, RFCGP ?JJCJC DPCOSCLAGCQ ?R LSKCPMSQ ECLCRGA JMAG EP?BS?JJW @CAMKC KMPC ?LB KMPC BGDDCPCLR ?Q LCU ?JJCJCQ GLBCNCLBCLRJW ?PGQC @W KSR?RGML GL C?AF NMNSJ?RGML. 5WNGA?JJW, CLTGPMLKCLR?J AMLBGRGMLQ, QSAF ?Q AJGK?RC, PCQMSPACQ, NPCB?RMPQ, ?LB AMKNCRGRMPQ DMP RFC RUM NMNSJ?RGMLQ UGJJ BGDDCP A?SQGLE L?RSP?J QCJCARGML RM D?TMP BGTCPECLR ?B?NR?RGMLQ GL C?AF EPMSN. *QMJ?RGML MD NMNSJ?RGMLQ JC?BGLE RM ?JJMN?RPGA QNCAG?RGML A?L MAASP GL ? T?PGCRW MD U?WQ: ? PGTCP DMPKGLE ? LCU @P?LAF, CPMQGML DMPKGLE ? LCU T?JJCW, ? EPMSN MD MPE?LGQKQ RP?TCJGLE RM ? LCU JMA?RGML UGRFMSR RFC ?@GJGRW RM PCRSPL, MP QCCBQ DJM?RGLE MTCP RFC MAC?L RM ?L GQJ?LB. 5FC L?RSPC MD RFC ECMEP?NFGA QCN?P?RGML LCACQQ?PW RM GQMJ?RC NMNSJ?RGMLQ BCNCLBQ CLRGPCJW ML RFC @GMJMEW MD RFC MPE?LGQK ?LB GRQ NMRCLRG?J DMP BGQNCPQ?J. *D RUM DJWGLE GLQCAR NMNSJ?RGMLQ RMMI SN PCQGBCLAC GL QCN?P?RC LC?P@W T?JJCWQ, AF?LACQ ?PC, GLBGTGBS?JQ DPMK C?AF NMNSJ?RGML UMSJB DJW @?AI ?LB DMPRF AMLRGLSGLE ECLC DJMU. )MUCTCP, GD RUM PMBCLR NMNSJ?RGMLQ @CA?KC BGTGBCB @W RFC DMPK?RGML MD ? LCU J?IC, AMLRGLSCB ECLC DJMU UMSJB @C SLJGICJW; RFCPCDMPC, QNCAG?RGML UMSJB @C KMPC JGICJW. #GMJMEGQRQ EPMSN ?JJMN?RPGA NPMACQQCQ GLRM RUM A?RCEMPGCQ: BGQNCPQ?J ?LB TGA?PG?LAC. Dispersal GQ UFCL ? DCU KCK@CPQ MD ? QNCAGCQ KMTC RM ? LCU ECMEP?NFGA?J ?PC?, ?LB vicariance GQ UFCL ? L?RSP?J QGRS?RGML ?PGQCQ RM NFWQGA?JJW BGTGBC MPE?LGQKQ. 4AGCLRGQRQ F?TC BMASKCLRCB LSKCPMSQ A?QCQ MD ?JJMN?RPGA QNCAG?RGML R?IGLE NJ?AC. 'MP CV?KNJC, ?JMLE RFC UCQR AM?QR MD RFC 6LGRCB 4R?RCQ, RUM QCN?P?RC QS@-QNCAGCQ MD QNMRRCB MUJQ CVGQR. 5FC LMPRFCPL QNMRRCB MUJ F?Q ECLCRGA ?LB NFCLMRWNGA BGDDCPCLACQ DPMK GRQ AJMQC PCJ?RGTC: RFC .CVGA?L QNMRRCB MUJ, UFGAF JGTCQ GL RFC QMSRF (Figure 18.12).
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490
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/MRGAC RFC BGDDCPCLACQ GL RFC QNCAGCQs @C?IQ GL Figure 18.13. &TMJSRGML GL PCQNMLQC RM L?RSP?J QCJCARGML @?QCB ML QNCAGDGA DMMB QMSPACQ GL C?AF LCU F?@GR?R JCB RM CTMJSRGML MD ? BGDDCPCLR @C?I QSGRCB RM RFC QNCAGDGA DMMB QMSPAC. 5FC QCCB-C?RGLE @GPB F?Q ? RFGAICP, QRPMLECP @C?I UFGAF GQ QSGRCB RM @PC?I F?PB LSRQ. 5FC LCAR?P-C?RGLE @GPBQ F?TC JMLE @C?IQ RM BGN GLRM DJMUCPQ RM PC?AF RFC LCAR?P. 5FC GLQCAR-C?RGLE @GPBQ F?TC @C?IQ JGIC QUMPBQ, ?NNPMNPG?RC DMP QR?@@GLE ?LB GKN?JGLE GLQCARQ. %?PUGLsQ DGLAFCQ ?PC ?LMRFCP CV?KNJC MD ?B?NRGTC P?BG?RGML GL ?L ?PAFGNCJ?EM.
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!@>LI< 18.14 !KBRMILFAV OBPRIQP TEBK QEB D>JBQBP E>SB QLL J>KV LO QLL CBT @EOLJLPLJBP ARB QL KLKAFPGRK@QFLK AROFKD JBFLPFP. )K QEB BU>JMIB PELTK EBOB, QEB OBPRIQFKD LCCPMOFKD TFII E>SB 25+1 LO 25-1 @EOLJLPLJBP
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492
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
-<GIF;L:K@M< $JFC8K@FE
(GTCL CLMSEF RGKC, RFC ECLCRGA ?LB NFCLMRWNGA BGTCPECLAC @CRUCCL NMNSJ?RGMLQ UGJJ ?DDCAR AF?P?ARCPQ RF?R GLDJSCLAC PCNPMBSARGML: GD GLBGTGBS?JQ MD RFC RUM NMNSJ?RGMLQ UCPC RM @C @PMSEFR RMECRFCP, K?RGLE UMSJB @C JCQQ JGICJW, @SR GD K?RGLE MAASPPCB, MDDQNPGLE UMSJB @C LML-TG?@JC MP GLDCPRGJC. .?LW RWNCQ MD BGTCPEGLE AF?P?ARCPQ K?W ?DDCAR RFC reproductive isolation, RFC ?@GJGRW RM GLRCP@PCCB, MD RFC RUM NMNSJ?RGMLQ. 3CNPMBSARGTC GQMJ?RGML A?L R?IC NJ?AC GL ? T?PGCRW MD U?WQ. 4AGCLRGQRQ MPE?LGXC RFCK GLRM RUM EPMSNQ: NPCXWEMRGA @?PPGCPQ ?LB NMQRXWEMRGA @?PPGCPQ. 3CA?JJ RF?R ? XWEMRC GQ ? DCPRGJGXCB CEE: RFC DGPQR ACJJ MD RFC BCTCJMNKCLR MD ?L MPE?LGQK RF?R PCNPMBSACQ QCVS?JJW. 5FCPCDMPC, ? prezygotic barrier GQ ? KCAF?LGQK RF?R @JMAIQ PCNPMBSARGML DPMK R?IGLE NJ?AC; RFGQ GLAJSBCQ @?PPGCPQ RF?R NPCTCLR DCPRGJGX?RGML UFCL MPE?LGQKQ ?RRCKNR PCNPMBSARGML. " postzygotic barrier MAASPQ ?DRCP XWEMRC DMPK?RGML; RFGQ GLAJSBCQ MPE?LGQKQ RF?R BMLsR QSPTGTC RFC CK@PWMLGA QR?EC ?LB RFMQC RF?R ?PC @MPL QRCPGJC. 4MKC RWNCQ MD NPCXWEMRGA @?PPGCPQ NPCTCLR PCNPMBSARGML CLRGPCJW. .?LW MPE?LGQKQ MLJW PCNPMBSAC ?R ACPR?GL RGKCQ MD RFC WC?P, MDRCL HSQR ?LLS?JJW. %GDDCPCLACQ GL @PCCBGLE QAFCBSJCQ, A?JJCB temporal isolation, A?L ?AR ?Q ? DMPK MD PCNPMBSARGTC GQMJ?RGML. 'MP CV?KNJC, RUM QNCAGCQ MD DPMEQ GLF?@GR RFC Q?KC ?PC?, @SR MLC PCNPMBSACQ DPMK +?LS?PW RM .?PAF, UFCPC?Q RFC MRFCP PCNPMBSACQ DPMK .?PAF RM .?W (Figure 18.17).
!@>LI< 18.17 4EBPB QTL OBI>QBA COLD PMB@FBP BUEF?FQ QBJMLO>I OBMOLAR@QFSB FPLI>QFLK. (>) R(5( (<969( ?OBBAP B>OIFBO FK QEB VB>O QE>K (?) R(5( )6@300. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V ->OH 2. *BKKFKDP, 53&73; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V !IBPP>KAOL #>QBK>WWF)
*L QMKC A?QCQ, NMNSJ?RGMLQ MD ? QNCAGCQ KMTC MP ?PC KMTCB RM ? LCU F?@GR?R ?LB R?IC SN PCQGBCLAC GL ? NJ?AC RF?R LM JMLECP MTCPJ?NQ UGRF RFC MRFCP NMNSJ?RGMLQ MD RFC Q?KC QNCAGCQ. 5FGQ QGRS?RGML GQ A?JJCB habitat isolation. 3CNPMBSARGML UGRF RFC N?PCLR QNCAGCQ AC?QCQ, ?LB ? LCU EPMSN CVGQRQ RF?R GQ LMU
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493
PCNPMBSARGTCJW ?LB ECLCRGA?JJW GLBCNCLBCLR. 'MP CV?KNJC, ? APGAICR NMNSJ?RGML RF?R U?Q BGTGBCB ?DRCP ? DJMMB AMSJB LM JMLECP GLRCP?AR UGRF C?AF MRFCP. 0TCP RGKC, RFC DMPACQ MD L?RSP?J QCJCARGML, KSR?RGML, ?LB ECLCRGA BPGDR UGJJ JGICJW PCQSJR GL RFC BGTCPECLAC MD RFC RUM EPMSNQ (Figure 18.18).
!@>LI< 18.18 3MB@F>QFLK @>K L@@RO TEBK QTL MLMRI>QFLKP L@@RMV AFCCBOBKQ E>?FQ>QP. 4EB E>?FQ>QP KBBA KLQ ?B C>O >M>OQ. 4EB @OF@HBQ (>) G9@33<: 7,55:@3=(50*<: MOBCBOP P>KAV PLFI, >KA QEB @OF@HBQ (?) G9@33<: -094<: MOBCBOP IL>JV PLFI. 4EB QTL PMB@FBP @>K IFSB FK @ILPB MOLUFJFQV, ?RQ ?B@>RPB LC QEBFO AFCCBOBKQ PLFI MOBCBOBK@BP, QEBV ?B@>JB DBKBQF@>IIV FPLI>QBA.
Behavioral isolation MAASPQ UFCL RFC NPCQCLAC MP ?@QCLAC MD ? QNCAGDGA @CF?TGMP NPCTCLRQ PCNPMBSARGML DPMK R?IGLE NJ?AC. 'MP CV?KNJC, K?JC DGPCDJGCQ SQC QNCAGDGA JGEFR N?RRCPLQ RM ?RRP?AR DCK?JCQ. 7?PGMSQ QNCAGCQ MD DGPCDJGCQ BGQNJ?W RFCGP JGEFRQ BGDDCPCLRJW. *D ? K?JC MD MLC QNCAGCQ RPGCB RM ?RRP?AR RFC DCK?JC MD ?LMRFCP, QFC UMSJB LMR PCAMELGXC RFC JGEFR N?RRCPL ?LB UMSJB LMR K?RC UGRF RFC K?JC. 0RFCP NPCXWEMRGA @?PPGCPQ UMPI UFCL BGDDCPCLACQ GL RFCGP E?KCRC ACJJQ (CEEQ ?LB QNCPK) NPCTCLR DCPRGJGX?RGML DPMK R?IGLE NJ?AC; RFGQ GQ A?JJCB ? gametic barrier. 4GKGJ?PJW, GL QMKC A?QCQ AJMQCJW PCJ?RCB MPE?LGQKQ RPW RM K?RC, @SR RFCGP PCNPMBSARGTC QRPSARSPCQ QGKNJW BM LMR DGR RMECRFCP. 'MP CV?KNJC, B?KQCJDJW K?JCQ MD BGDDCPCLR QNCAGCQ F?TC BGDDCPCLRJW QF?NCB PCNPMBSARGTC MPE?LQ. *D MLC QNCAGCQ RPGCQ RM K?RC UGRF RFC DCK?JC MD ?LMRFCP, RFCGP @MBW N?PRQ QGKNJW BM LMR DGR RMECRFCP. (Figure 18.19).
!@>LI< 18.19 4EB PE>MB LC QEB J>IB OBMOLAR@QFSB LOD>K S>OFBP >JLKD J>IB A>JPBICIV PMB@FBP, >KA FP LKIV @LJM>QF?IB TFQE QEB CBJ>IB LC QE>Q PMB@FBP. 2BMOLAR@QFSB LOD>K FK@LJM>QF?FIFQV HBBMP QEB PMB@FBP OBMOLAR@QFSBIV FPLI>QBA.
*L NJ?LRQ, ACPR?GL QRPSARSPCQ ?GKCB RM ?RRP?AR MLC RWNC MD NMJJGL?RMP QGKSJR?LCMSQJW NPCTCLR ? BGDDCPCLR NMJJGL?RMP DPMK ?AACQQGLE RFC NMJJCL. 5FC RSLLCJ RFPMSEF UFGAF ?L ?LGK?J KSQR ?AACQQ LCAR?P A?L T?PW UGBCJW GL JCLERF ?LB BG?KCRCP, UFGAF NPCTCLRQ RFC NJ?LR DPMK @CGLE APMQQ-NMJJGL?RCB UGRF ? BGDDCPCLR QNCAGCQ (Figure 18.20).
!@>LI< 18.20 3LJB CILTBOP E>SB BSLISBA QL >QQO>@Q @BOQ>FK MLIIFK>QLOP. 4EB (>) TFAB CLUDILSB CILTBO FP >A>MQBA CLO MLIIFK>QFLK ?V ?BBP, TEFIB QEB (?) ILKD, QR?B-PE>MBA QORJMBQ @OBBMBO CILTBO FP >A>MQBA CLO MLIIFK>QFLK ?V ERJJFKD ?FOAP.
8FCL DCPRGJGX?RGML R?ICQ NJ?AC ?LB ? XWEMRC DMPKQ, NMQRXWEMRGA @?PPGCPQ A?L NPCTCLR PCNPMBSARGML. )W@PGB GLBGTGBS?JQ GL K?LW A?QCQ A?LLMR DMPK LMPK?JJW GL RFC UMK@ ?LB QGKNJW BM LMR QSPTGTC N?QR RFC CK@PWMLGA QR?ECQ. 5FGQ GQ A?JJCB hybrid inviability @CA?SQC RFC FW@PGB MPE?LGQKQ QGKNJW ?PC LMR TG?@JC. *L ?LMRFCP NMQRXWEMRGA QGRS?RGML, PCNPMBSARGML JC?BQ RM RFC @GPRF ?LB EPMURF MD ? FW@PGB RF?R GQ QRCPGJC ?LB SL?@JC RM PCNPMBSAC MDDQNPGLE MD RFCGP MUL; RFGQ GQ A?JJCB FW@PGB QRCPGJGRW.
494
#(!04%2 18 X %6/,54)/. !.$ 4(% /2)'). /& 30%#)%3
H'(/9'9 I3,1:+3)+ 43 !5+)/'9/43 4WKN?RPGA QNCAG?RGML K?W ?JQM R?IC NJ?AC GL U?WQ MRFCP RF?L NMJWNJMGBW. 'MP CV?KNJC, AMLQGBCP ? QNCAGCQ MD DGQF RF?R JGTCQ GL ? J?IC. "Q RFC NMNSJ?RGML EPMUQ, AMKNCRGRGML DMP DMMB ?JQM EPMUQ. 6LBCP NPCQQSPC RM DGLB DMMB, QSNNMQC RF?R ? EPMSN MD RFCQC DGQF F?B RFC ECLCRGA DJCVG@GJGRW RM BGQAMTCP ?LB DCCB MDD ?LMRFCP PCQMSPAC RF?R U?Q SLSQCB @W RFC MRFCP DGQF. 8F?R GD RFGQ LCU DMMB QMSPAC U?Q DMSLB ?R ? BGDDCPCLR BCNRF MD RFC J?IC 0TCP RGKC, RFMQC DCCBGLE ML RFC QCAMLB DMMB QMSPAC UMSJB GLRCP?AR KMPC UGRF C?AF MRFCP RF?L RFC MRFCP DGQF; RFCPCDMPC, RFCW UMSJB @PCCB RMECRFCP ?Q UCJJ. 0DDQNPGLE MD RFCQC DGQF UMSJB JGICJW @CF?TC ?Q RFCGP N?PCLRQ: DCCBGLE ?LB JGTGLE GL RFC Q?KC ?PC? ?LB ICCNGLE QCN?P?RC DPMK RFC MPGEGL?J NMNSJ?RGML. *D RFGQ EPMSN MD DGQF AMLRGLSCB RM PCK?GL QCN?P?RC DPMK RFC DGPQR NMNSJ?RGML, CTCLRS?JJW QWKN?RPGA QNCAG?RGML KGEFR MAASP ?Q KMPC ECLCRGA BGDDCPCLACQ ?AASKSJ?RCB @CRUCCL RFCK. 5FGQ QACL?PGM BMCQ NJ?W MSR GL L?RSPC, ?Q BM MRFCPQ RF?R JC?B RM PCNPMBSARGTC GQMJ?RGML. 0LC QSAF NJ?AC GQ -?IC 7GARMPG? GL "DPGA?, D?KMSQ DMP GRQ QWKN?RPGA QNCAG?RGML MD AGAFJGB DGQF. 3CQC?PAFCPQ F?TC DMSLB FSLBPCBQ MD QWKN?RPGA QNCAG?RGML CTCLRQ GL RFCQC DGQF, UFGAF F?TC LMR MLJW F?NNCLCB GL EPC?R LSK@CP, @SR ?JQM MTCP ? QFMPR NCPGMB MD RGKC. Figure 18.21 QFMUQ RFGQ RWNC MD QNCAG?RGML ?KMLE ? AGAFJGB DGQF NMNSJ?RGML GL /GA?P?ES?. *L RFGQ JMA?JC, RUM RWNCQ MD AGAFJGBQ JGTC GL RFC Q?KC ECMEP?NFGA JMA?RGML @SR F?TC AMKC RM F?TC BGDDCPCLR KMPNFMJMEGCQ RF?R ?JJMU RFCK RM C?R T?PGMSQ DMMB QMSPACQ.
!@>LI< 18.21 #F@EIFA CFPE COLJ ,>HB !MLVBNRB, .F@>O>DR>, PELT BSFABK@B LC PVJM>QOF@ PMB@F>QFLK. ,>HB !MLVBNRB, > @O>QBO I>HB, FP 1800 VB>OP LIA, ?RQ DBKBQF@ BSFABK@B FKAF@>QBP QE>Q QEB I>HB T>P MLMRI>QBA LKIV 100 VB>OP >DL ?V > PFKDIB MLMRI>QFLK LC @F@EIFA CFPE. .BSBOQEBIBPP, QTL MLMRI>QFLKP TFQE AFPQFK@Q JLOMELILDFBP >KA AFBQP KLT BUFPQ FK QEB I>HB, >KA P@FBKQFPQP ?BIFBSB QEBPB MLMRI>QFLKP J>V ?B FK >K B>OIV PQ>DB LC PMB@F>QFLK.
18.3 R -<:FEE<:K@FE 8E; -8K<J F= .G<:@8K@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C N?RFU?WQ MD QNCAGCQ CTMJSRGML GL FW@PGB XMLCQ z &VNJ?GL RFC RUM K?HMP RFCMPGCQ ML P?RCQ MD QNCAG?RGML 4NCAG?RGML MAASPQ MTCP ? QN?L MD CTMJSRGML?PW RGKC, QM UFCL ? LCU QNCAGCQ ?PGQCQ, RFCPC GQ ? RP?LQGRGML NCPGMB BSPGLE UFGAF RFC AJMQCJW PCJ?RCB QNCAGCQ AMLRGLSC RM GLRCP?AR.
-<:FEE<:K@FE
"DRCP QNCAG?RGML, RUM QNCAGCQ K?W PCAMK@GLC MP CTCL AMLRGLSC GLRCP?ARGLE GLBCDGLGRCJW. *LBGTGBS?J MPE?LGQKQ UGJJ K?RC UGRF ?LW LC?P@W GLBGTGBS?J UFM RFCW ?PC A?N?@JC MD @PCCBGLE UGRF. "L ?PC? UFCPC RUM AJMQCJW PCJ?RCB QNCAGCQ AMLRGLSC RM GLRCP?AR ?LB PCNPMBSAC, DMPKGLE FW@PGBQ, GQ A?JJCB ? hybrid zone. 0TCP RGKC, RFC FW@PGB XMLC K?W AF?LEC BCNCLBGLE ML RFC DGRLCQQ MD RFC FW@PGBQ ?LB RFC PCNPMBSARGTC @?PPGCPQ (Figure 18.22). *D RFC FW@PGBQ ?PC JCQQ DGR RF?L RFC N?PCLRQ, PCGLDMPACKCLR MD QNCAG?RGML MAASPQ, ?LB RFC QNCAGCQ AMLRGLSC RM BGTCPEC SLRGJ RFCW A?L LM JMLECP K?RC ?LB NPMBSAC TG?@JC MDDQNPGLE. *D PCNPMBSARGTC @?PPGCPQ UC?ICL, DSQGML MAASPQ ?LB RFC RUM QNCAGCQ @CAMKC MLC. #?PPGCPQ PCK?GL RFC Q?KC GD FW@PGBQ ?PC DGR ?LB PCNPMBSARGTC: QR?@GJGRW K?W MAASP ?LB FW@PGBGX?RGML AMLRGLSCQ.
#(!04%2 18 X %6/,54)/. !.$ 4(% /2)'). /& 30%#)%3
495
!@>LI< 18.22 !CQBO PMB@F>QFLK E>P L@@ROOBA, QEB QTL PBM>O>QB ?RQ @ILPBIV OBI>QBA PMB@FBP J>V @LKQFKRB QL MOLAR@B LCCPMOFKD FK >K >OB> @>IIBA QEB EV?OFA WLKB. 2BFKCLO@BJBKQ, CRPFLK, LO PQ>?FIFQV J>V OBPRIQ, ABMBKAFKD LK OBMOLAR@QFSB ?>OOFBOP >KA QEB OBI>QFSB CFQKBPP LC QEB EV?OFAP.
)C QTL PMB@FBP B>Q > AFCCBOBKQ AFBQ ?RQ LKB LC QEB CLLA PLRO@BP FP BIFJFK>QBA >KA ?LQE PMB@FBP >OB CLO@BA QL B>Q QEB P>JB CLLAP, TE>Q @E>KDB FK QEB EV?OFA WLKB FP JLPQ IFHBIV QL L@@RO )W@PGBQ A?L @C CGRFCP JCQQ DGR RF?L RFC N?PCLRQ, KMPC DGR, MP ?@MSR RFC Q?KC. 6QS?JJW FW@PGBQ RCLB RM @C JCQQ DGR; RFCPCDMPC, QSAF PCNPMBSARGML BGKGLGQFCQ MTCP RGKC, LSBEGLE RFC RUM QNCAGCQ RM BGTCPEC DSPRFCP GL ? NPMACQQ A?JJCB reinforcement. 5FGQ RCPK GQ SQCB @CA?SQC RFC JMU QSAACQQ MD RFC FW@PGBQ PCGLDMPACQ RFC MPGEGL?J QNCAG?RGML. *D RFC FW@PGBQ ?PC ?Q DGR MP KMPC DGR RF?L RFC N?PCLRQ, RFC RUM QNCAGCQ K?W DSQC @?AI GLRM MLC QNCAGCQ (Figure 18.23). 4AGCLRGQRQ F?TC ?JQM M@QCPTCB RF?R QMKCRGKCQ RUM QNCAGCQ UGJJ PCK?GL QCN?P?RC @SR ?JQM AMLRGLSC RM GLRCP?AR RM NPMBSAC QMKC FW@PGB GLBGTGBS?JQ; RFGQ GQ AJ?QQGDGCB ?Q QR?@GJGRW @CA?SQC LM PC?J LCR AF?LEC GQ R?IGLE NJ?AC.
18IP@E> -8K<J F= .G<:@8K@FE

4AGCLRGQRQ ?PMSLB RFC UMPJB QRSBW QNCAG?RGML, BMASKCLRGLE M@QCPT?RGMLQ @MRF MD JGTGLE MPE?LGQKQ ?LB RFMQC DMSLB GL RFC DMQQGJ PCAMPB. "Q RFCGP GBC?Q R?IC QF?NC ?LB ?Q PCQC?PAF PCTC?JQ LCU BCR?GJQ ?@MSR FMU JGDC CTMJTCQ, RFCW BCTCJMN KMBCJQ RM FCJN CVNJ?GL P?RCQ MD QNCAG?RGML. *L RCPKQ MD FMU OSGAIJW QNCAG?RGML MAASPQ, RUM N?RRCPLQ ?PC ASPPCLRJW M@QCPTCB: EP?BS?J QNCAG?RGML KMBCJ ?LB NSLARS?RCB COSGJG@PGSK KMBCJ. *L RFC gradual speciation model, QNCAGCQ BGTCPEC EP?BS?JJW MTCP RGKC GL QK?JJ QRCNQ. *L RFC punctuated equilibrium KMBCJ, ? LCU QNCAGCQ SLBCPEMCQ AF?LECQ OSGAIJW DPMK RFC N?PCLR QNCAGCQ, ?LB RFCL PCK?GLQ J?PECJW SLAF?LECB DMP JMLE NCPGMBQ MD RGKC ?DRCPU?PB (Figure 18.23). 5FGQ C?PJW AF?LEC KMBCJ GQ A?JJCB NSLARS?RCB COSGJG@PGSK, @CA?SQC GR @CEGLQ UGRF ? NSLARS?RCB MP NCPGMBGA AF?LEC ?LB RFCL PCK?GLQ GL @?J?LAC ?DRCPU?PB. 8FGJC NSLARS?RCB COSGJG@PGSK QSEECQRQ ? D?QRCP RCKNM, GR BMCQ LMR LCACQQ?PGJW CVAJSBC EP?BS?JGQK.
496
#(!04%2 18 X %6/,54)/. !.$ 4(% /2)'). /& 30%#)%3
!@>LI< 18.23 )K (>) DO>AR>I PMB@F>QFLK, PMB@FBP AFSBODB >Q > PILT, PQB>AV M>@B >P QO>FQP @E>KDB FK@OBJBKQ>IIV. )K (?) MRK@QR>QBA BNRFIF?OFRJ, PMB@FBP AFSBODB NRF@HIV >KA QEBK OBJ>FK RK@E>KDBA CLO ILKD MBOFLAP LC QFJB.
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qQSAF ?Q ? BPMN GL RFC U?RCP JCTCJq? QK?JJ LSK@CP MD MPE?LGQKQ ?PC QCN?P?RCB DPMK RFC PCQR GL ? @PGCD NCPGMB MD RGKC, CQQCLRG?JJW DMPKGLE MLC J?PEC ?LB MLC RGLW NMNSJ?RGML. 5FC RGLW NMNSJ?RGML D?ACQ LCU CLTGPMLKCLR?J AMLBGRGMLQ. #CA?SQC GRQ ECLC NMMJ OSGAIJW @CA?KC QM QK?JJ, ?LW T?PG?RGML RF?R QSPD?ACQ ?LB RF?R ?GBQ GL QSPTGTGLE RFC LCU AMLBGRGMLQ @CAMKCQ RFC NPCBMKGL?LR DMPK.
7GQGR this website (http://openstaxcollege.org/l/snails) RM AMLRGLSC RFC QNCAG?RGML QRMPW MD RFC QL?GJQ.
#(!04%2 18 X %6/,54)/. !.$ 4(% /2)'). /& 30%#)%3
497
& 4 / -(.
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
498
#(!04%2 18 X %6/,54)/. !.$ 4(% /2)'). /& 30%#)%3
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
C#A+/ - .0((A-4
18.1 0E;<IJK8E;@E> MFCLK@FE
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
18.2 !FID8K@FE F= )<N .G<:@<J

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
18.3 -<:FEE<:K@FE 8E; -8K<J F= .G<:@8K@FE

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
A-/ C*)) C/$*) ,0 ./$*).

1. Figure 18.14 8FGAF GQ KMQR JGICJW RM QSPTGTC, MDDQNPGLE UGRF 25+1 AFPMKMQMKCQ MP MDDQNPGLE UGRF 25-1 AFPMKMQMKCQ 2. Figure 18.22 *D RUM QNCAGCQ C?R ? BGDDCPCLR BGCR @SR MLC MD RFC DMMB QMSPACQ GQ CJGKGL?RCB ?LB @MRF QNCAGCQ ?PC DMPACB RM C?R RFC Q?KC DMMBQ, UF?R AF?LEC GL RFC FW@PGB XMLC GQ KMQR JGICJW RM MAASP 3. Figure 18.23 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ GQ D?JQC >. 1SLARS?RCB COSGJG@PGSK GQ KMQR JGICJW RM MAASP GL ? QK?JJ NMNSJ?RGML RF?R
CVNCPGCLACQ ? P?NGB AF?LEC GL GRQ CLTGPMLKCLR. ?. 1SLARS?RCB COSGJG@PGSK GQ KMQR JGICJW RM MAASP GL ? J?PEC NMNSJ?RGML RF?R JGTCQ GL ? QR?@JC AJGK?RC. @. (P?BS?J QNCAG?RGML GQ KMQR JGICJW RM MAASP GL QNCAGCQ RF?R JGTC GL ? QR?@JC AJGK?RC. A. (P?BS?J QNCAG?RGML ?LB NSLARS?RCB COSGJG@PGSK @MRF PCQSJR GL RFC CTMJSRGML MD LCU QNCAGCQ.
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4. 8FGAF QAGCLRGDGA AMLACNR BGB $F?PJCQ %?PUGL ?LB "JDPCB 8?JJ?AC GLBCNCLBCLRJW BGQAMTCP >. ?. @. A. KSR?RGML L?RSP?J QCJCARGML MTCP@PCCBGLE QCVS?J PCNPMBSARGML 10. 8FGAF T?PG?@JC GLAPC?QCQ RFC JGICJGFMMB MD ?JJMN?RPGA QNCAG?RGML R?IGLE NJ?AC KMPC OSGAIJW >. ?. @. A. JMUCP P?RC MD KSR?RGML JMLECP BGQR?LAC @CRUCCL BGTGBCB EPMSNQ GLAPC?QCB GLQR?LACQ MD FW@PGB DMPK?RGML COSGT?JCLR LSK@CPQ MD GLBGTGBS?JQ GL C?AF NMNSJ?RGML
5. 8FGAF MD RFC DMJJMUGLE QGRS?RGMLQ UGJJ JC?B RM L?RSP?J QCJCARGML >. 5FC QCCBQ MD RUM NJ?LRQ J?LB LC?P C?AF MRFCP ?LB MLC EPMUQ J?PECP RF?L RFC MRFCP. ?. 5UM RWNCQ MD DGQF C?R RFC Q?KC IGLB MD DMMB, ?LB MLC GQ @CRRCP ?@JC RM E?RFCP DMMB RF?L RFC MRFCP. @. .?JC JGMLQ AMKNCRC DMP RFC PGEFR RM K?RC UGRF DCK?JCQ, UGRF MLJW MLC NMQQG@JC UGLLCP. A. ?JJ MD RFC ?@MTC 6. 8FGAF BCQAPGNRGML GQ ?L CV?KNJC MD ? NFCLMRWNC >. " ACPR?GL BSAI F?Q ? @JSC @C?I. ?. " KSR?RGML MAASPPCB RM ? DJMUCP. @. .MQR AFCCR?FQ JGTC QMJGR?PW JGTCQ. A. @MRF ? ?LB A 7. 8FGAF QGRS?RGML GQ KMQR JGICJW ?L CV?KNJC MD AMLTCPECLR CTMJSRGML >. 4OSGB ?LB FSK?LQ F?TC CWCQ QGKGJ?P GL QRPSARSPC. ?. 8MPKQ ?LB QL?ICQ @MRF KMTC UGRFMSR JCEQ. @. 4MKC @?RQ ?LB @GPBQ F?TC UGLEQ RF?R ?JJMU RFCK RM DJW A. ?JJ MD RFC ?@MTC 8. 8FGAF QGRS?RGML UMSJB KMQR JGICJW JC?B RM ?JJMN?RPGA QNCAG?RGML >. DJMMB A?SQCQ RFC DMPK?RGML MD ? LCU J?IC. ?. " QRMPK A?SQCQ QCTCP?J J?PEC RPCCQ RM D?JJ BMUL. @. " KSR?RGML A?SQCQ ? LCU RP?GR RM BCTCJMN. A. "L GLHSPW A?SQCQ ?L MPE?LGQK RM QCCI MSR ? LCU DMMB QMSPAC. 9. 8F?R GQ RFC K?GL BGDDCPCLAC @CRUCCL BGQNCPQ?J ?LB TGA?PG?LAC >. 0LC JC?BQ RM ?JJMN?RPGA QNCAG?RGML, UFCPC?Q RFC MRFCP JC?BQ RM QWKN?RPGA QNCAG?RGML. ?. 0LC GLTMJTCQ RFC KMTCKCLR MD RFC MPE?LGQK, ?LB RFC MRFCP GLTMJTCQ ? AF?LEC GL RFC CLTGPMLKCLR. @. 0LC BCNCLBQ ML ? ECLCRGA KSR?RGML MAASPPGLE, ?LB RFC MRFCP BMCQ LMR. A. 0LC GLTMJTCQ AJMQCJW PCJ?RCB MPE?LGQKQ, ?LB RFC MRFCP GLTMJTCQ MLJW GLBGTGBS?JQ MD RFC Q?KC QNCAGCQ.
11. 8F?R GQ RFC K?GL BGDDCPCLAC @CRUCCL ?SRMNMJWNJMGB ?LB ?JJMNMJWNJMGB >. RFC LSK@CP MD AFPMKMQMKCQ ?. RFC DSLARGML?JGRW MD RFC AFPMKMQMKCQ @. RFC QMSPAC MD RFC CVRP? AFPMKMQMKCQ A. RFC LSK@CP MD KSR?RGMLQ GL RFC CVRP? AFPMKMQMKCQ 12. 8FGAF PCNPMBSARGTC AMK@GL?RGML NPMBSACQ FW@PGBQ >. UFCL GLBGTGBS?JQ MD RFC Q?KC QNCAGCQ GL BGDDCPCLR ECMEP?NFGA?J ?PC?Q PCNPMBSAC ?. UFCL ?LW RUM GLBGTGBS?JQ QF?PGLE RFC Q?KC F?@GR?R PCNPMBSAC @. UFCL KCK@CPQ MD AJMQCJW PCJ?RCB QNCAGCQ PCNPMBSAC A. UFCL MDDQNPGLE MD RFC Q?KC N?PCLRQ PCNPMBSAC 13. 8FGAF AMLBGRGML GQ RFC @?QGQ DMP ? QNCAGCQ RM @C PCNPMBSARGTCJW GQMJ?RCB DPMK MRFCP KCK@CPQ >. *R BMCQ LMR QF?PC GRQ F?@GR?R UGRF PCJ?RCB QNCAGCQ. ?. *R BMCQ LMR CVGQR MSR MD ? QGLEJC F?@GR?R. @. *R BMCQ LMR CVAF?LEC ECLCRGA GLDMPK?RGML UGRF MRFCP QNCAGCQ. A. *R BMCQ LMR SLBCPEM CTMJSRGML?PW AF?LECQ DMP ? QGELGDGA?LR NCPGMB MD RGKC. 14. 8FGAF QGRS?RGML GQ 56; ?L CV?KNJC MD ? NPCXWEMRGA @?PPGCP >. 5UM QNCAGCQ MD RSPRJCQ @PCCB ?R BGDDCPCLR RGKCQ MD RFC WC?P. ?. 5UM QNCAGCQ MD DJMUCPQ ?RRP?AR BGDDCPCLR NMJJGL?RMPQ. @. 5UM QNCAGCQ MD @GPBQ BGQNJ?W BGDDCPCLR K?RGLE B?LACQ. A. 5UM QNCAGCQ MD GLQCARQ NPMBSAC GLDCPRGJC MDDQNPGLE. 15. 8FGAF RCPK GQ SQCB RM BCQAPG@C RFC AMLRGLSCB BGTCPECLAC MD QNCAGCQ @?QCB ML RFC JMU DGRLCQQ MD FW@PGB MDDQNPGLE >. PCGLDMPACKCLR ?. DSQGML @. QR?@GJGRW A. NSLARS?RCB COSGJG@PGSK 16. 8FGAF AMKNMLCLRQ MD QNCAG?RGML UMSJB @C JC?QR JGICJW RM @C ? N?PR MD NSLARS?RCB COSGJG@PGSK >. ? BGTGQGML MD NMNSJ?RGMLQ ?. ? AF?LEC GL CLTGPMLKCLR?J AMLBGRGMLQ
500
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@. MLEMGLE ECLC DJMU ?KMLE ?JJ GLBGTGBS?JQ
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17. *D ? NCPQML QA?RRCPQ ? F?LBDSJ MD E?PBCL NC? NJ?LR QCCBQ GL MLC ?PC?, FMU UMSJB L?RSP?J QCJCARGML UMPI GL RFGQ QGRS?RGML 18. 8FW BM QAGCLRGQRQ AMLQGBCP TCQRGEG?J QRPSARSPCQ CTGBCLAC DMP CTMJSRGML 19. )MU BMCQ RFC QAGCLRGDGA KC?LGLE MD tRFCMPWu BGDDCP DPMK RFC AMKKML TCPL?ASJ?P KC?LGLE 20. &VNJ?GL UFW RFC QR?RCKCLR RF?R ? KMLICW GQ KMPC CTMJTCB RF?L ? KMSQC GQ GLAMPPCAR. 21. 8FW BM GQJ?LB AF?GLQ NPMTGBC GBC?J AMLBGRGMLQ DMP ?B?NRGTC P?BG?RGML RM MAASP 22. 5UM QNCAGCQ MD DGQF F?B PCACLRJW SLBCPEMLC QWKN?RPGA QNCAG?RGML. 5FC K?JCQ MD C?AF QNCAGCQ F?B ? BGDDCPCLR AMJMPGLE RFPMSEF UFGAF RFC DCK?JCQ AMSJB GBCLRGDW ?LB AFMMQC ? N?PRLCP DPMK FCP MUL QNCAGCQ. "DRCP QMKC RGKC, NMJJSRGML K?BC RFC J?IC QM AJMSBW RF?R GR U?Q F?PB DMP DCK?JCQ RM BGQRGLESGQF AMJMPQ. 8F?R KGEFR R?IC NJ?AC GL RFGQ QGRS?RGML 23. 8FW A?L NMJWNJMGBW GLBGTGBS?JQ JC?B RM QNCAG?RGML D?GPJW OSGAIJW 24. 8F?R BM @MRF P?RC MD QNCAG?RGML KMBCJQ F?TC GL AMKKML 25. %CQAPG@C ? QGRS?RGML UFCPC FW@PGB PCNPMBSARGML UMSJB A?SQC RUM QNCAGCQ RM DSQC GLRM MLC.
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19.1: +FGLC8K@FE MFCLK@FE 19.2: +FGLC8K@FE "<E<K@:J 19.3: A;8GK@M< MFCLK@FE
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502
#(!04%2 19 X 4(% %6/,54)/. /& 0/05,!4)/.3
19.1 R +FGLC8K@FE MFCLK@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CDGLC NMNSJ?RGML ECLCRGAQ ?LB BCQAPG@C FMU NMNSJ?RGML ECLCRGAQ GQ SQCB GL RFC QRSBW MD RFC CTMJSRGML MD NMNSJ?RGMLQ z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sQ UMPI U?Q PCBGQAMTCPCB GL RFC C?PJW RUCLRGCRF ACLRSPW ?R UFGAF RGKC ECLCRGAGQRQ UCPC P?NGBJW AMKGLE RM ?L SLBCPQR?LBGLE MD RFC @?QGAQ MD GLFCPGR?LAC. *LGRG?JJW, RFC LCUJW BGQAMTCPCB N?PRGASJ?RC L?RSPC MD ECLCQ K?BC GR BGDDGASJR DMP @GMJMEGQRQ RM SLBCPQR?LB FMU EP?BS?J CTMJSRGML AMSJB MAASP. #SR MTCP RFC LCVR DCU BCA?BCQ ECLCRGAQ ?LB CTMJSRGML UCPC GLRCEP?RCB GL UF?R @CA?KC ILMUL ?Q RFC modern synthesisqRFC AMFCPCLR SLBCPQR?LBGLE MD RFC PCJ?RGMLQFGN @CRUCCL L?RSP?J QCJCARGML ?LB ECLCRGAQ RF?R RMMI QF?NC @W RFC 1940Q ?LB GQ ECLCP?JJW ?AACNRCB RMB?W. *L QSK, RFC KMBCPL QWLRFCQGQ BCQAPG@CQ FMU CTMJSRGML?PW NPMACQQCQ, QSAF ?Q L?RSP?J QCJCARGML, A?L ?DDCAR ? NMNSJ?RGMLsQ ECLCRGA K?ICSN, ?LB, GL RSPL, FMU RFGQ A?L PCQSJR GL RFC EP?BS?J CTMJSRGML MD NMNSJ?RGMLQ ?LB QNCAGCQ. 5FC RFCMPW ?JQM AMLLCARQ RFGQ AF?LEC MD ? NMNSJ?RGML MTCP RGKC, A?JJCB microevolution, UGRF RFC NPMACQQCQ RF?R E?TC PGQC RM LCU QNCAGCQ ?LB FGEFCP R?VMLMKGA EPMSNQ UGRF UGBCJW BGTCPECLR AF?P?ARCPQ, A?JJCB macroevolution.
MFCLK@FE 8E; !CL 18::@E<J

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+FGLC8K@FE "<E<K@:J
3CA?JJ RF?R ? ECLC DMP ? N?PRGASJ?P AF?P?ARCP K?W F?TC QCTCP?J ?JJCJCQ, MP T?PG?LRQ, RF?R AMBC DMP BGDDCPCLR RP?GRQ ?QQMAG?RCB UGRF RF?R AF?P?ARCP. 'MP CV?KNJC, GL RFC "#0 @JMMB RWNC QWQRCK GL FSK?LQ, RFPCC ?JJCJCQ BCRCPKGLC RFC N?PRGASJ?P @JMMB-RWNC NPMRCGL ML RFC QSPD?AC MD PCB @JMMB ACJJQ. &?AF GLBGTGBS?J GL ? NMNSJ?RGML MD BGNJMGB MPE?LGQKQ A?L MLJW A?PPW RUM ?JJCJCQ DMP ? N?PRGASJ?P ECLC, @SR KMPC RF?L RUM K?W @C NPCQCLR GL RFC GLBGTGBS?JQ RF?R K?IC SN RFC NMNSJ?RGML. .CLBCJ DMJJMUCB ?JJCJCQ ?Q RFCW UCPC GLFCPGRCB DPMK N?PCLR RM MDDQNPGLE. *L RFC C?PJW RUCLRGCRF ACLRSPW, @GMJMEGQRQ GL ? DGCJB MD QRSBW ILMUL ?Q
#(!04%2 19 X 4(% %6/,54)/. /& 0/05,!4)/.3
503
population genetics @CE?L RM QRSBW FMU QCJCARGTC DMPACQ AF?LEC ? NMNSJ?RGML RFPMSEF AF?LECQ GL ?JJCJC ?LB ECLMRWNGA DPCOSCLAGCQ. 5FC allele frequency (MP ECLC DPCOSCLAW) GQ RFC P?RC ?R UFGAF ? QNCAGDGA ?JJCJC ?NNC?PQ UGRFGL ? NMNSJ?RGML. 6LRGJ LMU UC F?TC BGQASQQCB CTMJSRGML ?Q ? AF?LEC GL RFC AF?P?ARCPGQRGAQ MD ? NMNSJ?RGML MD MPE?LGQKQ, @SR @CFGLB RF?R NFCLMRWNGA AF?LEC GQ ECLCRGA AF?LEC. *L NMNSJ?RGML ECLCRGAQ, RFC RCPK CTMJSRGML GQ BCDGLCB ?Q ? AF?LEC GL RFC DPCOSCLAW MD ?L ?JJCJC GL ? NMNSJ?RGML. 6QGLE RFC "#0 @JMMB RWNC QWQRCK ?Q ?L CV?KNJC, RFC DPCOSCLAW MD MLC MD RFC ?JJCJCQ, I", GQ RFC LSK@CP MD AMNGCQ MD RF?R ?JJCJC <1= BGTGBCB @W ?JJ RFC AMNGCQ MD RFC "#0 ECLC GL RFC NMNSJ?RGML. 'MP CV?KNJC, ? QRSBW GL +MPB?L DMSLB ? DPCOSCLAW MD I" RM @C 26.1 NCPACLR. 5FC I# ?LB I0 ?JJCJCQ K?BC SN 13.4 NCPACLR ?LB 60.5 NCPACLR MD RFC ?JJCJCQ PCQNCARGTCJW, ?LB ?JJ MD RFC DPCOSCLAGCQ ?BBCB SN RM 100 NCPACLR. " AF?LEC GL RFGQ DPCOSCLAW MTCP RGKC UMSJB AMLQRGRSRC CTMJSRGML GL RFC NMNSJ?RGML. 5FC ?JJCJC DPCOSCLAW UGRFGL ? EGTCL NMNSJ?RGML A?L AF?LEC BCNCLBGLE ML CLTGPMLKCLR?J D?ARMPQ; RFCPCDMPC, ACPR?GL ?JJCJCQ @CAMKC KMPC UGBCQNPC?B RF?L MRFCPQ BSPGLE RFC NPMACQQ MD L?RSP?J QCJCARGML. /?RSP?J QCJCARGML A?L ?JRCP RFC NMNSJ?RGMLs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gene pool GQ RFC QSK MD ?JJ RFC ?JJCJCQ GL ? NMNSJ?RGML. 4MKCRGKCQ, ?JJCJC DPCOSCLAGCQ UGRFGL ? NMNSJ?RGML AF?LEC P?LBMKJW UGRF LM ?BT?LR?EC RM RFC NMNSJ?RGML MTCP CVGQRGLE ?JJCJC DPCOSCLAGCQ. 5FGQ NFCLMKCLML GQ A?JJCB ECLCRGA BPGDR. /?RSP?J QCJCARGML ?LB ECLCRGA BPGDR SQS?JJW MAASP QGKSJR?LCMSQJW GL NMNSJ?RGMLQ ?LB ?PC LMR GQMJ?RCB CTCLRQ. *R GQ F?PB RM BCRCPKGLC UFGAF NPMACQQ BMKGL?RCQ @CA?SQC GR GQ MDRCL LC?PJW GKNMQQG@JC RM BCRCPKGLC RFC A?SQC MD AF?LEC GL ?JJCJC DPCOSCLAGCQ ?R C?AF MAASPPCLAC. "L CTCLR RF?R GLGRG?RCQ ?L ?JJCJC DPCOSCLAW AF?LEC GL ?L GQMJ?RCB N?PR MD RFC NMNSJ?RGML, UFGAF GQ LMR RWNGA?J MD RFC MPGEGL?J NMNSJ?RGML, GQ A?JJCB RFC founder effect. /?RSP?J QCJCARGML, P?LBMK BPGDR, ?LB DMSLBCP CDDCARQ A?L JC?B RM QGELGDGA?LR AF?LECQ GL RFC ECLMKC MD ? NMNSJ?RGML.
#8I;P-2<@E9 +I@E:@GC< F= HL@C@9I@LD

*L RFC C?PJW RUCLRGCRF ACLRSPW, &LEJGQF K?RFCK?RGAG?L (MBDPCW )?PBW ?LB (CPK?L NFWQGAG?L 8GJFCJK 8CGL@CPE QR?RCB RFC NPGLAGNJC MD COSGJG@PGSK RM BCQAPG@C RFC ECLCRGA K?ICSN MD ? NMNSJ?RGML. 5FC RFCMPW, UFGAF J?RCP @CA?KC ILMUL ?Q RFC )?PBW-8CGL@CPE NPGLAGNJC MD COSGJG@PGSK, QR?RCQ RF?R ? NMNSJ?RGMLsQ ?JJCJC ?LB ECLMRWNC DPCOSCLAGCQ ?PC GLFCPCLRJW QR?@JCq SLJCQQ QMKC IGLB MD CTMJSRGML?PW DMPAC GQ ?ARGLE SNML RFC NMNSJ?RGML, LCGRFCP RFC ?JJCJC LMP RFC ECLMRWNGA DPCOSCLAGCQ UMSJB AF?LEC. 5FC )?PBW-8CGL@CPE NPGLAGNJC ?QQSKCQ AMLBGRGMLQ UGRF LM KSR?RGMLQ, KGEP?RGML, CKGEP?RGML, MP QCJCARGTC NPCQQSPC DMP MP ?E?GLQR ECLMRWNC, NJSQ ?L GLDGLGRC NMNSJ?RGML; UFGJC LM NMNSJ?RGML A?L Q?RGQDW RFMQC AMLBGRGMLQ, RFC NPGLAGNJC MDDCPQ ? SQCDSJ KMBCJ ?E?GLQR UFGAF RM AMKN?PC PC?J NMNSJ?RGML AF?LECQ. 8MPIGLE SLBCP RFGQ RFCMPW, NMNSJ?RGML ECLCRGAGQRQ PCNPCQCLR BGDDCPCLR ?JJCJCQ ?Q BGDDCPCLR T?PG?@JCQ GL RFCGP K?RFCK?RGA?J KMBCJQ. 5FC T?PG?@JC N, DMP CV?KNJC, MDRCL PCNPCQCLRQ RFC DPCOSCLAW MD ? N?PRGASJ?P ?JJCJC, Q?W : DMP RFC RP?GR MD WCJJMU GL .CLBCJsQ NC?Q, UFGJC RFC T?PG?@JC O PCNPCQCLRQ RFC DPCOSCLAW MD W ?JJCJCQ RF?R AMLDCP RFC AMJMP EPCCL. *D RFCQC ?PC RFC MLJW RUM NMQQG@JC ?JJCJCQ DMP ? EGTCL JMASQ GL RFC NMNSJ?RGML, N + O = 1. *L MRFCP UMPBQ, ?JJ RFC N ?JJCJCQ ?LB ?JJ RFC O ?JJCJCQ K?IC SN ?JJ MD RFC ?JJCJCQ DMP RF?R JMASQ RF?R ?PC DMSLB GL RFC NMNSJ?RGML. #SR UF?R SJRGK?RCJW GLRCPCQRQ KMQR @GMJMEGQRQ GQ LMR RFC DPCOSCLAGCQ MD BGDDCPCLR ?JJCJCQ, @SR RFC DPCOSCLAGCQ MD RFC PCQSJRGLE ECLMRWNCQ, ILMUL ?Q RFC NMNSJ?RGMLsQ genetic structure, DPMK UFGAF QAGCLRGQRQ A?L QSPKGQC RFC BGQRPG@SRGML MD NFCLMRWNCQ. *D RFC NFCLMRWNC GQ M@QCPTCB, MLJW RFC ECLMRWNC MD RFC FMKMXWEMSQ PCACQQGTC ?JJCJCQ A?L @C ILMUL; RFC A?JASJ?RGMLQ NPMTGBC ?L CQRGK?RC MD RFC PCK?GLGLE ECLMRWNCQ. 4GLAC C?AF GLBGTGBS?J A?PPGCQ RUM ?JJCJCQ NCP ECLC, GD RFC ?JJCJC DPCOSCLAGCQ (N ?LB O) ?PC ILMUL, NPCBGARGLE RFC DPCOSCLAGCQ MD RFCQC ECLMRWNCQ GQ ? QGKNJC K?RFCK?RGA?J A?JASJ?RGML RM BCRCPKGLC RFC NPM@?@GJGRW MD ECRRGLE RFCQC ECLMRWNCQ GD RUM ?JJCJCQ ?PC BP?UL ?R P?LBMK DPMK RFC ECLC NMMJ. 4M GL RFC ?@MTC QACL?PGM, ?L GLBGTGBS?J NC? NJ?LR AMSJB @C NN (::), ?LB RFSQ NPMBSAC WCJJMU NC?Q; NO (:W), ?JQM WCJJMU; MP OO (WW), ?LB RFSQ NPMBSAGLE EPCCL NC?Q (Figure 19.2). *L MRFCP UMPBQ, RFC DPCOSCLAW MD NN GLBGTGBS?JQ GQ QGKNJW N2; RFC DPCOSCLAW MD NO GLBGTGBS?JQ GQ 2NO; ?LB RFC DPCOSCLAW MD OO GLBGTGBS?JQ GQ O2. "LB, ?E?GL, GD N ?LB O ?PC RFC MLJW RUM NMQQG@JC ?JJCJCQ DMP ? EGTCL RP?GR GL RFC NMNSJ?RGML, RFCQC ECLMRWNCQ DPCOSCLAGCQ UGJJ QSK RM MLC: N2 + 2NO + O2 = 1.
1. 4?F?P 4. )?L?LG?, %FG? 4. )?QQ?UG, ?LB /GB?J .. *PQF?GB, t"JJCJC 'PCOSCLAW ?LB .MJCASJ?P (CLMRWNCQ MD "#0 #JMMB (PMSN 4WQRCK GL ? +MPB?LG?L 1MNSJ?RGML,u J6<95(3 6- M,+0*(3 *0,5*,: 7 (2007): 51-58, BMG:10.3923/HKQ.2007.51.58.
504
#(!04%2 19 X 4(% %6/,54)/. /& 0/05,!4)/.3
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)K MI>KQP, SFLIBQ CILTBO @LILO (6) FP ALJFK>KQ LSBO TEFQB (S). )C M = 0.8 >KA N = 0.2 FK > MLMRI>QFLK LC 500 MI>KQP, ELT J>KV FKAFSFAR>IP TLRIA VLR BUMB@Q QL ?B ELJLWVDLRP ALJFK>KQ (66), EBQBOLWVDLRP (6S), >KA ELJLWVDLRP OB@BPPFSB (SS) (LT J>KV MI>KQP TLRIA VLR BUMB@Q QL E>SB SFLIBQ CILTBOP, >KA ELT J>KV TLRIA E>SB TEFQB CILTBOP *L RFCMPW, GD ? NMNSJ?RGML GQ ?R COSGJG@PGSKqRF?R GQ, RFCPC ?PC LM CTMJSRGML?PW DMPACQ ?ARGLE SNML GRqECLCP?RGML ?DRCP ECLCP?RGML UMSJB F?TC RFC Q?KC ECLC NMMJ ?LB ECLCRGA QRPSARSPC, ?LB RFCQC COS?RGMLQ UMSJB ?JJ FMJB RPSC ?JJ MD RFC RGKC. 0D AMSPQC, CTCL )?PBW ?LB 8CGL@CPE PCAMELGXCB RF?R LM L?RSP?J NMNSJ?RGML GQ GKKSLC RM CTMJSRGML. 1MNSJ?RGMLQ GL L?RSPC ?PC AMLQR?LRJW AF?LEGLE GL ECLCRGA K?ICSN BSC RM BPGDR, KSR?RGML, NMQQG@JW KGEP?RGML, ?LB QCJCARGML. "Q ? PCQSJR, RFC MLJW U?W RM BCRCPKGLC RFC CV?AR BGQRPG@SRGML MD NFCLMRWNCQ GL ? NMNSJ?RGML GQ RM EM MSR ?LB AMSLR RFCK. #SR RFC )?PBW-8CGL@CPE NPGLAGNJC EGTCQ QAGCLRGQRQ ? K?RFCK?RGA?J @?QCJGLC MD ? LML-CTMJTGLE NMNSJ?RGML RM UFGAF RFCW A?L AMKN?PC CTMJTGLE NMNSJ?RGMLQ ?LB RFCPC@W GLDCP UF?R CTMJSRGML?PW DMPACQ KGEFR @C ?R NJ?W. *D RFC DPCOSCLAGCQ MD ?JJCJCQ MP ECLMRWNCQ BCTG?RC DPMK RFC T?JSC CVNCARCB DPMK RFC )?PBW-8CGL@CPE COS?RGML, RFCL RFC NMNSJ?RGML GQ CTMJTGLE.
#(!04%2 19 X 4(% %6/,54)/. /& 0/05,!4)/.3
505
6QC RFGQ online calculator (http://openstaxcollege.org/l/hardy-weinberg) RM BCRCPKGLC RFC ECLCRGA QRPSARSPC MD ? NMNSJ?RGML.
19.2 R +FGLC8K@FE "<E<K@:J

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC BGDDCPCLR RWNCQ MD T?PG?RGML GL ? NMNSJ?RGML z &VNJ?GL UFW MLJW FCPGR?@JC T?PG?RGML A?L @C ?ARCB SNML @W L?RSP?J QCJCARGML z %CQAPG@C ECLCRGA BPGDR ?LB RFC @MRRJCLCAI CDDCAR z &VNJ?GL FMU C?AF CTMJSRGML?PW DMPAC A?L GLDJSCLAC RFC ?JJCJC DPCOSCLAGCQ MD ? NMNSJ?RGML *LBGTGBS?JQ MD ? NMNSJ?RGML MDRCL BGQNJ?W BGDDCPCLR NFCLMRWNCQ, MP CVNPCQQ BGDDCPCLR ?JJCJCQ MD ? N?PRGASJ?P ECLC, PCDCPPCB RM ?Q NMJWKMPNFGQKQ. 1MNSJ?RGMLQ UGRF RUM MP KMPC T?PG?RGMLQ MD N?PRGASJ?P AF?P?ARCPGQRGAQ ?PC A?JJCB NMJWKMPNFGA. 5FC BGQRPG@SRGML MD NFCLMRWNCQ ?KMLE GLBGTGBS?JQ, ILMUL ?Q RFC population variation, GQ GLDJSCLACB @W ? LSK@CP MD D?ARMPQ, GLAJSBGLE RFC NMNSJ?RGMLsQ ECLCRGA QRPSARSPC ?LB RFC CLTGPMLKCLR (Figure 19.3). 6LBCPQR?LBGLE RFC QMSPACQ MD ? NFCLMRWNGA T?PG?RGML GL ? NMNSJ?RGML GQ GKNMPR?LR DMP BCRCPKGLGLE FMU ? NMNSJ?RGML UGJJ CTMJTC GL PCQNMLQC RM BGDDCPCLR CTMJSRGML?PW NPCQQSPCQ.
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/?RSP?J QCJCARGML ?LB QMKC MD RFC MRFCP CTMJSRGML?PW DMPACQ A?L MLJW ?AR ML FCPGR?@JC RP?GRQ, L?KCJW ?L MPE?LGQKsQ ECLCRGA AMBC. #CA?SQC ?JJCJCQ ?PC N?QQCB DPMK N?PCLR RM MDDQNPGLE, RFMQC RF?R AMLDCP @CLCDGAG?J RP?GRQ MP @CF?TGMPQ K?W @C QCJCARCB DMP, UFGJC BCJCRCPGMSQ ?JJCJCQ K?W @C QCJCARCB ?E?GLQR. "AOSGPCB RP?GRQ, DMP RFC KMQR N?PR, ?PC LMR FCPGR?@JC. 'MP CV?KNJC, GD ?L ?RFJCRC UMPIQ MSR GL RFC EWK CTCPW B?W, @SGJBGLE SN KSQAJC QRPCLERF, RFC ?RFJCRCsQ MDDQNPGLE UGJJ LMR LCACQQ?PGJW EPMU SN RM @C ? @MBW @SGJBCP. *D RFCPC GQ ? ECLCRGA @?QGQ DMP RFC ?@GJGRW RM PSL D?QR, ML RFC MRFCP F?LB, RFGQ K?W @C N?QQCB RM ? AFGJB.
506
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL RFC BGDDCPCLR U?WQ L?RSP?J QCJCARGML A?L QF?NC NMNSJ?RGMLQ z %CQAPG@C FMU RFCQC BGDDCPCLR DMPACQ A?L JC?B RM BGDDCPCLR MSRAMKCQ GL RCPKQ MD RFC NMNSJ?RGML T?PG?RGML /?RSP?J QCJCARGML MLJW ?ARQ ML RFC NMNSJ?RGMLsQ FCPGR?@JC RP?GRQ: QCJCARGLE DMP @CLCDGAG?J ?JJCJCQ ?LB RFSQ GLAPC?QGLE RFCGP DPCOSCLAW GL RFC NMNSJ?RGML, UFGJC QCJCARGLE ?E?GLQR BCJCRCPGMSQ ?JJCJCQ ?LB RFCPC@W BCAPC?QGLE RFCGP DPCOSCLAWq? NPMACQQ ILMUL ?Q adaptive evolution. /?RSP?J QCJCARGML BMCQ LMR ?AR ML GLBGTGBS?J ?JJCJCQ, FMUCTCP, @SR ML CLRGPC MPE?LGQKQ. "L GLBGTGBS?J K?W A?PPW ? TCPW @CLCDGAG?J ECLMRWNC UGRF ? PCQSJRGLE NFCLMRWNC RF?R, DMP CV?KNJC, GLAPC?QCQ RFC ?@GJGRW RM PCNPMBSAC (DCASLBGRW), @SR GD RF?R Q?KC GLBGTGBS?J ?JQM A?PPGCQ ?L ?JJCJC RF?R PCQSJRQ GL ? D?R?J AFGJBFMMB BGQC?QC, RF?R DCASLBGRW NFCLMRWNC UGJJ LMR @C N?QQCB ML RM RFC LCVR ECLCP?RGML @CA?SQC RFC GLBGTGBS?J UGJJ LMR JGTC RM PC?AF PCNPMBSARGTC ?EC. /?RSP?J QCJCARGML ?ARQ ?R RFC JCTCJ MD RFC GLBGTGBS?J; GR QCJCARQ DMP GLBGTGBS?JQ UGRF EPC?RCP AMLRPG@SRGMLQ RM RFC ECLC NMMJ MD RFC LCVR ECLCP?RGML, ILMUL ?Q ?L MPE?LGQKsQ evolutionary (Darwinian) fitness. 'GRLCQQ GQ MDRCL OS?LRGDG?@JC ?LB GQ KC?QSPCB @W QAGCLRGQRQ GL RFC DGCJB. )MUCTCP, GR GQ LMR RFC ?@QMJSRC DGRLCQQ MD ?L GLBGTGBS?J RF?R AMSLRQ, @SR P?RFCP FMU GR AMKN?PCQ RM RFC MRFCP MPE?LGQKQ GL RFC NMNSJ?RGML. 5FGQ AMLACNR, A?JJCB relative fitness, ?JJMUQ PCQC?PAFCPQ RM BCRCPKGLC UFGAF GLBGTGBS?JQ ?PC AMLRPG@SRGLE ?BBGRGML?J MDDQNPGLE RM RFC LCVR ECLCP?RGML, ?LB RFSQ, FMU RFC NMNSJ?RGML KGEFR CTMJTC. 5FCPC ?PC QCTCP?J U?WQ QCJCARGML A?L ?DDCAR NMNSJ?RGML T?PG?RGML: QR?@GJGXGLE QCJCARGML, BGPCARGML?J QCJCARGML, BGTCPQGDWGLE QCJCARGML, DPCOSCLAW-BCNCLBCLR QCJCARGML, ?LB QCVS?J QCJCARGML. "Q L?RSP?J QCJCARGML GLDJSCLACQ RFC ?JJCJC DPCOSCLAGCQ GL ? NMNSJ?RGML, GLBGTGBS?JQ A?L CGRFCP @CAMKC KMPC MP JCQQ ECLCRGA?JJW QGKGJ?P ?LB RFC NFCLMRWNCQ BGQNJ?WCB A?L @CAMKC KMPC QGKGJ?P MP KMPC BGQN?P?RC.
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*D L?RSP?J QCJCARGML D?TMPQ ?L ?TCP?EC NFCLMRWNC, QCJCARGLE ?E?GLQR CVRPCKC T?PG?RGML, RFC NMNSJ?RGML UGJJ SLBCPEM stabilizing selection (Figure 19.8). *L ? NMNSJ?RGML MD KGAC RF?R JGTC GL RFC UMMBQ, DMP CV?KNJC, L?RSP?J QCJCARGML GQ JGICJW RM D?TMP GLBGTGBS?JQ RF?R @CQR @JCLB GL UGRF RFC DMPCQR DJMMP ?LB ?PC JCQQ JGICJW
512
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8FCL RFC CLTGPMLKCLR AF?LECQ, NMNSJ?RGMLQ UGJJ MDRCL SLBCPEM directional selection (Figure 19.8), UFGAF QCJCARQ DMP NFCLMRWNCQ ?R MLC CLB MD RFC QNCARPSK MD CVGQRGLE T?PG?RGML. " AJ?QQGA CV?KNJC MD RFGQ RWNC MD QCJCARGML GQ RFC CTMJSRGML MD RFC NCNNCPCB KMRF GL CGEFRCCLRF- ?LB LGLCRCCLRF-ACLRSPW &LEJ?LB. 1PGMP RM RFC *LBSQRPG?J 3CTMJSRGML, RFC KMRFQ UCPC NPCBMKGL?RCJW JGEFR GL AMJMP, UFGAF ?JJMUCB RFCK RM @JCLB GL UGRF RFC JGEFR-AMJMPCB RPCCQ ?LB JGAFCLQ GL RFCGP CLTGPMLKCLR. #SR ?Q QMMR @CE?L QNCUGLE DPMK D?ARMPGCQ, RFC RPCCQ @CA?KC B?PICLCB, ?LB RFC JGEFR-AMJMPCB KMRFQ @CA?KC C?QGCP DMP NPCB?RMPW @GPBQ RM QNMR. 0TCP RGKC, RFC DPCOSCLAW MD RFC KCJ?LGA DMPK MD RFC KMRF GLAPC?QCB @CA?SQC RFCW F?B ? FGEFCP QSPTGT?J P?RC GL F?@GR?RQ ?DDCARCB @W ?GP NMJJSRGML @CA?SQC RFCGP B?PICP AMJMP?RGML @JCLBCB UGRF RFC QMMRW RPCCQ. 4GKGJ?PJW, RFC FWNMRFCRGA?J KMSQC NMNSJ?RGML K?W CTMJTC RM R?IC ML ? BGDDCPCLR AMJMP?RGML GD QMKCRFGLE UCPC RM A?SQC RFC DMPCQR DJMMP UFCPC RFCW JGTC RM AF?LEC AMJMP. 5FC PCQSJR MD RFGQ RWNC MD QCJCARGML GQ ? QFGDR GL RFC NMNSJ?RGMLsQ ECLCRGA T?PG?LAC RMU?PB RFC LCU, DGR NFCLMRWNC.
*L QAGCLAC, QMKCRGKCQ RFGLEQ ?PC @CJGCTCB RM @C RPSC, ?LB RFCL LCU GLDMPK?RGML AMKCQ RM JGEFR RF?R AF?LECQ MSP SLBCPQR?LBGLE. 5FC QRMPW MD RFC NCNNCPCB KMRF GQ ?L CV?KNJC: RFC D?ARQ @CFGLB RFC QCJCARGML RMU?PB B?PICP KMRFQ F?TC PCACLRJW @CCL A?JJCB GLRM OSCQRGML. 3C?B RFGQ article (http://openstaxcollege.org/l/peppered_moths) RM JC?PL KMPC.
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4MKCRGKCQ RUM MP KMPC BGQRGLAR NFCLMRWNCQ A?L C?AF F?TC RFCGP ?BT?LR?ECQ ?LB @C QCJCARCB DMP @W L?RSP?J QCJCARGML, UFGJC RFC GLRCPKCBG?RC NFCLMRWNCQ ?PC, ML ?TCP?EC, JCQQ DGR. ,LMUL ?Q diversifying selection (Figure 19.8), RFGQ GQ QCCL GL K?LW NMNSJ?RGMLQ MD ?LGK?JQ RF?R F?TC KSJRGNJC K?JC DMPKQ. -?PEC, BMKGL?LR ?JNF? K?JCQ M@R?GL K?RCQ @W @PSRC DMPAC, UFGJC QK?JJ K?JCQ A?L QLC?I GL DMP DSPRGTC AMNSJ?RGMLQ UGRF RFC DCK?JCQ GL ?L ?JNF? K?JCsQ RCPPGRMPW. *L RFGQ A?QC, @MRF RFC ?JNF? K?JCQ ?LB RFC tQLC?IGLEu K?JCQ UGJJ @C QCJCARCB DMP, @SR KCBGSK-QGXCB K?JCQ, UFGAF A?LsR MTCPR?IC RFC ?JNF? K?JCQ ?LB ?PC RMM @GE RM QLC?I AMNSJ?RGMLQ, ?PC QCJCARCB ?E?GLQR. %GTCPQGDWGLE QCJCARGML A?L ?JQM MAASP UFCL CLTGPMLKCLR?J AF?LECQ D?TMP GLBGTGBS?JQ ML CGRFCP CLB MD RFC NFCLMRWNGA QNCARPSK. *K?EGLC ? NMNSJ?RGML MD KGAC JGTGLE ?R RFC @C?AF UFCPC RFCPC GQ JGEFR-AMJMPCB Q?LB GLRCPQNCPQCB UGRF N?RAFCQ MD R?JJ EP?QQ. *L RFGQ QACL?PGM, JGEFR-AMJMPCB KGAC RF?R @JCLB GL UGRF RFC Q?LB UMSJB @C D?TMPCB, ?Q UCJJ ?Q B?PI-AMJMPCB KGAC RF?R A?L FGBC GL RFC EP?QQ. .CBGSK-AMJMPCB KGAC, ML RFC MRFCP F?LB, UMSJB LMR @JCLB GL UGRF CGRFCP RFC EP?QQ MP RFC Q?LB, ?LB UMSJB RFSQ @C KMPC JGICJW RM @C C?RCL @W NPCB?RMPQ. 5FC PCQSJR MD RFGQ RWNC MD QCJCARGML GQ GLAPC?QCB ECLCRGA T?PG?LAC ?Q RFC NMNSJ?RGML @CAMKCQ KMPC BGTCPQC.
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514
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*L RFGQ QACL?PGM, MP?LEC K?JCQ UGJJ @C D?TMPCB @W L?RSP?J QCJCARGML UFCL RFC NMNSJ?RGML GQ BMKGL?RCB @W @JSC K?JCQ, @JSC K?JCQ UGJJ RFPGTC UFCL RFC NMNSJ?RGML GQ KMQRJW WCJJMU K?JCQ, ?LB WCJJMU K?JCQ UGJJ @C QCJCARCB DMP UFCL MP?LEC K?JCQ ?PC RFC KMQR NMNSJMSQ. "Q ? PCQSJR, NMNSJ?RGMLQ MD QGBC-@JMRAFCB JGX?PBQ AWAJC GL RFC BGQRPG@SRGML MD RFCQC NFCLMRWNCQqGL MLC ECLCP?RGML, MP?LEC KGEFR @C NPCBMKGL?LR, ?LB RFCL WCJJMU K?JCQ UGJJ @CEGL RM PGQC GL DPCOSCLAW. 0LAC WCJJMU K?JCQ K?IC SN ? K?HMPGRW MD RFC NMNSJ?RGML, @JSC K?JCQ UGJJ @C QCJCARCB DMP. 'GL?JJW, UFCL @JSC K?JCQ @CAMKC AMKKML, MP?LEC K?JCQ UGJJ MLAC ?E?GL @C D?TMPCB. /CE?RGTC DPCOSCLAW-BCNCLBCLR QCJCARGML QCPTCQ RM GLAPC?QC RFC NMNSJ?RGMLsQ ECLCRGA T?PG?LAC @W QCJCARGLE DMP P?PC NFCLMRWNCQ, UFCPC?Q NMQGRGTC DPCOSCLAW-BCNCLBCLR QCJCARGML SQS?JJW BCAPC?QCQ ECLCRGA T?PG?LAC @W QCJCARGLE DMP AMKKML NFCLMRWNCQ.
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.?JCQ ?LB DCK?JCQ MD ACPR?GL QNCAGCQ ?PC MDRCL OSGRC BGDDCPCLR DPMK MLC ?LMRFCP GL U?WQ @CWMLB RFC PCNPMBSARGTC MPE?LQ. .?JCQ ?PC MDRCL J?PECP, DMP CV?KNJC, ?LB BGQNJ?W K?LW CJ?@MP?RC AMJMPQ ?LB ?BMPLKCLRQ, JGIC RFC NC?AMAIsQ R?GJ, UFGJC DCK?JCQ RCLB RM @C QK?JJCP ?LB BSJJCP GL BCAMP?RGML. 4SAF BGDDCPCLACQ ?PC ILMUL ?Q sexual dimorphisms (Figure 19.10), UFGAF ?PGQC DPMK RFC D?AR RF?R GL K?LW NMNSJ?RGMLQ, N?PRGASJ?PJW ?LGK?J NMNSJ?RGMLQ, RFCPC GQ KMPC T?PG?LAC GL RFC PCNPMBSARGTC QSAACQQ MD RFC K?JCQ RF?L RFCPC GQ MD RFC DCK?JCQ. 5F?R GQ, QMKC K?JCQqMDRCL RFC @GEECP, QRPMLECP, MP KMPC BCAMP?RCB K?JCQqECR RFC T?QR K?HMPGRW MD RFC RMR?J K?RGLEQ, UFGJC MRFCPQ PCACGTC LMLC. 5FGQ A?L MAASP @CA?SQC RFC K?JCQ ?PC @CRRCP ?R DGEFRGLE MDD MRFCP K?JCQ, MP @CA?SQC DCK?JCQ UGJJ AFMMQC RM K?RC UGRF RFC @GEECP MP KMPC BCAMP?RCB K?JCQ. *L CGRFCP A?QC, RFGQ T?PG?RGML GL PCNPMBSARGTC QSAACQQ ECLCP?RCQ ? QRPMLE QCJCARGML NPCQQSPC ?KMLE K?JCQ RM ECR RFMQC K?RGLEQ, PCQSJRGLE GL RFC CTMJSRGML MD @GEECP @MBW QGXC ?LB CJ?@MP?RC MPL?KCLRQ RM ECR RFC DCK?JCQs ?RRCLRGML. 'CK?JCQ, ML RFC MRFCP F?LB, RCLB RM ECR ? F?LBDSJ MD QCJCARCB K?RGLEQ; RFCPCDMPC, RFCW ?PC KMPC JGICJW RM QCJCAR KMPC BCQGP?@JC K?JCQ. 4CVS?J BGKMPNFGQK T?PGCQ UGBCJW ?KMLE QNCAGCQ, MD AMSPQC, ?LB QMKC QNCAGCQ ?PC CTCL QCV-PMJC PCTCPQCB. *L QSAF A?QCQ, DCK?JCQ RCLB RM F?TC ? EPC?RCP T?PG?LAC GL RFCGP PCNPMBSARGTC QSAACQQ RF?L K?JCQ ?LB ?PC AMPPCQNMLBGLEJW QCJCARCB DMP RFC @GEECP @MBW QGXC ?LB CJ?@MP?RC RP?GRQ SQS?JJW AF?P?ARCPGQRGA MD K?JCQ.
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*L 1915, @GMJMEGQR 3ML?JB 'GQFCP NPMNMQCB ?LMRFCP KMBCJ MD QCVS?J QCJCARGML: RFC Fisherian runaway model (http://openstaxcollege.org/l/sexual_select) , UFGAF QSEECQRQ RF?R QCJCARGML MD ACPR?GL RP?GRQ GQ ? PCQSJR MD QCVS?J NPCDCPCLAC. *L @MRF RFC F?LBGA?N NPGLAGNJC ?LB RFC EMMB ECLCQ FWNMRFCQGQ, RFC RP?GR GQ Q?GB RM @C ?L honest signal MD RFC K?JCQs OS?JGRW, RFSQ EGTGLE DCK?JCQ ? U?W RM DGLB RFC DGRRCQR K?RCQq K?JCQ RF?R UGJJ N?QQ RFC @CQR ECLCQ RM RFCGP MDDQNPGLE.
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516
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5FGQ @CC ?LB E*/05(*,( DJMUCP (Figure 20.1) AMSJB LMR JMMI KMPC BGDDCPCLR, WCR RFCW ?PC PCJ?RCB, ?Q ?PC ?JJ JGTGLE MPE?LGQKQ ML &?PRF. #W DMJJMUGLE N?RFU?WQ MD QGKGJ?PGRGCQ ?LB AF?LECQq@MRF TGQG@JC ?LB ECLCRGAqQAGCLRGQRQ QCCI RM K?N RFC CTMJSRGML?PW N?QR MD FMU JGDC BCTCJMNCB DPMK QGLEJC-ACJJCB MPE?LGQKQ RM RFC RPCKCLBMSQ AMJJCARGML MD APC?RSPCQ RF?R F?TC ECPKGL?RCB, AP?UJCB, DJM?RCB, QU?K, DJMUL, ?LB U?JICB ML RFGQ NJ?LCR.
20.1 R *I>8E@Q@E> '@=< FE 8IK?

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %GQASQQ RFC LCCB DMP ? AMKNPCFCLQGTC AJ?QQGDGA?RGML QWQRCK z -GQR RFC BGDDCPCLR JCTCJQ MD RFC R?VMLMKGA AJ?QQGDGA?RGML QWQRCK z %CQAPG@C FMU QWQRCK?RGAQ ?LB R?VMLMKW PCJ?RC RM NFWJMECLW z %GQASQQ RFC AMKNMLCLRQ ?LB NSPNMQC MD ? NFWJMECLCRGA RPCC
522
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*L QAGCLRGDGA RCPKQ, RFC CTMJSRGML?PW FGQRMPW ?LB PCJ?RGMLQFGN MD ?L MPE?LGQK MP EPMSN MD MPE?LGQKQ GQ A?JJCB NFWJMECLW. Phylogeny BCQAPG@CQ RFC PCJ?RGMLQFGNQ MD ?L MPE?LGQK, QSAF ?Q DPMK UFGAF MPE?LGQKQ GR GQ RFMSEFR RM F?TC CTMJTCB, RM UFGAF QNCAGCQ GR GQ KMQR AJMQCJW PCJ?RCB, ?LB QM DMPRF. 1FWJMECLCRGA PCJ?RGMLQFGNQ NPMTGBC GLDMPK?RGML ML QF?PCB ?LACQRPW @SR LMR LCACQQ?PGJW ML FMU MPE?LGQKQ ?PC QGKGJ?P MP BGDDCPCLR.
+?PCF><E<K@: /I<<J
4AGCLRGQRQ SQC ? RMMJ A?JJCB ? NFWJMECLCRGA RPCC RM QFMU RFC CTMJSRGML?PW N?RFU?WQ ?LB AMLLCARGMLQ ?KMLE MPE?LGQKQ. " phylogenetic tree GQ ? BG?EP?K SQCB RM PCDJCAR CTMJSRGML?PW PCJ?RGMLQFGNQ ?KMLE MPE?LGQKQ MP EPMSNQ MD MPE?LGQKQ. 4AGCLRGQRQ AMLQGBCP NFWJMECLCRGA RPCCQ RM @C ? FWNMRFCQGQ MD RFC CTMJSRGML?PW N?QR QGLAC MLC A?LLMR EM @?AI RM AMLDGPK RFC NPMNMQCB PCJ?RGMLQFGNQ. *L MRFCP UMPBQ, ? tRPCC MD JGDCu A?L @C AMLQRPSARCB RM GJJSQRP?RC UFCL BGDDCPCLR MPE?LGQKQ CTMJTCB ?LB RM QFMU RFC PCJ?RGMLQFGNQ ?KMLE BGDDCPCLR MPE?LGQKQ (Figure 20.2). 6LJGIC ? R?VMLMKGA AJ?QQGDGA?RGML BG?EP?K, ? NFWJMECLCRGA RPCC A?L @C PC?B JGIC ? K?N MD CTMJSRGML?PW FGQRMPW. .?LW NFWJMECLCRGA RPCCQ F?TC ? QGLEJC JGLC?EC ?R RFC @?QC PCNPCQCLRGLE ? AMKKML ?LACQRMP. 4AGCLRGQRQ A?JJ QSAF RPCCQ rooted, UFGAF KC?LQ RFCPC GQ ? QGLEJC ?LACQRP?J JGLC?EC (RWNGA?JJW BP?UL DPMK RFC @MRRMK MP JCDR) RM UFGAF ?JJ MPE?LGQKQ PCNPCQCLRCB GL RFC BG?EP?K PCJ?RC. /MRGAC GL RFC PMMRCB NFWJMECLCRGA RPCC RF?R RFC RFPCC BMK?GLQq #?ARCPG?, "PAF?C?, ?LB &SI?PW?qBGTCPEC DPMK ? QGLEJC NMGLR ?LB @P?LAF MDD. 5FC QK?JJ @P?LAF RF?R NJ?LRQ ?LB ?LGK?JQ (GLAJSBGLE FSK?LQ) MAASNW GL RFGQ BG?EP?K QFMUQ FMU PCACLR ?LB KGLGQASJC RFCQC EPMSNQ ?PC AMKN?PCB UGRF MRFCP MPE?LGQKQ. 6LPMMRCB RPCCQ BMLsR QFMU ? AMKKML ?LACQRMP @SR BM QFMU PCJ?RGMLQFGNQ ?KMLE QNCAGCQ.
!@>LI< 20.2 "LQE LC QEBPB MEVILDBKBQF@ QOBBP PELTP QEB OBI>QFLKPEFM LC QEB QEOBB ALJ>FKP LC IFCBu">@QBOF>, !O@E>B>, >KA %RH>OV>u?RQ QEB (>) OLLQBA QOBB >QQBJMQP QL FABKQFCV TEBK S>OFLRP PMB@FBP AFSBODBA COLJ > @LJJLK >K@BPQLO TEFIB QEB (?) RKOLLQBA QOBB ALBP KLQ. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V %OF@ '>?>)
*L ? PMMRCB RPCC, RFC @P?LAFGLE GLBGA?RCQ CTMJSRGML?PW PCJ?RGMLQFGNQ (Figure 20.3). 5FC NMGLR UFCPC ? QNJGR MAASPQ, A?JJCB ? branch point, PCNPCQCLRQ UFCPC ? QGLEJC JGLC?EC CTMJTCB GLRM ? BGQRGLAR LCU MLC. " JGLC?EC RF?R CTMJTCB C?PJW DPMK RFC PMMR ?LB PCK?GLQ SL@P?LAFCB GQ A?JJCB basal taxon. 8FCL RUM JGLC?ECQ QRCK DPMK RFC Q?KC @P?LAF NMGLR, RFCW ?PC A?JJCB sister taxa. " @P?LAF UGRF KMPC RF?L RUM JGLC?ECQ GQ A?JJCB ? polytomy ?LB QCPTCQ RM GJJSQRP?RC UFCPC QAGCLRGQRQ F?TC LMR BCDGLGRGTCJW BCRCPKGLCB ?JJ MD RFC PCJ?RGMLQFGNQ. *R GQ GKNMPR?LR RM LMRC RF?R ?JRFMSEF QGQRCP R?V? ?LB NMJWRMKW BM QF?PC ?L ?LACQRMP, GR BMCQ LMR KC?L RF?R RFC EPMSNQ MD MPE?LGQKQ QNJGR MP CTMJTCB DPMK C?AF MRFCP. 0PE?LGQKQ GL RUM R?V? K?W F?TC QNJGR ?N?PR ?R ? QNCAGDGA @P?LAF NMGLR, @SR LCGRFCP R?V? E?TC PGQC RM RFC MRFCP.
!@>LI< 20.3 4EB OLLQ LC > MEVILDBKBQF@ QOBB FKAF@>QBP QE>Q >K >K@BPQO>I IFKB>DB D>SB OFPB QL >II LOD>KFPJP LK QEB QOBB. ! ?O>K@E MLFKQ FKAF@>QBP TEBOB QTL IFKB>DBP AFSBODBA. ! IFKB>DB QE>Q BSLISBA B>OIV >KA OBJ>FKP RK?O>K@EBA FP > ?>P>I Q>ULK. 7EBK QTL IFKB>DBP PQBJ COLJ QEB P>JB ?O>K@E MLFKQ, QEBV >OB PFPQBO Q>U>. ! ?O>K@E TFQE JLOB QE>K QTL IFKB>DBP FP > MLIVQLJV.
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523
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tRPCC MD JGDC.u *LDMPK?RGML GQ SQCB RM MPE?LGXC ?LB AJ?QQGDW MPE?LGQKQ @?QCB ML CTMJSRGML?PW PCJ?RGMLQFGNQ GL ? QAGCLRGDGA DGCJB A?JJCB systematics. %?R? K?W @C AMJJCARCB DPMK DMQQGJQ, DPMK QRSBWGLE RFC QRPSARSPC MD @MBW N?PRQ MP KMJCASJCQ SQCB @W ?L MPE?LGQK, ?LB @W %/" ?L?JWQGQ. #W AMK@GLGLE B?R? DPMK K?LW QMSPACQ, QAGCLRGQRQ A?L NSR RMECRFCP RFC NFWJMECLW MD ?L MPE?LGQK; QGLAC NFWJMECLCRGA RPCCQ ?PC FWNMRFCQCQ, RFCW UGJJ AMLRGLSC RM AF?LEC ?Q LCU RWNCQ MD JGDC ?PC BGQAMTCPCB ?LB LCU GLDMPK?RGML GQ JC?PLCB.
'@D@K8K@FEJ F= +?PCF><E<K@: /I<<J

*R K?W @C C?QW RM ?QQSKC RF?R KMPC AJMQCJW PCJ?RCB MPE?LGQKQ JMMI KMPC ?JGIC, ?LB UFGJC RFGQ GQ MDRCL RFC A?QC, GR GQ LMR ?JU?WQ RPSC. *D RUM AJMQCJW PCJ?RCB JGLC?ECQ CTMJTCB SLBCP QGELGDGA?LRJW T?PGCB QSPPMSLBGLEQ MP ?DRCP RFC CTMJSRGML MD ? K?HMP LCU ?B?NR?RGML, GR GQ NMQQG@JC DMP RFC RUM EPMSNQ RM ?NNC?P KMPC BGDDCPCLR RF?L MRFCP EPMSNQ RF?R ?PC LMR ?Q AJMQCJW PCJ?RCB. 'MP CV?KNJC, RFC NFWJMECLCRGA RPCC GL Figure 20.4 QFMUQ RF?R JGX?PBQ ?LB P?@@GRQ @MRF F?TC ?KLGMRGA CEEQ, UFCPC?Q DPMEQ BM LMR; WCR JGX?PBQ ?LB DPMEQ ?NNC?P KMPC QGKGJ?P RF?L JGX?PBQ ?LB P?@@GRQ.
!@>LI< 20.4 4EFP I>AABO-IFHB MEVILDBKBQF@ QOBB LC SBOQB?O>QBP FP OLLQBA ?V >K LOD>KFPJ QE>Q I>@HBA > SBOQB?O>I @LIRJK. !Q B>@E ?O>K@E MLFKQ, LOD>KFPJP TFQE AFCCBOBKQ @E>O>@QBOP >OB MI>@BA FK AFCCBOBKQ DOLRMP ?>PBA LK QEB @E>O>@QBOFPQF@P QEBV PE>OB.
"LMRFCP ?QNCAR MD NFWJMECLCRGA RPCCQ GQ RF?R, SLJCQQ MRFCPUGQC GLBGA?RCB, RFC @P?LAFCQ BM LMR ?AAMSLR DMP JCLERF MD RGKC, MLJW RFC CTMJSRGML?PW MPBCP. *L MRFCP UMPBQ, RFC JCLERF MD ? @P?LAF BMCQ LMR RWNGA?JJW KC?L KMPC RGKC N?QQCB, LMP BMCQ ? QFMPR @P?LAF KC?L JCQQ RGKC N?QQCBq SLJCQQ QNCAGDGCB ML RFC BG?EP?K. 'MP CV?KNJC, GL Figure 20.4, RFC RPCC BMCQ LMR GLBGA?RC FMU KSAF RGKC N?QQCB @CRUCCL RFC CTMJSRGML MD ?KLGMRGA CEEQ ?LB F?GP. 8F?R RFC RPCC BMCQ QFMU GQ RFC MPBCP GL UFGAF RFGLEQ RMMI NJ?AC. "E?GL SQGLE Figure 20.4, RFC RPCC QFMUQ RF?R RFC MJBCQR RP?GR GQ RFC TCPRC@P?J AMJSKL, DMJJMUCB @W FGLECB H?UQ, ?LB QM DMPRF. 3CKCK@CP RF?R ?LW NFWJMECLCRGA RPCC GQ ? N?PR MD RFC EPC?RCP UFMJC, ?LB JGIC ? PC?J RPCC, GR BMCQ LMR EPMU GL MLJW MLC BGPCARGML ?DRCP ? LCU @P?LAF BCTCJMNQ. 4M, DMP RFC MPE?LGQKQ GL Figure 20.4, HSQR @CA?SQC ? TCPRC@P?J AMJSKL CTMJTCB BMCQ LMR KC?L RF?R GLTCPRC@P?RC CTMJSRGML AC?QCB, GR MLJW KC?LQ RF?R ? LCU @P?LAF DMPKCB. "JQM, EPMSNQ RF?R ?PC LMR AJMQCJW PCJ?RCB, @SR CTMJTC SLBCP QGKGJ?P AMLBGRGMLQ, K?W ?NNC?P KMPC NFCLMRWNGA?JJW QGKGJ?P RM C?AF MRFCP RF?L RM ? AJMQC PCJ?RGTC.
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Taxonomy (UFGAF JGRCP?JJW KC?LQ t?PP?LECKCLR J?Uu) GQ RFC QAGCLAC MD AJ?QQGDWGLE MPE?LGQKQ RM AMLQRPSAR GLRCPL?RGML?JJW QF?PCB AJ?QQGDGA?RGML QWQRCKQ UGRF C?AF MPE?LGQK NJ?ACB GLRM KMPC ?LB KMPC GLAJSQGTC EPMSNGLEQ. 5FGLI ?@MSR FMU ? EPMACPW QRMPC GQ MPE?LGXCB. 0LC J?PEC QN?AC GQ BGTGBCB GLRM BCN?PRKCLRQ, QSAF ?Q NPMBSAC, B?GPW, ?LB KC?RQ. 5FCL C?AF BCN?PRKCLR DSPRFCP BGTGBCQ GLRM ?GQJCQ, RFCL C?AF ?GQJC GLRM A?RCEMPGCQ ?LB @P?LBQ, ?LB RFCL DGL?JJW ? QGLEJC NPMBSAR. 5FGQ MPE?LGX?RGML DPMK J?PECP RM QK?JJCP, KMPC QNCAGDGA A?RCEMPGCQ GQ A?JJCB ? FGCP?PAFGA?J QWQRCK. 5FC R?VMLMKGA AJ?QQGDGA?RGML QWQRCK (?JQM A?JJCB RFC -GLL?C?L QWQRCK ?DRCP GRQ GLTCLRMP, $?PJ -GLL?CSQ, ? 4UCBGQF @MR?LGQR, XMMJMEGQR, ?LB NFWQGAG?L) SQCQ ? FGCP?PAFGA?J KMBCJ. .MTGLE DPMK RFC NMGLR MD MPGEGL, RFC EPMSNQ @CAMKC KMPC QNCAGDGA, SLRGJ MLC @P?LAF CLBQ ?Q ? QGLEJC QNCAGCQ. 'MP CV?KNJC, ?DRCP RFC AMKKML @CEGLLGLE MD ?JJ JGDC, QAGCLRGQRQ BGTGBC MPE?LGQKQ GLRM RFPCC J?PEC A?RCEMPGCQ A?JJCB ? BMK?GL: #?ARCPG?, "PAF?C?, ?LB &SI?PW?. 8GRFGL C?AF BMK?GL GQ ? QCAMLB A?RCEMPW A?JJCB ? kingdom. "DRCP IGLEBMKQ, RFC QS@QCOSCLR A?RCEMPGCQ MD GLAPC?QGLE QNCAGDGAGRW ?PC: phylum, class, order, family, genus, ?LB species (Figure 20.5).
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525
!@>LI< 20.5 4EB Q>ULKLJF@ @I>PPFCF@>QFLK PVPQBJ RPBP > EFBO>O@EF@>I JLABI QL LOD>KFWB IFSFKD LOD>KFPJP FKQL FK@OB>PFKDIV PMB@FCF@ @>QBDLOFBP. 4EB @LJJLK ALD, C(50: 3<7<: -(4030(90:, FP > PR?PMB@FBP LC C(50: 3<7<:, TEF@E >IPL FK@IRABP QEB TLIC >KA AFKDL. (@OBAFQ xALDy: JLAFCF@>QFLK LC TLOH ?V *>KKBHB 6OBRDABKEFI)
5FC IGLEBMK "LGK?JG? QRCKQ DPMK RFC &SI?PW? BMK?GL. 'MP RFC AMKKML BME, RFC AJ?QQGDGA?RGML JCTCJQ UMSJB @C ?Q QFMUL GL Figure 20.5. 5FCPCDMPC, RFC DSJJ L?KC MD ?L MPE?LGQK RCAFLGA?JJW F?Q CGEFR RCPKQ. 'MP RFC BME, GR GQ: &SI?PW?, "LGK?JG?, $FMPB?R?, .?KK?JG?, $?PLGTMP?, $?LGB?C, C(50:, ?LB 3<7<:. /MRGAC RF?R C?AF L?KC GQ A?NGR?JGXCB CVACNR DMP QNCAGCQ, ?LB RFC ECLSQ ?LB QNCAGCQ L?KCQ ?PC GR?JGAGXCB. 4AGCLRGQRQ ECLCP?JJW PCDCP RM ?L MPE?LGQK MLJW @W GRQ ECLSQ ?LB QNCAGCQ, UFGAF GQ GRQ RUM-UMPB QAGCLRGDGA L?KC, GL UF?R GQ A?JJCB binomial nomenclature. 5FCPCDMPC, RFC QAGCLRGDGA L?KC MD RFC BME GQ C(50: 3<7<:. 5FC L?KC ?R C?AF JCTCJ GQ ?JQM A?JJCB ? taxon. *L MRFCP UMPBQ, BMEQ ?PC GL MPBCP $?PLGTMP?. $?PLGTMP? GQ RFC L?KC MD RFC R?VML ?R RFC MPBCP JCTCJ; $?LGB?C GQ RFC R?VML ?R RFC D?KGJW JCTCJ, ?LB QM DMPRF. 0PE?LGQKQ ?JQM F?TC ? AMKKML L?KC RF?R NCMNJC RWNGA?JJW SQC, GL RFGQ A?QC, BME. /MRC RF?R RFC BME GQ ?BBGRGML?JJW ? QS@QNCAGCQ: RFC t-(4030(90:u GL C(50: 3<7<: -(4030(90:. 4S@QNCAGCQ ?PC KCK@CPQ MD RFC Q?KC QNCAGCQ RF?R ?PC A?N?@JC MD K?RGLE ?LB PCNPMBSAGLE TG?@JC MDDQNPGLE, @SR RFCW ?PC AMLQGBCPCB QCN?P?RC QS@QNCAGCQ BSC RM ECMEP?NFGA MP @CF?TGMP?J GQMJ?RGML MP MRFCP D?ARMPQ. Figure 20.6 QFMUQ FMU RFC JCTCJQ KMTC RMU?PB QNCAGDGAGRW UGRF MRFCP MPE?LGQKQ. /MRGAC FMU RFC BME QF?PCQ ? BMK?GL UGRF RFC UGBCQR BGTCPQGRW MD MPE?LGQKQ, GLAJSBGLE NJ?LRQ ?LB @SRRCPDJGCQ. "R C?AF QS@JCTCJ, RFC MPE?LGQKQ @CAMKC KMPC QGKGJ?P @CA?SQC RFCW ?PC KMPC AJMQCJW PCJ?RCB. )GQRMPGA?JJW,
526
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!@>LI< 20.6 !Q B>@E PR?IBSBI FK QEB Q>ULKLJF@ @I>PPFCF@>QFLK PVPQBJ, LOD>KFPJP ?B@LJB JLOB PFJFI>O. $LDP >KA TLISBP >OB QEB P>JB PMB@FBP ?B@>RPB QEBV @>K ?OBBA >KA MOLAR@B SF>?IB LCCPMOFKD, ?RQ QEBV >OB AFCCBOBKQ BKLRDE QL ?B @I>PPFCFBA >P AFCCBOBKQ PR?PMB@FBP. (@OBAFQ xMI>KQy: JLAFCF@>QFLK LC TLOH ?V "?BOAR@ET>I"/&IF@HO; @OBAFQ xFKPB@Qy: JLAFCF@>QFLK LC TLOH ?V *LK 3RIIFS>K; @OBAFQ xCFPEy: JLAFCF@>QFLK LC TLOH ?V #EOFPQF>K -BEICiEOBO; @OBAFQ xO>??FQy: JLAFCF@>QFLK LC TLOH ?V !FA>K 7LGQ>P; @OBAFQ x@>Qy: JLAFCF@>QFLK LC TLOH ?V *LK>QE>K ,FA?B@H; @OBAFQ xCLUy: JLAFCF@>QFLK LC TLOH ?V +BSFK ">@EBO, .03; @OBAFQ xG>@H>Iy: JLAFCF@>QFLK LC TLOH ?V 4ELJ>P !. (BOJ>KK, .")), 53'3; @OBAFQ xTLICy: JLAFCF@>QFLK LC TLOH ?V 2L?BOQ $BT>O; @OBAFQ xALDy: JLAFCF@>QFLK LC TLOH ?V "AFDFQ>I=FJ>DB=C>K"/&IF@HO)
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527
7GQGR RFGQ website (http://openstaxcollege.org/l/classify_life) RM AJ?QQGDW RFPCC MPE?LGQKQq@C?P, MPAFGB, ?LB QC? ASASK@CPq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
20.2 R D<K<ID@E@E> MFCLK@FE8IP -<C8K@FEJ?@GJ

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528
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530
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534
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5FC PCAMELGRGML MD RFC GKNMPR?LAC MD )(5, CQNCAG?JJW GL RFC CTMJSRGML MD NPMI?PWMRCQ, F?Q A?SQCB QMKC RM NPMNMQC ?@?LBMLGLE RFC AJ?QQGA tRPCC MD JGDCu KMBCJ. *L 1999, 8. 'MPB %MMJGRRJC NPMNMQCB ? NFWJMECLCRGA KMBCJ RF?R PCQCK@JCQ ? UC@ MP ? LCRUMPI KMPC RF?L ? RPCC. 5FC FWNMRFCQGQ GQ RF?R CSI?PWMRCQ CTMJTCB LMR DPMK ? QGLEJC NPMI?PWMRGA ?LACQRMP, @SR DPMK ? NMMJ MD K?LW QNCAGCQ RF?R UCPC QF?PGLE ECLCQ @W )(5 KCAF?LGQKQ. "Q QFMUL GL Figure 20.16a, QMKC GLBGTGBS?J NPMI?PWMRCQ UCPC PCQNMLQG@JC DMP RP?LQDCPPGLE RFC @?ARCPG? RF?R A?SQCB KGRMAFMLBPG?J BCTCJMNKCLR RM RFC LCU CSI?PWMRCQ, UFCPC?Q MRFCP QNCAGCQ RP?LQDCPPCB RFC @?ARCPG? RF?R E?TC PGQC RM AFJMPMNJ?QRQ. 5FGQ KMBCJ GQ MDRCL A?JJCB RFC t web of life.u *L ?L CDDMPR RM Q?TC RFC RPCC ?L?JMEW, QMKC F?TC NPMNMQCB SQGLE RFC F0*<: RPCC (Figure 20.16b) UGRF GRQ KSJRGNJC RPSLIQ ?Q ? NFWJMECLCRGA RM PCNPCQCLR ? BGKGLGQFCB CTMJSRGML?PW PMJC DMP )(5.
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0RFCPQ F?TC NPMNMQCB ?@?LBMLGLE ?LW RPCC-JGIC KMBCJ MD NFWJMECLW GL D?TMP MD ? PGLE QRPSARSPC, RFC QMA?JJCB t ring of lifeu (Figure 20.17); ? NFWJMECLCRGA KMBCJ UFCPC ?JJ RFPCC BMK?GLQ MD JGDC CTMJTCB DPMK ? NMMJ MD NPGKGRGTC NPMI?PWMRCQ. -?IC, ?E?GL SQGLE RFC AMLBGRGMLCB PCAMLQRPSARGML ?JEMPGRFK, NPMNMQCQ ? PGLE-JGIC KMBCJ GL UFGAF QNCAGCQ MD ?JJ RFPCC BMK?GLQq"PAF?C?, #?ARCPG?, ?LB &SI?PW?qCTMJTCB DPMK ? QGLEJC NMMJ MD ECLC-QU?NNGLE NPMI?PWMRCQ. )GQ J?@MP?RMPW NPMNMQCQ RF?R RFGQ QRPSARSPC GQ RFC @CQR DGR DMP B?R? DPMK CVRCLQGTC %/" ?L?JWQCQ NCPDMPKCB GL FGQ J?@MP?RMPW, ?LB RF?R RFC PGLE KMBCJ GQ RFC MLJW MLC RF?R ?BCOS?RCJW R?ICQ )(5 ?LB ECLMKGA DSQGML GLRM ?AAMSLR. )MUCTCP, MRFCP NFWJMECLCRGAGQRQ PCK?GL FGEFJW QICNRGA?J MD RFGQ KMBCJ.
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542
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20.2 D<K<ID@E@E> MFCLK@FE8IP -<C8K@FEJ?@GJ

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1. Figure 20.6 "R UF?R JCTCJQ ?PC A?RQ ?LB BMEQ AMLQGBCPCB RM @C N?PR MD RFC Q?KC EPMSN 2. Figure 20.10 8FGAF ?LGK?JQ GL RFGQ DGESPC @CJMLE RM ? AJ?BC RF?R GLAJSBCQ ?LGK?JQ UGRF F?GP 8FGAF CTMJTCB DGPQR, F?GP MP RFC ?KLGMRGA CEE 3. Figure 20.11 8F?R GQ RFC J?PECQR AJ?BC GL RFGQ BG?EP?K
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6. 8FGAF QR?RCKCLR ?@MSR RFC R?VMLMKGA AJ?QQGDGA?RGML QWQRCK GQ AMPPCAR >. 5FCPC ?PC KMPC BMK?GLQ RF?L IGLEBMKQ. ?. ,GLEBMKQ ?PC RFC RMN A?RCEMPW MD AJ?QQGDGA?RGML. @. $J?QQCQ ?PC BGTGQGMLQ MD MPBCPQ. A. 4S@QNCAGCQ ?PC RFC KMQR QNCAGDGA A?RCEMPW MD AJ?QQGDGA?RGML. 7. 0L ? NFWJMECLCRGA RPCC, UFGAF RCPK PCDCPQ RM JGLC?ECQ RF?R BGTCPECB DPMK RFC Q?KC NJ?AC >. ?. @. A. QGQRCP R?V? @?Q?J R?V? PMMRCB R?V? BGAFMRMKMSQ R?V?
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21.1 R 1@I8C MFCLK@FE, (FIG?FCF>P, 8E; CC8JJ@=@:8K@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C FMU TGPSQCQ UCPC DGPQR BGQAMTCPCB ?LB FMU RFCW ?PC BCRCARCB z %GQASQQ RFPCC FWNMRFCQCQ ?@MSR FMU TGPSQCQ CTMJTCB z 3CAMELGXC RFC @?QGA QF?NCQ MD TGPSQCQ z 6LBCPQR?LB N?QR ?LB CKCPEGLE AJ?QQGDGA?RGML QWQRCKQ DMP TGPSQCQ 7GPSQCQ ?PC BGTCPQC CLRGRGCQ. 5FCW T?PW GL RFCGP QRPSARSPC, RFCGP PCNJGA?RGML KCRFMBQ, ?LB GL RFCGP R?PECR FMQRQ. /C?PJW ?JJ DMPKQ MD JGDCqDPMK @?ARCPG? ?LB ?PAF?C? RM CSI?PWMRCQ QSAF ?Q NJ?LRQ, ?LGK?JQ, ?LB DSLEGqF?TC TGPSQCQ RF?R GLDCAR RFCK. 8FGJC KMQR @GMJMEGA?J BGTCPQGRW A?L @C SLBCPQRMMB RFPMSEF CTMJSRGML?PW FGQRMPW, QSAF ?Q FMU QNCAGCQ F?TC ?B?NRCB RM AMLBGRGMLQ ?LB CLTGPMLKCLRQ, KSAF ?@MSR TGPSQ MPGEGLQ ?LB CTMJSRGML PCK?GLQ SLILMUL.
546
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1@I8C (FIG?FCF>P
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8FGJC UC BM F?TC JGKGRCB LSK@CPQ MD CDDCARGTC ?LRGTGP?J BPSEQ, QSAF ?Q RFMQC SQCB RM RPC?R )*7 ?LB GLDJSCLX?, RFC NPGK?PW KCRFMB MD AMLRPMJJGLE TGP?J BGQC?QC GQ @W T?AAGL?RGML, UFGAF GQ GLRCLBCB RM NPCTCLR MSR@PC?IQ @W @SGJBGLE GKKSLGRW RM ? TGPSQ MP TGPSQ D?KGJW (Figure 21.14). Vaccines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back mutations. -GTC T?AAGLCQ ?PC SQS?JJW K?BC @W attenuating (UC?ICLGLE) RFC tUGJB-RWNCu (BGQC?QC-A?SQGLE) TGPSQ @W EPMUGLE GR GL RFC J?@MP?RMPW GL RGQQSCQ MP ?R RCKNCP?RSPCQ BGDDCPCLR DPMK UF?R RFC TGPSQ GQ ?AASQRMKCB RM GL RFC FMQR. "B?NR?RGMLQ RM RFCQC LCU ACJJQ MP RCKNCP?RSPCQ GLBSAC KSR?RGMLQ GL RFC ECLMKCQ MD RFC TGPSQ, ?JJMUGLE GR RM EPMU @CRRCP GL RFC J?@MP?RMPW UFGJC GLFG@GRGLE GRQ ?@GJGRW RM A?SQC BGQC?QC UFCL PCGLRPMBSACB GLRM AMLBGRGMLQ DMSLB GL RFC FMQR. 5FCQC ?RRCLS?RCB TGPSQCQ RFSQ QRGJJ A?SQC GLDCARGML, @SR RFCW BM LMR EPMU TCPW UCJJ, ?JJMUGLE RFC GKKSLC PCQNMLQC RM BCTCJMN GL RGKC RM NPCTCLR K?HMP BGQC?QC. #?AI KSR?RGMLQ MAASP UFCL RFC T?AAGLC SLBCPEMCQ KSR?RGMLQ GL RFC FMQR QSAF RF?R GR PC?B?NRQ RM RFC FMQR ?LB A?L ?E?GL A?SQC BGQC?QC, UFGAF A?L RFCL @C QNPC?B RM MRFCP FSK?LQ GL ?L CNGBCKGA. 5FGQ RWNC MD QACL?PGM F?NNCLCB ?Q PCACLRJW ?Q 2007 GL /GECPG? UFCPC KSR?RGMLQ GL ? NMJGM T?AAGLC JCB RM ?L CNGBCKGA MD NMJGM GL RF?R AMSLRPW. 4MKC T?AAGLCQ ?PC GL AMLRGLSMSQ BCTCJMNKCLR @CA?SQC ACPR?GL TGPSQCQ, QSAF ?Q GLDJSCLX? ?LB )*7, F?TC ? FGEF KSR?RGML P?RC AMKN?PCB RM MRFCP TGPSQCQ ?LB LMPK?J FMQR ACJJQ. 8GRF GLDJSCLX?, KSR?RGMLQ GL RFC QSPD?AC KMJCASJCQ MD RFC TGPSQ FCJN RFC MPE?LGQK CT?BC RFC NPMRCARGTC GKKSLGRW RF?R K?W F?TC @CCL M@R?GLCB GL ? NPCTGMSQ GLDJSCLX? QC?QML, K?IGLE GR LCACQQ?PW DMP GLBGTGBS?JQ RM ECR T?AAGL?RCB CTCPW WC?P. 0RFCP TGPSQCQ, QSAF ?Q RFMQC RF?R A?SQC RFC AFGJBFMMB BGQC?QCQ KC?QJCQ, KSKNQ, ?LB PS@CJJ?, KSR?RC QM GLDPCOSCLRJW RF?R RFC Q?KC T?AAGLC GQ SQCB WC?P ?DRCP WC?P.
#(!04%2 21 X 6)253%3
561
!@>LI< 21.14 6>@@FK>QFLKP >OB ABPFDKBA QL ?LLPQ FJJRKFQV QL > SFORP QL MOBSBKQ FKCB@QFLK. (@OBAFQ: 53!#% %ROLMB $FPQOF@Q)
8?RAF RFGQ /07" video (http://openstaxcollege.org/l/1918_flu) RM JC?PL FMU KGAPM@GMJMEGQRQ ?PC ?RRCKNRGLE RM PCNJGA?RC RFC BC?BJW 1918 4N?LGQF GLDJSCLX? TGPSQ QM RFCW A?L SLBCPQR?LB KMPC ?@MSR TGPMJMEW.
18::@E<J 8E; AEK@-M@I8C DIL>J =FI /I<8KD<EK

*L QMKC A?QCQ, T?AAGLCQ A?L @C SQCB RM RPC?R ?L ?ARGTC TGP?J GLDCARGML. 5FC AMLACNR @CFGLB RFGQ GQ RF?R @W EGTGLE RFC T?AAGLC, GKKSLGRW GQ @MMQRCB UGRFMSR ?BBGLE KMPC BGQC?QC-A?SQGLE TGPSQ. *L RFC A?QC MD P?@GCQ, ? D?R?J LCSPMJMEGA?J BGQC?QC RP?LQKGRRCB TG? RFC Q?JGT? MD P?@GCQ TGPSQ-GLDCARCB ?LGK?JQ, RFC NPMEPCQQGML MD RFC BGQC?QC DPMK RFC RGKC MD RFC ?LGK?J @GRC RM RFC RGKC GR CLRCPQ RFC ACLRP?J LCPTMSQ QWQRCK K?W @C 2 UCCIQ MP JMLECP. 5FGQ GQ CLMSEF RGKC RM T?AAGL?RC ?L GLBGTGBS?J UFM QSQNCARQ RF?R RFCW F?TC @CCL @GRRCL @W ? P?@GB ?LGK?J, ?LB RFCGP @MMQRCB GKKSLC PCQNMLQC GQ QSDDGAGCLR RM NPCTCLR RFC TGPSQ DPMK CLRCPGLE LCPTMSQ RGQQSC. 5FSQ, RFC NMRCLRG?JJW D?R?J LCSPMJMEGA?J AMLQCOSCLACQ MD RFC BGQC?QC ?PC ?TCPRCB, ?LB RFC GLBGTGBS?J MLJW F?Q RM PCAMTCP DPMK RFC GLDCARCB @GRC. 5FGQ ?NNPM?AF GQ ?JQM @CGLE SQCB DMP RFC RPC?RKCLR MD &@MJ?, MLC MD RFC D?QRCQR ?LB KMQR BC?BJW TGPSQCQ ML C?PRF. 5P?LQKGRRCB @W @?RQ ?LB EPC?R ?NCQ, RFGQ BGQC?QC A?L A?SQC BC?RF GL 70p90 NCPACLR MD GLDCARCB FSK?LQ UGRFGL 2 UCCIQ. 6QGLE LCUJW BCTCJMNCB T?AAGLCQ RF?R @MMQR RFC GKKSLC PCQNMLQC GL RFGQ U?W, RFCPC GQ FMNC RF?R ?DDCARCB GLBGTGBS?JQ UGJJ @C @CRRCP ?@JC RM AMLRPMJ RFC TGPSQ, NMRCLRG?JJW Q?TGLE ? EPC?RCP NCPACLR?EC MD GLDCARCB NCPQMLQ DPMK ? P?NGB ?LB TCPW N?GLDSJ BC?RF. "LMRFCP U?W MD RPC?RGLE TGP?J GLDCARGMLQ GQ RFC SQC MD ?LRGTGP?J BPSEQ. 5FCQC BPSEQ MDRCL F?TC JGKGRCB QSAACQQ GL ASPGLE TGP?J BGQC?QC, @SR GL K?LW A?QCQ, RFCW F?TC @CCL SQCB RM AMLRPMJ ?LB PCBSAC QWKNRMKQ DMP ? UGBC T?PGCRW MD TGP?J BGQC?QCQ. 'MP KMQR TGPSQCQ, RFCQC BPSEQ A?L GLFG@GR RFC TGPSQ @W @JMAIGLE RFC ?ARGMLQ MD MLC MP KMPC MD GRQ NPMRCGLQ. *R GQ GKNMPR?LR RF?R RFC R?PECRCB NPMRCGLQ @C CLAMBCB @W TGP?J ECLCQ ?LB RF?R RFCQC KMJCASJCQ ?PC LMR NPCQCLR GL ? FC?JRFW FMQR ACJJ. *L RFGQ U?W, TGP?J EPMURF GQ GLFG@GRCB UGRFMSR B?K?EGLE RFC FMQR. 5FCPC ?PC J?PEC LSK@CPQ MD ?LRGTGP?J BPSEQ ?T?GJ?@JC RM RPC?R GLDCARGMLQ, QMKC QNCAGDGA DMP ? N?PRGASJ?P TGPSQ ?LB MRFCPQ RF?R A?L ?DDCAR KSJRGNJC TGPSQCQ. "LRGTGP?JQ F?TC @CCL BCTCJMNCB RM RPC?R ECLGR?J FCPNCQ (FCPNCQ QGKNJCV **) ?LB GLDJSCLX?. 'MP ECLGR?J FCPNCQ, BPSEQ QSAF ?Q ?AWAJMTGP A?L PCBSAC RFC LSK@CP ?LB BSP?RGML MD CNGQMBCQ MD ?ARGTC TGP?J BGQC?QC, BSPGLE UFGAF N?RGCLRQ BCTCJMN TGP?J JCQGMLQ GL RFCGP QIGL ACJJQ. "Q RFC TGPSQ PCK?GLQ J?RCLR GL LCPTMSQ RGQQSC MD RFC @MBW DMP JGDC, RFGQ BPSE GQ LMR ASP?RGTC @SR A?L K?IC RFC QWKNRMKQ MD RFC BGQC?QC KMPC K?L?EC?@JC. 'MP GLDJSCLX?, BPSEQ JGIC 5?KGDJS (MQCJR?KGTGP) (Figure 21.15) A?L PCBSAC RFC BSP?RGML MD
562
#(!04%2 21 X 6)253%3
tDJSu QWKNRMKQ @W 1 MP 2 B?WQ, @SR RFC BPSE BMCQ LMR NPCTCLR QWKNRMKQ CLRGPCJW. 5?KGDJS UMPIQ @W GLFG@GRGLE ?L CLXWKC (TGP?J LCSP?KGLGB?QC) RF?R ?JJMUQ LCU TGPGMLQ RM JC?TC RFCGP GLDCARCB ACJJQ. 5FSQ, 5?KGDJS GLFG@GRQ RFC QNPC?B MD TGPSQ DPMK GLDCARCB RM SLGLDCARCB ACJJQ. 0RFCP ?LRGTGP?J BPSEQ, QSAF ?Q 3G@?TGPGL, F?TC @CCL SQCB RM RPC?R ? T?PGCRW MD TGP?J GLDCARGMLQ, ?JRFMSEF GRQ KCAF?LGQK MD ?ARGML ?E?GLQR ACPR?GL TGPSQCQ PCK?GLQ SLAJC?P.
!@>LI< 21.15 (>) 4>JFCIR FKEF?FQP > SFO>I BKWVJB @>IIBA KBRO>JFKFA>PB (.!) CLRKA FK QEB FKCIRBKW> SFO>I BKSBILMB. (?) .BRO>JFKFA>PB @IB>SBP QEB @LKKB@QFLK ?BQTBBK SFO>I EBJ>DDIRQFKFK ((!), >IPL CLRKA FK QEB SFO>I BKSBILMB, >KA DIV@LMOLQBFKP LK QEB ELPQ @BII PROC>@B. )KEF?FQFLK LC KBRO>JFKFA>PB MOBSBKQP QEB SFORP COLJ ABQ>@EFKD COLJ QEB ELPQ @BII, QEBOB?V ?IL@HFKD CROQEBO FKCB@QFLK. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V -. %F@HJ>KK)
#W D?P, RFC KMQR QSAACQQDSJ SQC MD ?LRGTGP?JQ F?Q @CCL GL RFC RPC?RKCLR MD RFC PCRPMTGPSQ )*7, UFGAF A?SQCQ ? BGQC?QC RF?R, GD SLRPC?RCB, GQ SQS?JJW D?R?J UGRFGL 10p12 WC?PQ ?DRCP GLDCARGML. "LRG-)*7 BPSEQ F?TC @CCL ?@JC RM AMLRPMJ TGP?J PCNJGA?RGML RM RFC NMGLR RF?R GLBGTGBS?JQ PCACGTGLE RFCQC BPSEQ QSPTGTC DMP ? QGELGDGA?LRJW JMLECP RGKC RF?L RFC SLRPC?RCB. "LRG-)*7 BPSEQ GLFG@GR TGP?J PCNJGA?RGML ?R K?LW BGDDCPCLR NF?QCQ MD RFC )*7 PCNJGA?RGTC AWAJC (Figure 21.16). %PSEQ F?TC @CCL BCTCJMNCB RF?R GLFG@GR RFC DSQGML MD RFC )*7 TGP?J CLTCJMNC UGRF RFC NJ?QK? KCK@P?LC MD RFC FMQR ACJJ (DSQGML GLFG@GRMPQ), RFC AMLTCPQGML MD GRQ 3/" ECLMKC GLRM BMS@JCQRP?LBCB %/" (PCTCPQC RP?LQAPGNR?QC GLFG@GRMPQ), RFC GLRCEP?RGML MD RFC TGP?J %/" GLRM RFC FMQR ECLMKC (GLRCEP?QC GLFG@GRMPQ), ?LB RFC NPMACQQGLE MD TGP?J NPMRCGLQ (NPMRC?QC GLFG@GRMPQ).
#(!04%2 21 X 6)253%3
563
!@>LI< 21.16 ()6, >K BKSBILMBA, F@LP>EBAO>I SFORP, >QQ>@EBP QL QEB #$4 OB@BMQLO LC >K FJJRKB @BII >KA CRPBP TFQE QEB @BII JBJ?O>KB. 6FO>I @LKQBKQP >OB OBIB>PBA FKQL QEB @BII, TEBOB SFO>I BKWVJBP @LKSBOQ QEB PFKDIB-PQO>KABA 2.! DBKLJB FKQL $.! >KA FK@LOMLO>QB FQ FKQL QEB ELPQ DBKLJB. (@OBAFQ: .)!)$, .)()
8FCL ?LW MD RFCQC BPSEQ ?PC SQCB GLBGTGBS?JJW, RFC FGEF KSR?RGML P?RC MD RFC TGPSQ ?JJMUQ GR RM C?QGJW ?LB P?NGBJW BCTCJMN PCQGQR?LAC RM RFC BPSE, JGKGRGLE RFC BPSEsQ CDDCARGTCLCQQ. 5FC @PC?IRFPMSEF GL RFC RPC?RKCLR MD )*7 U?Q RFC BCTCJMNKCLR MD )""35, FGEFJW ?ARGTC ?LRG-PCRPMTGP?J RFCP?NW, UFGAF GLTMJTCQ ? KGVRSPC MD BGDDCPCLR BPSEQ, QMKCRGKCQ A?JJCB ? BPSE tAMAIR?GJ.u #W ?RR?AIGLE RFC TGPSQ ?R BGDDCPCLR QR?ECQ MD GRQ PCNJGA?RGTC AWAJC, GR GQ KSAF KMPC BGDDGASJR DMP RFC TGPSQ RM BCTCJMN PCQGQR?LAC RM KSJRGNJC BPSEQ ?R RFC Q?KC RGKC. 4RGJJ, CTCL UGRF RFC SQC MD AMK@GL?RGML )""35 RFCP?NW, RFCPC GQ AMLACPL RF?R, MTCP RGKC, RFC TGPSQ UGJJ BCTCJMN PCQGQR?LAC RM RFGQ RFCP?NW. 5FSQ, LCU ?LRG-)*7 BPSEQ ?PC AMLQR?LRJW @CGLE BCTCJMNCB UGRF RFC FMNC MD AMLRGLSGLE RFC @?RRJC ?E?GLQR RFGQ FGEFJW D?R?J TGPSQ.
564
#(!04%2 21 X 6)253%3
AGGC@<; 1@IFCF>P
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xJBOP>y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
21.4 R *K?<I A:<CCLC8I EK@K@<J: +I@FEJ 8E; 1@IF@;J

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C NPGMLQ ?LB RFCGP @?QGA NPMNCPRGCQ z %CDGLC TGPMGBQ ?LB RFCGP R?PECRQ MD GLDCARGML 1PGMLQ ?LB TGPMGBQ ?PC pathogens (?ECLRQ UGRF RFC ?@GJGRW RM A?SQC BGQC?QC) RF?R F?TC QGKNJCP QRPSARSPCQ RF?L TGPSQCQ @SR, GL RFC A?QC MD NPGMLQ, QRGJJ A?L NPMBSAC BC?BJW BGQC?QCQ.
+I@FEJ
Prions, QM-A?JJCB @CA?SQC RFCW ?PC NPMRCGL?ACMSQ, ?PC GLDCARGMSQ N?PRGAJCQqQK?JJCP RF?L TGPSQCQqRF?R AMLR?GL LM LSAJCGA ?AGBQ (LCGRFCP %/" LMP 3/"). )GQRMPGA?JJW, RFC GBC? MD ?L GLDCARGMSQ ?ECLR RF?R BGB
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#3):19:7'(1+ P740'7>49+8 '3* 9.+ $/'(1+-(:9-N43-C:19:7'(1+ !9'9+ .GAPM@GMJMEGQRQ RWNGA?JJW EPMU NPMI?PWMRCQ GL RFC J?@MP?RMPW SQGLE ?L ?NNPMNPG?RC ASJRSPC KCBGSK AMLR?GLGLE ?JJ RFC LSRPGCLRQ LCCBCB @W RFC R?PECR MPE?LGQK. 5FC KCBGSK A?L @C JGOSGB, @PMRF, MP QMJGB. "DRCP ?L GLAS@?RGML RGKC ?R RFC PGEFR RCKNCP?RSPC, RFCPC QFMSJB @C CTGBCLAC MD KGAPM@G?J EPMURF (Figure
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viable-but-non-culturable (7#/$) QR?RC. 5FC 7#/$ QR?RC MAASPQ UFCL NPMI?PWMRCQ PCQNMLB RM CLTGPMLKCLR?J QRPCQQMPQ @W CLRCPGLE ? BMPK?LR QR?RC RF?R ?JJMUQ RFCGP QSPTGT?J. 5FC APGRCPG? DMP CLRCPGLE GLRM RFC 7#/$ QR?RC ?PC LMR AMKNJCRCJW SLBCPQRMMB. *L ? NPMACQQ A?JJCB resuscitation, RFC NPMI?PWMRC A?L EM @?AI RM tLMPK?Ju JGDC UFCL CLTGPMLKCLR?J AMLBGRGMLQ GKNPMTC. *Q RFC 7#/$ QR?RC ?L SLSQS?J U?W MD JGTGLE DMP NPMI?PWMRCQ *L D?AR, KMQR MD RFC NPMI?PWMRCQ JGTGLE GL RFC QMGJ MP GL MAC?LGA U?RCPQ ?PC LML-ASJRSP?@JC. *R F?Q @CCL Q?GB RF?R MLJW ? QK?JJ DP?ARGML, NCPF?NQ MLC NCPACLR, MD NPMI?PWMRCQ A?L @C ASJRSPCB SLBCP J?@MP?RMPW AMLBGRGMLQ. *D RFCQC MPE?LGQKQ ?PC LMLASJRSP?@JC, RFCL FMU GQ GR ILMUL UFCRFCP RFCW ?PC NPCQCLR ?LB ?JGTC .GAPM@GMJMEGQRQ SQC KMJCASJ?P RCAFLGOSCQ, QSAF ?Q RFC NMJWKCP?QC AF?GL PC?ARGML (1$3), RM ?KNJGDW QCJCARCB NMPRGMLQ MD %/" MD NPMI?PWMRCQ, BCKMLQRP?RGLE RFCGP CVGQRCLAC. 3CA?JJ RF?R 1$3 A?L K?IC @GJJGMLQ MD AMNGCQ MD ? %/" QCEKCLR GL ? NPMACQQ A?JJCB ?KNJGDGA?RGML.
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6LRGJ ? AMSNJC MD BCA?BCQ ?EM, KGAPM@GMJMEGQRQ SQCB RM RFGLI MD NPMI?PWMRCQ ?Q GQMJ?RCB CLRGRGCQ JGTGLE ?N?PR. 5FGQ KMBCJ, FMUCTCP, BMCQ LMR PCDJCAR RFC RPSC CAMJMEW MD NPMI?PWMRCQ, KMQR MD UFGAF NPCDCP RM JGTC GL AMKKSLGRGCQ UFCPC RFCW A?L GLRCP?AR. " biofilm GQ ? KGAPM@G?J AMKKSLGRW (Figure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igure 22.14).
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585
!@>LI< 22.15 !O@E>B>I MELPMELIFMFAP AFCCBO COLJ QELPB CLRKA FK ">@QBOF> >KA %RH>OV> FK QTL T>VP. &FOPQ, QEBV E>SB ?O>K@EBA MEVQ>KVI PFAB@E>FKP FKPQB>A LC IFKB>O LKBP. 3B@LKA, >K BQEBO ?LKA FKPQB>A LC >K BPQBO ?LKA @LKKB@QP QEB IFMFA QL QEB DIV@BOLI.
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peptidoglycan, AMKNMQCB MD NMJWQ?AAF?PGBC AF?GLQ RF?R ?PC APMQQJGLICB @W SLSQS?J NCNRGBCQ AMLR?GLGLE @MRF -- ?LB %-?KGLM ?AGBQ GLAJSBGLE %-EJSR?KGA ?AGB ?LB %?J?LGLC. 1PMRCGLQ LMPK?JJW F?TC MLJW --?KGLM ?AGBQ; ?Q ? AMLQCOSCLAC, K?LW MD MSP ?LRG@GMRGAQ UMPI @W KGKGAIGLE %-?KGLM ?AGBQ ?LB RFCPCDMPC F?TC QNCAGDGA CDDCARQ ML @?ARCPG?J ACJJ U?JJ BCTCJMNKCLR. 5FCPC ?PC KMPC RF?L 100 BGDDCPCLR DMPKQ MD NCNRGBMEJWA?L. S-layer (QSPD?AC J?WCP) NPMRCGLQ ?PC ?JQM NPCQCLR ML RFC MSRQGBC MD ACJJ U?JJQ MD @MRF ?PAF?C? ?LB @?ARCPG?. #?ARCPG? ?PC BGTGBCB GLRM RUM K?HMP EPMSNQ: Gram positive ?LB Gram negative, @?QCB ML RFCGP PC?ARGML RM (P?K QR?GLGLE. /MRC RF?R ?JJ (P?K-NMQGRGTC @?ARCPG? @CJMLE RM MLC NFWJSK; @?ARCPG? GL RFC MRFCP DMSP NFWJ? (1PMRCM@?ARCPG?, $FJ?KWBG?Q, 4NGPMAFCRCQ, ?LB $W?LM@?ARCPG?) ?PC (P?K-LCE?RGTC. 5FC (P?K QR?GLGLE KCRFMB GQ L?KCB ?DRCP GRQ GLTCLRMP, %?LGQF QAGCLRGQR )?LQ $FPGQRG?L (P?K (1853p1938). 5FC BGDDCPCLR @?ARCPG?J PCQNMLQCQ RM RFC QR?GLGLE NPMACBSPC ?PC SJRGK?RCJW BSC RM ACJJ U?JJ QRPSARSPC. (P?KNMQGRGTC MPE?LGQKQ RWNGA?JJW J?AI RFC MSRCP KCK@P?LC DMSLB GL (P?K-LCE?RGTC MPE?LGQKQ (Figure 22.15). 6N RM 90 NCPACLR MD RFC ACJJ U?JJ GL (P?K-NMQGRGTC @?ARCPG? GQ AMKNMQCB MD NCNRGBMEJWA?L, ?LB KMQR MD RFC PCQR GQ AMKNMQCB MD ?AGBGA QS@QR?LACQ A?JJCB teichoic acids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
586
#(!04%2 22 X 02/+!29/4%3: "!#4%2)! !.$ !2#(!%!
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587
*L transformation, RFC NPMI?PWMRC R?ICQ GL %/" DMSLB GL GRQ CLTGPMLKCLR RF?R GQ QFCB @W MRFCP NPMI?PWMRCQ. *D ? LMLN?RFMECLGA @?ARCPGSK R?ICQ SN %/" DMP ? RMVGL ECLC DPMK ? N?RFMECL ?LB GLAMPNMP?RCQ RFC LCU %/" GLRM GRQ MUL AFPMKMQMKC, GR RMM K?W @CAMKC N?RFMECLGA. *L transduction, @?ARCPGMNF?ECQ, RFC TGPSQCQ RF?R GLDCAR @?ARCPG?, QMKCRGKCQ ?JQM KMTC QFMPR NGCACQ MD AFPMKMQMK?J %/" DPMK MLC @?ARCPGSK RM ?LMRFCP. 5P?LQBSARGML PCQSJRQ GL ? PCAMK@GL?LR MPE?LGQK. "PAF?C? ?PC LMR ?DDCARCB @W @?ARCPGMNF?ECQ @SR GLQRC?B F?TC RFCGP MUL TGPSQCQ RF?R RP?LQJMA?RC ECLCRGA K?RCPG?J DPMK MLC GLBGTGBS?J RM ?LMRFCP. *L conjugation, %/" GQ RP?LQDCPPCB DPMK MLC NPMI?PWMRC RM ?LMRFCP @W KC?LQ MD ? NGJSQ, UFGAF @PGLEQ RFC MPE?LGQKQ GLRM AMLR?AR UGRF MLC ?LMRFCP. 5FC %/" RP?LQDCPPCB A?L @C GL RFC DMPK MD ? NJ?QKGB MP ?Q ? FW@PGB, AMLR?GLGLE @MRF NJ?QKGB ?LB AFPMKMQMK?J %/". 5FCQC RFPCC NPMACQQCQ MD %/" CVAF?LEC ?PC QFMUL GL Figure 22.16. 3CNPMBSARGML A?L @C TCPW P?NGB: ? DCU KGLSRCQ DMP QMKC QNCAGCQ. 5FGQ QFMPR ECLCP?RGML RGKC AMSNJCB UGRF KCAF?LGQKQ MD ECLCRGA PCAMK@GL?RGML ?LB FGEF P?RCQ MD KSR?RGML PCQSJR GL RFC P?NGB CTMJSRGML MD NPMI?PWMRCQ, ?JJMUGLE RFCK RM PCQNMLB RM CLTGPMLKCLR?J AF?LECQ (QSAF ?Q RFC GLRPMBSARGML MD ?L ?LRG@GMRGA) TCPW OSGAIJW.
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588
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5FC BGQRPG@SRGML MD ? N?PRGASJ?P BGQC?QC GQ BWL?KGA. 5FCPCDMPC, AF?LECQ GL RFC CLTGPMLKCLR, RFC N?RFMECL, MP RFC FMQR NMNSJ?RGML A?L BP?K?RGA?JJW GKN?AR RFC QNPC?B MD ? BGQC?QC. "AAMPBGLE RM RFC 8MPJB )C?JRF 0PE?LGX?RGML (8)0) ?L emerging disease (Figure 22.22) GQ MLC RF?R F?Q ?NNC?PCB GL ? NMNSJ?RGML DMP RFC DGPQR RGKC, MP RF?R K?W F?TC CVGQRCB NPCTGMSQJW @SR GQ P?NGBJW GLAPC?QGLE GL GLAGBCLAC MP ECMEP?NFGA P?LEC. 5FGQ BCDGLGRGML ?JQM GLAJSBCQ PC-CKCPEGLE BGQC?QCQ RF?R UCPC NPCTGMSQJW SLBCP AMLRPMJ. "NNPMVGK?RCJW 75 NCPACLR MD PCACLRJW CKCPEGLE GLDCARGMSQ BGQC?QCQ ?DDCARGLE FSK?LQ ?PC XMMLMRGA BGQC?QCQ, zoonoses, BGQC?QCQ RF?R NPGK?PGJW GLDCAR ?LGK?JQ ?LB ?PC RP?LQKGRRCB RM FSK?LQ; QMKC ?PC MD TGP?J MPGEGL ?LB QMKC ?PC MD @?ARCPG?J MPGEGL. #PSACJJMQGQ GQ ?L CV?KNJC MD ? NPMI?PWMRGA XMMLMQGQ RF?R GQ PC-CKCPEGLE GL QMKC PCEGMLQ, ?LB LCAPMRGXGLE D?QAGGRGQ (AMKKMLJW ILMUL ?Q DJCQF-C?RGLE @?ARCPG?) F?Q @CCL GLAPC?QGLE GL TGPSJCLAC DMP RFC J?QR 80 WC?PQ DMP SLILMUL PC?QMLQ.
594
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3CA?JJ RF?R @GMDGJKQ ?PC KGAPM@G?J AMKKSLGRGCQ RF?R ?PC TCPW BGDDGASJR RM BCQRPMW. 5FCW ?PC PCQNMLQG@JC DMP BGQC?QCQ QSAF ?Q GLDCARGMLQ GL N?RGCLRQ UGRF AWQRGA DG@PMQGQ, -CEGMLL?GPCQs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sR ?BCOS?RCJW AJC?LCB. #GMDGJK GLDCARGMLQ BCTCJMN EP?BS?JJW; QMKCRGKCQ, RFCW BM LMR A?SQC QWKNRMKQ GKKCBG?RCJW. 5FCW ?PC P?PCJW PCQMJTCB @W FMQR BCDCLQC KCAF?LGQKQ. 0LAC ?L GLDCARGML @W ? @GMDGJK GQ CQR?@JGQFCB, GR GQ
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8?RAF ? PCACLR LCUQ report (http://openstaxcollege.org/l/antibiotics) ML RFC NPM@JCK MD PMSRGLC ?LRG@GMRGA ?BKGLGQRP?RGML RM JGTCQRMAI ?LB ?LRG@GMRGA-PCQGQR?LR @?ARCPG?. O3+ 4, 9.+ !:5+7(:-8: M !A 5FC GKNPSBCLR SQC MD ?LRG@GMRGAQ F?Q N?TCB RFC U?W DMP @?ARCPG? RM CVN?LB NMNSJ?RGMLQ MD PCQGQR?LR DMPKQ. 'MP CV?KNJC, ;(7/@36*6**<: (<9,<Q, MDRCL A?JJCB tQR?NF,u GQ ? AMKKML @?ARCPGSK RF?R A?L JGTC GL RFC FSK?L @MBW ?LB GQ SQS?JJW C?QGJW RPC?RCB UGRF ?LRG@GMRGAQ. " TCPW B?LECPMSQ QRP?GL, FMUCTCP, methicillinresistant Staphylococcus aureus (MRSA) F?Q K?BC RFC LCUQ MTCP RFC N?QR DCU WC?PQ (Figure 22.24). 5FGQ QRP?GL GQ PCQGQR?LR RM K?LW AMKKMLJW SQCB ?LRG@GMRGAQ, GLAJSBGLE KCRFGAGJJGL, ?KMVGAGJJGL, NCLGAGJJGL, ?LB MV?AGJJGL. .34" A?L A?SQC GLDCARGMLQ MD RFC QIGL, @SR GR A?L ?JQM GLDCAR RFC @JMMBQRPC?K, JSLEQ, SPGL?PW RP?AR, MP QGRCQ MD GLHSPW. 8FGJC .34" GLDCARGMLQ ?PC AMKKML ?KMLE NCMNJC GL FC?JRFA?PC D?AGJGRGCQ, RFCW F?TC ?JQM ?NNC?PCB GL FC?JRFW NCMNJC UFM F?TCLsR @CCL FMQNGR?JGXCB @SR UFM JGTC MP UMPI GL RGEFR NMNSJ?RGMLQ (JGIC KGJGR?PW NCPQMLLCJ ?LB NPGQMLCPQ). 3CQC?PAFCPQ F?TC CVNPCQQCB AMLACPL ?@MSR RFC U?W RFGQ J?RRCP QMSPAC MD .34" R?PECRQ ? KSAF WMSLECP NMNSJ?RGML RF?L RFMQC PCQGBGLE GL A?PC D?AGJGRGCQ. !/, J6<95(3 6- ;/, A4,90*(5 M,+0*(3 A::6*0(;065 PCNMPRCB RF?R, ?KMLE .34"-?DDJGARCB NCPQMLQ GL FC?JRFA?PC D?AGJGRGCQ, RFC ?TCP?EC ?EC GQ 68, UFCPC?Q NCMNJC UGRF tAMKKSLGRW-?QQMAG?RCB .34"u ( CA-MRSA) F?TC ?L ?TCP?EC ?EC MD 23.
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4. /?GKG, 54, -C%CJJ, ,), $MKM-4?@CRRG, ,, CR ?J. $MKN?PGQML MD AMKKSLGRW- ?LB FC?JRF A?PC-?QQMAG?RCB KCRFGAGJJGL-PCQGQR?LR ;(7/@36*6**<: (<9,<: GLDCARGML. JAMA 290 (2003): 2976p84, BMG: 10.1001/ H?K?.290.22.2976.
596
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22.3 +IFB8IPFK@: (<K89FC@JD

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616
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618
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#(!04%2 23 X 02/4)343
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thallus. %GKMPNFGA DSLEG A?L AF?LEC DPMK RFC SLGACJJSJ?P RM KSJRGACJJSJ?P QR?RC BCNCLBGLE ML CLTGPMLKCLR?J AMLBGRGMLQ. 6LGACJJSJ?P DSLEG ?PC ECLCP?JJW PCDCPPCB RM ?Q yeasts. (**/(964@*,: *,9,=0:0(, (@?ICPsQ WC?QR) ?LB C(5+0+( QNCAGCQ (RFC ?ECLRQ MD RFPSQF, ? AMKKML DSLE?J GLDCARGML) ?PC CV?KNJCQ MD SLGACJJSJ?P DSLEG (Figure 24.3).
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.MQR DSLEG ?PC KSJRGACJJSJ?P MPE?LGQKQ. 5FCW BGQNJ?W RUM BGQRGLAR KMPNFMJMEGA?J QR?ECQ: RFC TCECR?RGTC ?LB PCNPMBSARGTC. 5FC TCECR?RGTC QR?EC AMLQGQRQ MD ? R?LEJC MD QJCLBCP RFPC?B-JGIC QRPSARSPCQ A?JJCB hyphae (QGLESJ?P, hypha), UFCPC?Q RFC PCNPMBSARGTC QR?EC A?L @C KMPC AMLQNGASMSQ. 5FC K?QQ MD FWNF?C GQ ? mycelium (Figure 24.4). *R A?L EPMU ML ? QSPD?AC, GL QMGJ MP BCA?WGLE K?RCPG?J, GL ? JGOSGB, MP CTCL ML JGTGLE RGQQSC. "JRFMSEF GLBGTGBS?J FWNF?C KSQR @C M@QCPTCB SLBCP ? KGAPMQAMNC, RFC KWACJGSK MD ? DSLESQ A?L @C TCPW J?PEC, UGRF QMKC QNCAGCQ RPSJW @CGLE tRFC DSLESQ FSKMLEMSQ.u 5FC EG?LR A94033(90( :630+07,: (FMLCW KSQFPMMK) GQ AMLQGBCPCB RFC J?PECQR MPE?LGQK ML &?PRF, QNPC?BGLE ?APMQQ KMPC RF?L 2,000 ?APCQ MD SLBCPEPMSLB QMGJ GL C?QRCPL 0PCEML; GR GQ CQRGK?RCB RM @C ?R JC?QR 2,400 WC?PQ MJB.
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645
DMPKCB @W J?PEC ACJJQ AMLR?GLGLE K?LW LSAJCG, ?L ?PP?LECKCLR BCQAPG@CB ?Q coenocytic hyphae (Figure 24.5b).
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646
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igure 24.6).
!@>LI< 24.6 4EB (>) DF>KQ MRCC ?>II JRPEOLLJ OBIB>PBP (?) > @ILRA LC PMLOBP TEBK FQ OB>@EBP J>QROFQV. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V 2LDBO 'OFCCFQE; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V 0B>OPLK 3@LQQ &LOBPJ>K, ALK>QBA QL QEB 7FHFJBAF> &LRKA>QFLK)
A8+=:'1 +574*:)9/43 'SLEG PCNPMBSAC ?QCVS?JJW @W DP?EKCLR?RGML, @SBBGLE, MP NPMBSAGLE QNMPCQ. 'P?EKCLRQ MD FWNF?C A?L EPMU LCU AMJMLGCQ. 4MK?RGA ACJJQ GL WC?QR DMPK @SBQ. %SPGLE @SBBGLE (? RWNC MD AWRMIGLCQGQ), ? @SJEC DMPKQ ML RFC QGBC MD RFC ACJJ, RFC LSAJCSQ BGTGBCQ KGRMRGA?JJW, ?LB RFC @SB SJRGK?RCJW BCR?AFCQ GRQCJD DPMK RFC KMRFCP ACJJ (Figure 24.7).
!@>LI< 24.7 4EB A>OH @BIIP FK QEFP ?OFDEQ CFBIA IFDEQ JF@OLDO>ME >OB QEB M>QELDBKF@ VB>PQ H0:;673(:4( *(7:<3(;<4, PBBK >D>FKPQ > ?>@HAOLM LC IFDEQ ?IRB QFPPRB. (FPQLMI>PJ> MOFJ>OFIV FKCB@QP IRKDP ?RQ @>K PMOB>A QL LQEBO QFPPRBP, @>RPFKD EFPQLMI>PJLPFP, > MLQBKQF>IIV C>Q>I AFPB>PB. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V $O. ,F?BOL !GBIIL, #$#; P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
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!@>LI< 24.9 4EFP ?OFDEQ CFBIA IFDEQ JF@OLDO>ME PELTP QEB OBIB>PB LC PMLOBP COLJ > PMLO>KDFRJ >Q QEB BKA LC > EVME> @>IIBA > PMLO>KDFLMELOB. 4EB LOD>KFPJ FP > M<*69 PM. CRKDRP, > JLIA LCQBK CLRKA FKALLOP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V $O. ,R@FIIB 'BLOD, #$#; P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
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648
#(!04%2 24 X &5.')
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3CTGCU RFC AF?P?ARCPGQRGAQ MD DSLEG @W TGQGRGLE RFGQ interactive site (http://openstaxcollege.org/l/ fungi_kingdom) DPMK 8GQAMLQGL-MLJGLC.
24.2 R CC8JJ@=@:8K@FEJ F= !LE>@

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5FC MLJW AJ?QQ GL RFC 1FWJSK $FWRPGBGMKWAMR? GQ RFC Chytridiomycetes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igure 24.10), UFGJC MRFCPQ ?PC Q?NPM@CQ. 5FC AFWRPGB QNCAGCQ A3364@*,: GQ UCJJ AF?P?ARCPGXCB ?Q ?L CVNCPGKCLR?J MPE?LGQK. *RQ PCNPMBSARGTC AWAJC GLAJSBCQ @MRF ?QCVS?J ?LB QCVS?J NF?QCQ. A3364@*,: NPMBSACQ BGNJMGB MP F?NJMGB DJ?ECJJ?RCB XMMQNMPCQ GL ? QNMP?LEGSK.
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qGLAJSBGLE QC? QR?PQ ?LB QC? ?LCKMLCQ, ?Q UCJJ ?Q QMKC GLQCARQ, PCNRGJCQ, ?LB DGQFq?PC A?N?@JC MD ?QCVS?J PCNPMBSARGML. 5FC KMQR AMKKML DMPKQ MD ?QCVS?J PCNPMBSARGML DMP QR?RGML?PW ?OS?RGA ?LGK?JQ GLAJSBC @SBBGLE ?LB DP?EKCLR?RGML, UFCPC N?PR MD ? N?PCLR GLBGTGBS?J A?L QCN?P?RC ?LB EPMU GLRM ? LCU GLBGTGBS?J. *L AMLRP?QR, ? DMPK MD ?QCVS?J PCNPMBSARGML DMSLB GL ACPR?GL GLQCARQ ?LB TCPRC@P?RCQ GQ A?JJCB N?PRFCLMECLCQGQ (MP tTGPEGL @CEGLLGLEu), UFCPC SLDCPRGJGXCB CEEQ A?L BCTCJMN GLRM LCU K?JC MDDQNPGLE. 5FGQ RWNC MD N?PRFCLMECLCQGQ GQ A?JJCB F?NJMBGNJMGBW. 5FCQC RWNCQ MD ?QCVS?J PCNPMBSARGML NPMBSAC ECLCRGA?JJW GBCLRGA?J MDDQNPGLE, UFGAF GQ BGQ?BT?LR?ECMSQ DPMK RFC NCPQNCARGTC MD CTMJSRGML?PW ?B?NR?@GJGRW @CA?SQC MD RFC NMRCLRG?J @SGJBSN MD BCJCRCPGMSQ KSR?RGMLQ. )MUCTCP, DMP ?LGK?JQ RF?R ?PC JGKGRCB GL RFCGP A?N?AGRW RM ?RRP?AR K?RCQ, ?QCVS?J PCNPMBSARGML A?L CLQSPC ECLCRGA NPMN?E?RGML. "DRCP DCPRGJGX?RGML, ? QCPGCQ MD BCTCJMNKCLR?J QR?ECQ MAASP BSPGLE UFGAF NPGK?PW ECPK J?WCPQ ?PC CQR?@JGQFCB ?LB PCMPE?LGXC RM DMPK ?L CK@PWM. %SPGLE RFGQ NPMACQQ, ?LGK?J RGQQSCQ @CEGL RM QNCAG?JGXC ?LB MPE?LGXC GLRM MPE?LQ ?LB MPE?L QWQRCKQ, BCRCPKGLGLE RFCGP DSRSPC KMPNFMJMEW ?LB NFWQGMJMEW. 4MKC ?LGK?JQ, QSAF ?Q EP?QQFMNNCPQ, SLBCPEM GLAMKNJCRC KCR?KMPNFMQGQ, GL UFGAF RFC WMSLE PCQCK@JC RFC ?BSJR. 0RFCP ?LGK?JQ, QSAF ?Q QMKC GLQCARQ, SLBCPEM AMKNJCRC KCR?KMPNFMQGQ UFCPC GLBGTGBS?JQ CLRCP MLC MP KMPC J?PT?J QR?ECQ RF?R K?W GL BGDDCP GL QRPSARSPC ?LB DSLARGML DPMK RFC ?BSJR (Figure 27.3). 'MP RFC J?RRCP, RFC WMSLE ?LB RFC ?BSJR K?W F?TC BGDDCPCLR BGCRQ, JGKGRGLE AMKNCRGRGML DMP DMMB @CRUCCL RFCK. 3CE?PBJCQQ MD UFCRFCP ? QNCAGCQ SLBCPEMCQ AMKNJCRC MP GLAMKNJCRC KCR?KMPNFMQGQ, RFC QCPGCQ MD BCTCJMNKCLR?J QR?ECQ MD RFC CK@PWM PCK?GLQ J?PECJW RFC Q?KC DMP KMQR KCK@CPQ MD RFC ?LGK?J IGLEBMK.
730
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!@>LI< 27.3 (>) 4EB DO>PPELMMBO RKABODLBP FK@LJMIBQB JBQ>JLOMELPFP. (?) 4EB ?RQQBOCIV RKABODLBP @LJMIBQB JBQ>JLOMELPFP. (@OBAFQ: 3.%. 3KLADO>PP, 53$!)
5FC NPMACQQ MD ?LGK?J BCTCJMNKCLR @CEGLQ UGRF RFC cleavage, MP QCPGCQ MD KGRMRGA ACJJ BGTGQGMLQ, MD RFC XWEMRC (Figure 27.4). 5FPCC ACJJ BGTGQGMLQ RP?LQDMPK RFC QGLEJC-ACJJCB XWEMRC GLRM ?L CGEFR-ACJJCB QRPSARSPC. "DRCP DSPRFCP ACJJ BGTGQGML ?LB PC?PP?LECKCLR MD CVGQRGLE ACJJQ, ? 6p32-ACJJCB FMJJMU QRPSARSPC A?JJCB ? blastula GQ DMPKCB. /CVR, RFC @J?QRSJ? SLBCPEMCQ DSPRFCP ACJJ BGTGQGML ?LB ACJJSJ?P PC?PP?LECKCLR BSPGLE ? NPMACQQ A?JJCB E?QRPSJ?RGML. 5FGQ JC?BQ RM RFC DMPK?RGML MD RFC LCVR BCTCJMNKCLR?J QR?EC, RFC gastrula, GL UFGAF RFC DSRSPC BGECQRGTC A?TGRW GQ DMPKCB. %GDDCPCLR ACJJ J?WCPQ (A?JJCB germ layers) ?PC DMPKCB BSPGLE E?QRPSJ?RGML. 5FCQC ECPK J?WCPQ ?PC NPMEP?KKCB RM BCTCJMN GLRM ACPR?GL RGQQSC RWNCQ, MPE?LQ, ?LB MPE?L QWQRCKQ BSPGLE ? NPMACQQ A?JJCB organogenesis.
!@>LI< 27.4 $ROFKD BJ?OVLKF@ ABSBILMJBKQ, QEB WVDLQB RKABODLBP > PBOFBP LC JFQLQF@ @BII AFSFPFLKP, LO @IB>S>DBP, QL CLOJ >K BFDEQ-@BII PQ>DB, QEBK > ELIILT ?I>PQRI>. $ROFKD > MOL@BPP @>IIBA D>PQORI>QFLK, QEB ?I>PQRI> CLIAP FKT>OA QL CLOJ > @>SFQV FK QEB D>PQORI>.
8?RAF RFC DMJJMUGLE video (http://openstaxcollege.org/l/embryo_evol) RM QCC FMU FSK?L CK@PWMLGA BCTCJMNKCLR (?DRCP RFC @J?QRSJ? ?LB E?QRPSJ? QR?ECQ MD BCTCJMNKCLR) PCDJCARQ CTMJSRGML.
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731
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tFMKCMRGA ECLCQ,u ?LB RFCW AMLR?GL %/" QCOSCLACQ A?JJCB FMKCM@MVCQ. 5FC ?LGK?J ECLCQ AMLR?GLGLE FMKCM@MV QCOSCLACQ ?PC QNCAGDGA?JJW PCDCPPCB RM ?Q Hox genes. 5FGQ D?KGJW MD ECLCQ GQ PCQNMLQG@JC DMP BCRCPKGLGLE RFC ECLCP?J @MBW NJ?L, QSAF ?Q RFC LSK@CP MD @MBW QCEKCLRQ MD ?L ?LGK?J, RFC LSK@CP ?LB NJ?ACKCLR MD ?NNCLB?ECQ, ?LB ?LGK?J FC?B-R?GJ BGPCARGML?JGRW. 5FC DGPQR H6? ECLCQ RM @C QCOSCLACB UCPC RFMQC DPMK RFC DPSGR DJW (D96:67/03( 4,3(56.(:;,9). " QGLEJC H6? KSR?RGML GL RFC DPSGR DJW A?L PCQSJR GL ?L CVRP? N?GP MD UGLEQ MP CTCL ?NNCLB?ECQ EPMUGLE DPMK RFC tUPMLEu @MBW N?PR. 8FGJC RFCPC ?PC ? EPC?R K?LW ECLCQ RF?R NJ?W PMJCQ GL RFC KMPNFMJMEGA?J BCTCJMNKCLR MD ?L ?LGK?J, UF?R K?ICQ H6? ECLCQ QM NMUCPDSJ GQ RF?R RFCW QCPTC ?Q K?QRCP AMLRPMJ ECLCQ RF?R A?L RSPL ML MP MDD J?PEC LSK@CPQ MD MRFCP ECLCQ. H6? ECLCQ BM RFGQ @W AMBGLE RP?LQAPGNRGML D?ARMPQ RF?R AMLRPMJ RFC CVNPCQQGML MD LSKCPMSQ MRFCP ECLCQ. H6? ECLCQ ?PC FMKMJMEMSQ GL RFC ?LGK?J IGLEBMK, RF?R GQ, RFC ECLCRGA QCOSCLACQ MD H6? ECLCQ ?LB RFCGP NMQGRGMLQ ML AFPMKMQMKCQ ?PC PCK?PI?@JW QGKGJ?P ?APMQQ KMQR ?LGK?JQ @CA?SQC MD RFCGP NPCQCLAC GL ? AMKKML ?LACQRMP, DPMK UMPKQ RM DJGCQ, KGAC, ?LB FSK?LQ (Figure 27.5). 0LC MD RFC AMLRPG@SRGMLQ RM GLAPC?QCB ?LGK?J @MBW AMKNJCVGRW GQ RF?R H6? ECLCQ F?TC SLBCPEMLC ?R JC?QR RUM BSNJGA?RGML CTCLRQ BSPGLE ?LGK?J CTMJSRGML, UGRF RFC ?BBGRGML?J ECLCQ ?JJMUGLE DMP KMPC AMKNJCV @MBW RWNCQ RM CTMJTC.
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
)C > H6? 13 DBKB FK > JLRPB T>P OBMI>@BA TFQE > H6? 1 DBKB, ELT JFDEQ QEFP >IQBO >KFJ>I ABSBILMJBKQ
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27.2 R !<8KLI<J 0J<; KF CC8JJ@=P AE@D8CJ

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL RFC BGDDCPCLACQ GL ?LGK?J @MBW NJ?LQ RF?R QSNNMPR @?QGA ?LGK?J AJ?QQGDGA?RGML z $MKN?PC ?LB AMLRP?QR RFC CK@PWMLGA BCTCJMNKCLR MD NPMRMQRMKCQ ?LB BCSRCPMQRMKCQ 4AGCLRGQRQ F?TC BCTCJMNCB ? AJ?QQGDGA?RGML QAFCKC RF?R A?RCEMPGXCQ ?JJ KCK@CPQ MD RFC ?LGK?J IGLEBMK, ?JRFMSEF RFCPC ?PC CVACNRGMLQ RM KMQR tPSJCQu EMTCPLGLE ?LGK?J AJ?QQGDGA?RGML (Figure 27.6). "LGK?JQ ?PC NPGK?PGJW AJ?QQGDGCB ?AAMPBGLE RM KMPNFMJMEGA?J ?LB BCTCJMNKCLR?J AF?P?ARCPGQRGAQ, QSAF ?Q ? @MBW NJ?L. 0LC MD RFC KMQR NPMKGLCLR DC?RSPCQ MD RFC @MBW NJ?L MD RPSC ?LGK?JQ GQ RF?R RFCW ?PC KMPNFMJMEGA?JJW QWKKCRPGA?J. 5FGQ KC?LQ RF?R RFCGP BGQRPG@SRGML MD @MBW N?PRQ GQ @?J?LACB ?JMLE ?L ?VGQ. "BBGRGML?J AF?P?ARCPGQRGAQ GLAJSBC RFC LSK@CP MD RGQQSC J?WCPQ DMPKCB BSPGLE BCTCJMNKCLR, RFC NPCQCLAC MP ?@QCLAC MD ?L GLRCPL?J @MBW A?TGRW, ?LB MRFCP DC?RSPCQ MD CK@PWMJMEGA?J BCTCJMNKCLR, QSAF ?Q RFC MPGEGL MD RFC KMSRF ?LB ?LSQ.
!@>LI< 27.6 4EB MEVILDBKBQF@ QOBB LC >KFJ>IP FP ?>PBA LK JLOMELILDF@>I, CLPPFI, >KA DBKBQF@ BSFABK@B.
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7EF@E LC QEB CLIILTFKD PQ>QBJBKQP FP C>IPB %RJBQ>WL>KP E>SB PMB@F>IFWBA QFPPRBP >KA M>O>WL>KP ALKwQ. ,LMELQOL@ELWL> >KA %@AVPLWL> >OB ?LQE "FI>Q>OF>. !@LBI> >KA #KFA>OF> ?LQE MLPPBPP O>AF>I PVJJBQOV. !OQEOLMLAP >OB JLOB @ILPBIV OBI>QBA QL KBJ>QLABP QE>K QEBV >OB QL >KKBIFAP.
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733
AE@D8C C?8I8:K<I@Q8K@FE B8J<; FE BF;P .PDD<KIP

"R ? TCPW @?QGA JCTCJ MD AJ?QQGDGA?RGML, RPSC ?LGK?JQ A?L @C J?PECJW BGTGBCB GLRM RFPCC EPMSNQ @?QCB ML RFC RWNC MD QWKKCRPW MD RFCGP @MBW NJ?L: P?BG?JJW QWKKCRPGA?J, @GJ?RCP?JJW QWKKCRPGA?J, ?LB ?QWKKCRPGA?J. "QWKKCRPW GQ ? SLGOSC DC?RSPC MD 1?P?XM? (Figure 27.7a). 0LJW ? DCU ?LGK?J EPMSNQ BGQNJ?W P?BG?J QWKKCRPW. "JJ RWNCQ MD QWKKCRPW ?PC UCJJ QSGRCB RM KCCR RFC SLGOSC BCK?LBQ MD ? N?PRGASJ?P ?LGK?JsQ JGDCQRWJC. Radial symmetry GQ RFC ?PP?LECKCLR MD @MBW N?PRQ ?PMSLB ? ACLRP?J ?VGQ, ?Q GQ QCCL GL ? BPGLIGLE EJ?QQ MP NGC. *R PCQSJRQ GL ?LGK?JQ F?TGLE RMN ?LB @MRRMK QSPD?ACQ @SR LM JCDR ?LB PGEFR QGBCQ, MP DPMLR MP @?AI. 5FC RUM F?JTCQ MD ? P?BG?JJW QWKKCRPGA?J ?LGK?J K?W @C BCQAPG@CB ?Q RFC QGBC UGRF ? KMSRF MP tMP?J QGBC,u ?LB RFC QGBC UGRFMSR ? KMSRF (RFC t?@MP?J QGBCu). 5FGQ DMPK MD QWKKCRPW K?PIQ RFC @MBW NJ?LQ MD ?LGK?JQ GL RFC NFWJ? $RCLMNFMP? ?LB $LGB?PG?, GLAJSBGLE HCJJWDGQF ?LB ?BSJR QC? ?LCKMLCQ (Figure 27.7bc). 3?BG?J QWKKCRPW COSGNQ RFCQC QC? APC?RSPCQ (UFGAF K?W @C QCBCLR?PW MP MLJW A?N?@JC MD QJMU KMTCKCLR MP DJM?RGLE) RM CVNCPGCLAC RFC CLTGPMLKCLR COS?JJW DPMK ?JJ BGPCARGMLQ.
!@>LI< 27.7 4EB (>) PMLKDB FP >PVJJBQOF@>I. 4EB (?) GBIIVCFPE >KA (@) >KBJLKB >OB O>AF>IIV PVJJBQOF@>I, >KA QEB (A) ?RQQBOCIV FP ?FI>QBO>IIV PVJJBQOF@>I. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V !KAOBT 4ROKBO; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V 2L?BOQ &OBF?RODBO; @OBAFQ @: JLAFCF@>QFLK LC TLOH ?V 3>JRBI #ELT; @OBAFQ A: JLAFCF@>QFLK LC TLOH ?V #LOV :>KHBO)
Bilateral symmetry GLTMJTCQ RFC BGTGQGML MD RFC ?LGK?J RFPMSEF ? Q?EGRR?J NJ?LC, PCQSJRGLE GL RUM KGPPMP GK?EC, PGEFR ?LB JCDR F?JTCQ, QSAF ?Q RFMQC MD ? @SRRCPDJW (Figure 27.7d), AP?@, MP FSK?L @MBW. "LGK?JQ UGRF @GJ?RCP?J QWKKCRPW F?TC ? tFC?Bu ?LB tR?GJu (?LRCPGMP TQ. NMQRCPGMP), DPMLR ?LB @?AI (BMPQ?J TQ. TCLRP?J), ?LB PGEFR ?LB JCDR QGBCQ (Figure 27.8). "JJ RPSC ?LGK?JQ CVACNR RFMQC UGRF P?BG?J QWKKCRPW ?PC @GJ?RCP?JJW QWKKCRPGA?J. 5FC CTMJSRGML MD @GJ?RCP?J QWKKCRPW RF?R ?JJMUCB DMP RFC DMPK?RGML MD ?LRCPGMP ?LB NMQRCPGMP (FC?B ?LB R?GJ) CLBQ NPMKMRCB ? NFCLMKCLML A?JJCB ACNF?JGX?RGML, UFGAF PCDCPQ RM RFC AMJJCARGML MD ?L MPE?LGXCB LCPTMSQ QWQRCK ?R RFC ?LGK?JsQ ?LRCPGMP CLB. *L AMLRP?QR RM P?BG?J QWKKCRPW, UFGAF GQ @CQR QSGRCB DMP QR?RGML?PW MP JGKGRCB-KMRGML JGDCQRWJCQ, @GJ?RCP?J QWKKCRPW ?JJMUQ DMP QRPC?KJGLCB ?LB BGPCARGML?J KMRGML. *L CTMJSRGML?PW RCPKQ, RFGQ QGKNJC DMPK MD QWKKCRPW NPMKMRCB ?ARGTC KM@GJGRW ?LB GLAPC?QCB QMNFGQRGA?RGML MD PCQMSPAC-QCCIGLE ?LB NPCB?RMP-NPCW PCJ?RGMLQFGNQ.
734
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!@>LI< 27.8 4EB ?FI>QBO>IIV PVJJBQOF@>I ERJ>K ?LAV @>K ?B AFSFABA FKQL MI>KBP.
"LGK?JQ GL RFC NFWJSK &AFGLMBCPK?R? (QSAF ?Q QC? QR?PQ, Q?LB BMJJ?PQ, ?LB QC? SPAFGLQ) BGQNJ?W P?BG?J QWKKCRPW ?Q ?BSJRQ, @SR RFCGP J?PT?J QR?ECQ CVFG@GR @GJ?RCP?J QWKKCRPW. 5FGQ GQ RCPKCB QCAMLB?PW P?BG?J QWKKCRPW. 5FCW ?PC @CJGCTCB RM F?TC CTMJTCB DPMK @GJ?RCP?JJW QWKKCRPGA?J ?LGK?JQ; RFSQ, RFCW ?PC AJ?QQGDGCB ?Q @GJ?RCP?JJW QWKKCRPGA?J.
8?RAF RFGQ video (http://openstaxcollege.org/l/symmetry) RM QCC ? OSGAI QICRAF MD RFC BGDDCPCLR RWNCQ MD @MBW QWKKCRPW.
AE@D8C C?8I8:K<I@Q8K@FE B8J<; FE !<8KLI<J F= D9IPFCF>@:8C D<M<CFGD<EK

.MQR ?LGK?J QNCAGCQ SLBCPEM ? QCN?P?RGML MD RGQQSCQ GLRM ECPK J?WCPQ BSPGLE CK@PWMLGA BCTCJMNKCLR. 3CA?JJ RF?R RFCQC ECPK J?WCPQ ?PC DMPKCB BSPGLE E?QRPSJ?RGML, ?LB RF?R RFCW ?PC NPCBCRCPKGLCB RM BCTCJMN GLRM RFC ?LGK?JsQ QNCAG?JGXCB RGQQSCQ ?LB MPE?LQ. "LGK?JQ BCTCJMN CGRFCP RUM MP RFPCC CK@PWMLGA ECPKQ J?WCPQ (Figure 27.9). 5FC ?LGK?JQ RF?R BGQNJ?W P?BG?J QWKKCRPW BCTCJMN RUM ECPK J?WCPQ, ?L GLLCP J?WCP (CLBMBCPK) ?LB ?L MSRCP J?WCP (CARMBCPK). 5FCQC ?LGK?JQ ?PC A?JJCB diploblasts. %GNJM@J?QRQ F?TC ? LMLJGTGLE J?WCP @CRUCCL RFC CLBMBCPK ?LB CARMBCPK. .MPC AMKNJCV ?LGK?JQ (RFMQC UGRF @GJ?RCP?J QWKKCRPW) BCTCJMN RFPCC RGQQSC J?WCPQ: ?L GLLCP J?WCP (CLBMBCPK), ?L MSRCP J?WCP (CARMBCPK), ?LB ? KGBBJC J?WCP (KCQMBCPK). "LGK?JQ UGRF RFPCC RGQQSC J?WCPQ ?PC A?JJCB triploblasts.
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735
!@>LI< 27.9 $ROFKD BJ?OVLDBKBPFP, AFMIL?I>PQP ABSBILM QTL BJ?OVLKF@ DBOJ I>VBOP: >K B@QLABOJ >KA >K BKALABOJ. 4OFMIL?I>PQP ABSBILM > QEFOA I>VBOuQEB JBPLABOJu?BQTBBK QEB BKALABOJ >KA B@QLABOJ.
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&?AF MD RFC RFPCC ECPK J?WCPQ GQ NPMEP?KKCB RM EGTC PGQC RM N?PRGASJ?P @MBW RGQQSCQ ?LB MPE?LQ. 5FC CLBMBCPK EGTCQ PGQC RM RFC JGLGLE MD RFC BGECQRGTC RP?AR (GLAJSBGLE RFC QRMK?AF, GLRCQRGLCQ, JGTCP, ?LB N?LAPC?Q), ?Q UCJJ ?Q RM RFC JGLGLE MD RFC RP?AFC?, @PMLAFG, ?LB JSLEQ MD RFC PCQNGP?RMPW RP?AR, ?JMLE UGRF ? DCU MRFCP QRPSARSPCQ. 5FC CARMBCPK BCTCJMNQ GLRM RFC MSRCP CNGRFCJG?J AMTCPGLE MD RFC @MBW QSPD?AC, RFC ACLRP?J LCPTMSQ QWQRCK, ?LB ? DCU MRFCP QRPSARSPCQ. 5FC KCQMBCPK GQ RFC RFGPB ECPK J?WCP; GR DMPKQ @CRUCCL RFC CLBMBCPK ?LB CARMBCPK GL RPGNJM@J?QRQ. 5FGQ ECPK J?WCP EGTCQ PGQC RM ?JJ KSQAJC RGQQSCQ (GLAJSBGLE RFC A?PBG?A RGQQSCQ ?LB KSQAJCQ MD RFC GLRCQRGLCQ), AMLLCARGTC RGQQSCQ QSAF ?Q RFC QICJCRML ?LB @JMMB ACJJQ, ?LB KMQR MRFCP TGQACP?J MPE?LQ QSAF ?Q RFC IGBLCWQ ?LB RFC QNJCCL. P7+8+3)+ 47 A(8+3)+ 4, ' C4+142 'SPRFCP QS@BGTGQGML MD ?LGK?JQ UGRF RFPCC ECPK J?WCPQ (RPGNJM@J?QRQ) PCQSJRQ GL RFC QCN?P?RGML MD ?LGK?JQ RF?R K?W BCTCJMN ?L GLRCPL?J @MBW A?TGRW BCPGTCB DPMK KCQMBCPK, A?JJCB ? coelom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acoelomates, ?LB RFCGP KCQMBCPK PCEGML GQ AMKNJCRCJW DGJJCB UGRF RGQQSC, ?JRFMSEF RFCW BM QRGJJ F?TC ? ESR A?TGRW. &V?KNJCQ MD ?AMCJMK?RCQ GLAJSBC ?LGK?JQ GL RFC NFWJSK 1J?RWFCJKGLRFCQ, ?JQM ILMUL ?Q DJ?RUMPKQ. "LGK?JQ UGRF ? RPSC AMCJMK ?PC A?JJCB eucoelomates (MP AMCJMK?RCQ) (Figure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pseudocoelomates. 5FC NFWJSK /CK?RMB? (PMSLBUMPKQ) GQ ?L CV?KNJC MD ? NQCSBMAMCJMK?RC. 5PSC AMCJMK?RCQ A?L @C DSPRFCP AF?P?ARCPGXCB @?QCB ML ACPR?GL DC?RSPCQ MD RFCGP C?PJW CK@PWMJMEGA?J BCTCJMNKCLR.
736
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!@>LI< 27.10 4OFMIL?I>PQP J>V ?B (>) >@LBILJ>QBP, (?) BR@LBILJ>QBP, LO (@) MPBRAL@LBILJ>QBP. !@LBILJ>QBP E>SB KL ?LAV @>SFQV. %R@LBILJ>QBP E>SB > ?LAV @>SFQV TFQEFK QEB JBPLABOJ, @>IIBA > @LBILJ, TEF@E FP IFKBA TFQE JBPLABOJ. 0PBRAL@LBILJ>QBP >IPL E>SB > ?LAV @>SFQV, ?RQ FQ FP P>KATF@EBA ?BQTBBK QEB BKALABOJ >KA JBPLABOJ. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V *>K $BOH; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V ./!!; @OBAFQ @: JLAFCF@>QFLK LC TLOH ?V 53$!, !23)
E2(7>43/) D+;+1452+39 4, 9.+ M4:9. #GJ?RCP?JJW QWKKCRPGA?J, RPG@JM@J?QRGA CSAMCJMK?RCQ A?L @C DSPRFCP BGTGBCB GLRM RUM EPMSNQ @?QCB ML BGDDCPCLACQ GL RFCGP C?PJW CK@PWMLGA BCTCJMNKCLR. Protostomes GLAJSBC ?PRFPMNMBQ, KMJJSQIQ, ?LB ?LLCJGBQ. Deuterostomes GLAJSBC KMPC AMKNJCV ?LGK?JQ QSAF ?Q AFMPB?RCQ @SR ?JQM QMKC QGKNJC ?LGK?JQ QSAF ?Q CAFGLMBCPKQ. 5FCQC RUM EPMSNQ ?PC QCN?P?RCB @?QCB ML UFGAF MNCLGLE MD RFC BGECQRGTC A?TGRW BCTCJMNQ DGPQR: KMSRF MP ?LSQ. 5FC UMPB NPMRMQRMKC AMKCQ DPMK RFC (PCCI UMPB KC?LGLE tKMSRF DGPQR,u ?LB BCSRCPMQRMKC MPGEGL?RCQ DPMK RFC UMPB KC?LGLE tKMSRF QCAMLBu (GL RFGQ A?QC, RFC ?LSQ BCTCJMNQ DGPQR). 5FC KMSRF MP ?LSQ BCTCJMNQ DPMK ? QRPSARSPC A?JJCB RFC @J?QRMNMPC (Figure 27.11). 5FC blastopore GQ RFC GLBCLR?RGML DMPKCB BSPGLE RFC GLGRG?J QR?ECQ MD E?QRPSJ?RGML. *L J?RCP QR?ECQ, ? QCAMLB MNCLGLE DMPKQ, ?LB RFCQC RUM MNCLGLEQ UGJJ CTCLRS?JJW EGTC PGQC RM RFC KMSRF ?LB ?LSQ (Figure 27.11). *R F?Q JMLE @CCL @CJGCTCB RF?R RFC @J?QRMNMPC BCTCJMNQ GLRM RFC KMSRF MD NPMRMQRMKCQ, UGRF RFC QCAMLB MNCLGLE BCTCJMNGLE GLRM RFC ?LSQ; RFC MNNMQGRC GQ RPSC DMP BCSRCPMQRMKCQ. 3CACLR CTGBCLAC F?Q AF?JJCLECB RFGQ TGCU MD RFC BCTCJMNKCLR MD RFC @J?QRMNMPC MD NPMRMQRMKCQ, FMUCTCP, ?LB RFC RFCMPW PCK?GLQ SLBCP BC@?RC. "LMRFCP BGQRGLARGML @CRUCCL NPMRMQRMKCQ ?LB BCSRCPMQRMKCQ GQ RFC KCRFMB MD AMCJMK DMPK?RGML, @CEGLLGLE DPMK RFC E?QRPSJ? QR?EC. 5FC AMCJMK MD KMQR NPMRMQRMKCQ GQ DMPKCB RFPMSEF ? NPMACQQ A?JJCB schizocoely, KC?LGLE RF?R BSPGLE BCTCJMNKCLR, ? QMJGB K?QQ MD RFC KCQMBCPK QNJGRQ ?N?PR ?LB DMPKQ RFC FMJJMU MNCLGLE MD RFC AMCJMK. %CSRCPMQRMKCQ BGDDCP GL RF?R RFCGP AMCJMK DMPKQ RFPMSEF ? NPMACQQ A?JJCB enterocoely. )CPC, RFC KCQMBCPK BCTCJMNQ ?Q NMSAFCQ RF?R ?PC NGLAFCB MDD DPMK RFC CLBMBCPK RGQQSC. 5FCQC NMSAFCQ CTCLRS?JJW DSQC RM DMPK RFC KCQMBCPK, UFGAF RFCL EGTCQ PGQC RM RFC AMCJMK. 5FC C?PJGCQR BGQRGLARGML @CRUCCL NPMRMQRMKCQ ?LB BCSRCPMQRMKCQ GQ RFC RWNC MD AJC?T?EC SLBCPEMLC @W RFC XWEMRC. 1PMRMQRMKCQ SLBCPEM spiral cleavage, KC?LGLE RF?R RFC ACJJQ MD MLC NMJC MD RFC CK@PWM ?PC PMR?RCB, ?LB RFSQ KGQ?JGELCB, UGRF PCQNCAR RM RFC ACJJQ MD RFC MNNMQGRC NMJC. 5FGQ GQ BSC RM RFC M@JGOSC ?LEJC MD RFC AJC?T?EC. %CSRCPMQRMKCQ SLBCPEM radial cleavage, UFCPC RFC AJC?T?EC ?VCQ ?PC CGRFCP N?P?JJCJ MP NCPNCLBGASJ?P RM RFC NMJ?P ?VGQ, PCQSJRGLE GL RFC ?JGELKCLR MD RFC ACJJQ @CRUCCL RFC RUM NMJCQ.
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!@>LI< 27.11 %R@LBILJ>QBP @>K ?B AFSFABA FKQL QTL DOLRMP ?>PBA LK QEBFO B>OIV BJ?OVLKF@ ABSBILMJBKQ. )K MOLQLPQLJBP, M>OQ LC QEB JBPLABOJ PBM>O>QBP QL CLOJ QEB @LBILJ FK > MOL@BPP @>IIBA P@EFWL@LBIV. )K ABRQBOLPQLJBP, QEB JBPLABOJ MFK@EBP LCC QL CLOJ QEB @LBILJ FK > MOL@BPP @>IIBA BKQBOL@LBIV. )Q T>P ILKD ?BIFBSBA QE>Q QEB ?I>PQLMLOB ABSBILMBA FKQL QEB JLRQE FK MOLQLPQLJBP >KA FKQL QEB >KRP FK ABRQBOLPQLJBP, ?RQ OB@BKQ BSFABK@B @E>IIBKDBP QEFP ?BIFBC.
5FCPC GQ ? QCAMLB BGQRGLARGML @CRUCCL RFC RWNCQ MD AJC?T?EC GL NPMRMQRMKCQ ?LB BCSRCPMQRMKCQ. *L ?BBGRGML RM QNGP?J AJC?T?EC, NPMRMQRMKCQ ?JQM SLBCPEM determinate cleavage. 5FGQ KC?LQ RF?R CTCL ?R RFGQ C?PJW QR?EC, RFC BCTCJMNKCLR?J D?RC MD C?AF CK@PWMLGA ACJJ GQ ?JPC?BW BCRCPKGLCB. " ACJJ BMCQ LMR F?TC RFC ?@GJGRW RM BCTCJMN GLRM ?LW ACJJ RWNC. *L AMLRP?QR, BCSRCPMQRMKCQ SLBCPEM indeterminate cleavage, GL UFGAF ACJJQ ?PC LMR WCR NPC-BCRCPKGLCB ?R RFGQ C?PJW QR?EC RM BCTCJMN GLRM QNCAGDGA ACJJ RWNCQ. 5FCQC ACJJQ ?PC PCDCPPCB RM ?Q SLBGDDCPCLRG?RCB ACJJQ. 5FGQ AF?P?ARCPGQRGA MD BCSRCPMQRMKCQ GQ PCDJCARCB GL RFC CVGQRCLAC MD D?KGJG?P CK@PWMLGA QRCK ACJJQ, UFGAF F?TC RFC ?@GJGRW RM BCTCJMN GLRM ?LW ACJJ RWNC SLRGJ RFCGP D?RC GQ NPMEP?KKCB ?R ? J?RCP BCTCJMNKCLR?J QR?EC.
738
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27.3 R AE@D8C +?PCF><EP

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z *LRCPNPCR RFC KCR?XM?L NFWJMECLCRGA RPCC z %CQAPG@C RFC RWNCQ MD B?R? RF?R QAGCLRGQRQ SQC RM AMLQRPSAR ?LB PCTGQC ?LGK?J NFWJMECLW z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
CFEJKIL:K@E> 8E AE@D8C +?PCF><E<K@: /I<<

5FC ASPPCLR SLBCPQR?LBGLE MD CTMJSRGML?PW PCJ?RGMLQFGNQ @CRUCCL ?LGK?J, MP Metazoa, NFWJ? @CEGLQ UGRF RFC BGQRGLARGML @CRUCCL tRPSCu ?LGK?JQ UGRF RPSC BGDDCPCLRG?RCB RGQQSCQ, A?JJCB Eumetazoa, ?LB ?LGK?J NFWJ? RF?R BM LMR F?TC RPSC BGDDCPCLRG?RCB RGQQSCQ (QSAF ?Q RFC QNMLECQ), A?JJCB Parazoa. #MRF 1?P?XM? ?LB &SKCR?XM? CTMJTCB DPMK ? AMKKML ?LACQRP?J MPE?LGQK RF?R PCQCK@JCQ RFC KMBCPL-B?W NPMRGQRQ
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A?JJCB AFM?LMDJ?ECJJ?RCQ. 5FCQC NPMRGQR ACJJQ QRPMLEJW PCQCK@JC RFC QNMLEC AFM?LMAWRC ACJJQ RMB?W (Figure 27.12).
!@>LI< 27.12 #BIIP LC QEB MOLQFPQ @EL>KLCI>DBII>QB OBPBJ?IB PMLKDB @EL>KL@VQB @BIIP. "B>QFKD LC @EL>KL@VQB CI>DBII> AO>TP T>QBO QEOLRDE QEB PMLKDB PL QE>Q KRQOFBKQP @>K ?B BUQO>@QBA >KA T>PQB OBJLSBA.
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igure 27.13?@). Ecdysozoa GLAJSBCQ LCK?RMBCQ ?LB ?PRFPMNMBQ; RFCW ?PC QM L?KCB DMP ? AMKKMLJW DMSLB AF?P?ARCPGQRGA ?KMLE RFC EPMSN: CVMQICJCR?J KMJRGLE (RCPKCB CABWQGQ). Lophotrochozoa GQ L?KCB DMP RUM QRPSARSP?J DC?RSPCQ, C?AF AMKKML RM ACPR?GL NFWJ? UGRFGL RFC AJ?BC. 4MKC JMNFMRPMAFMXM?L NFWJ? ?PC AF?P?ARCPGXCB @W ? J?PT?J QR?EC A?JJCB RPMAFMNFMPC J?PT?C, ?LB MRFCP NFWJ? ?PC AF?P?ARCPGXCB @W RFC NPCQCLAC MD ? DCCBGLE QRPSARSPC A?JJCB ? JMNFMNFMPC.
!@>LI< 27.13 !KFJ>IP QE>Q JLIQ QEBFO BULPHBIBQLKP, PR@E >P QEBPB (>) ->A>D>P@>O EFPPFKD @L@HOL>@EBP, >OB FK QEB @I>AB %@AVPLWL>. (?) 0ELOLKFAP >OB FK QEB @I>AB ,LMELQOL@ELWL>. 4EB QBKQ>@IBP >OB M>OQ LC > CBBAFKD PQOR@QROB @>IIBA > ILMELMELOB. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V 7EFQKBV #O>KPE>T, #LILO>AL 3Q>QB 5KFSBOPFQV, "RDTLLA.LOD; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V ./!!)
&VNJMPC ?L GLRCP?ARGTC tree (http://openstaxcollege.org/l/tree_of_life2) MD JGDC FCPC. ;MMK ?LB AJGAI RM JC?PL KMPC ?@MSR RFC MPE?LGQKQ ?LB RFCGP CTMJSRGML?PW PCJ?RGMLQFGNQ.
(F;<IE A;M8E:<J @E +?PCF><E<K@: 0E;<IJK8E;@E> CFD< =IFD (FC<:LC8I AE8CPJ<J

5FC NFWJMECLCRGA EPMSNGLEQ ?PC AMLRGLS?JJW @CGLE BC@?RCB ?LB PCDGLCB @W CTMJSRGML?PW @GMJMEGQRQ. &?AF WC?P, LCU CTGBCLAC CKCPECQ RF?R DSPRFCP ?JRCPQ RFC PCJ?RGMLQFGNQ BCQAPG@CB @W ? NFWJMECLCRGA RPCC BG?EP?K.
740
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8?RAF RFC DMJJMUGLE video (http://openstaxcollege.org/l/build_phylogeny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tDJ?RUMPKu KMPNFMJMEW. )MUCTCP, KMJCASJ?P ?L?JWQCQ PCTC?JCB RFGQ RM @C ? D?JQC PCJ?RGMLQFGN ?LB MPGEGL?JJW QSEECQRCB RF?R ?AMCJQ PCNPCQCLRCB JGTGLE QNCAGCQ MD QMKC MD RFC C?PJGCQR BGTCPECLR @GJ?RCPG?LQ. .MPC PCACLR PCQC?PAF GLRM RFC ?AMCJMKMPNFQ F?Q A?JJCB RFGQ FWNMRFCQGQ GLRM OSCQRGML ?LB QSEECQRCB ? AJMQCP PCJ?RGMLQFGN UGRF BCSRCPMQRMKCQ. 5FC NJ?ACKCLR MD RFGQ LCU NFWJSK PCK?GLQ BGQNSRCB, @SR QAGCLRGQRQ ?EPCC RF?R UGRF QSDDGAGCLR KMJCASJ?P B?R?, RFCGP RPSC NFWJMECLW UGJJ @C BCRCPKGLCB.
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742
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&V?KNJCQ MD RFCQC AF?LECQ GLAJSBC PGQGLE ?RKMQNFCPGA MVWECL JCTCJQ ?LB J?PEC GLAPC?QCQ GL MAC?LGA A?JAGSK AMLACLRP?RGMLQ RF?R NPCACBCB RFC $?K@PG?L NCPGMB (Figure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8:F<CFD8K< ?LGK?J UGRFMSR ? @MBW A?TGRW 9@C8K<I8C JPDD<KIP RWNC MD QWKKCRPW GL UFGAF RFCPC GQ MLJW MLC NJ?LC MD QWKKCRPW, QM RFC JCDR ?LB PGEFR F?JTCQ MD ?L ?LGK?J ?PC KGPPMP GK?ECQ 9C8JKFGFI< GLBCLR?RGML DMPKCB BSPGLE E?QRPSJ?RGML, CTGBCLR GL RFC E?QRPSJ? QR?EC 9C8JKLC8 16p32 ACJJ QR?EC MD BCTCJMNKCLR MD ?L ?LGK?J CK@PWM 9F;P GC8E KMPNFMJMEW MP AMLQR?LR QF?NC MD ?L MPE?LGQK C8D9I@8E <OGCFJ@FE RGKC BSPGLE RFC $?K@PG?L NCPGMB (542p488 KGJJGML WC?PQ ?EM) UFCL KMQR MD RFC ?LGK?J NFWJ? GL CVGQRCLAC RMB?W CTMJTCB CIPF><E@8E G<I@F; ECMJMEGA NCPGMB (850p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p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tQRCK ACJJQu ?PC LMR WCR NPC-BCRCPKGLCB RM BCTCJMN GLRM QNCAGDGA ACJJ RWNCQ 'FG?FKIF:?FQF8 AJ?BC MD NPMRMQRMKCQ RF?R CVFG@GR ? RPMAFMNFMPC J?PT?C QR?EC MP ? JMNFMNFMPC DCCBGLE QRPSARSPC (<K8QF8 EPMSN AMLR?GLGLE ?JJ ?LGK?JQ D8JJ <OK@E:K@FE CTCLR RF?R UGNCQ MSR RFC K?HMPGRW MD QNCAGCQ UGRFGL ? PCJ?RGTCJW QFMPR ECMJMEGA?J RGKC NCPGMB FI>8EF><E<J@J DMPK?RGML MD MPE?LQ GL ?LGK?J CK@PWMECLCQGQ +8I8QF8 EPMSN MD ?LGK?JQ UGRFMSR RPSC BGDDCPCLRG?RCB RGQQSCQ
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
27.3 AE@D8C +?PCF><EP

4AGCLRGQRQ ?PC GLRCPCQRCB GL RFC CTMJSRGML?PW FGQRMPW MD ?LGK?JQ ?LB RFC CTMJSRGML?PW PCJ?RGMLQFGNQ ?KMLE RFCK. 5FCPC ?PC RFPCC K?GL QMSPACQ MD B?R? RF?R QAGCLRGQRQ SQC RM APC?RC NFWJMECLCRGA CTMJSRGML?PW RPCC BG?EP?KQ RF?R GJJSQRP?RC QSAF PCJ?RGMLQFGNQ: KMPNFMJMEGA?J GLDMPK?RGML (UFGAF GLAJSBCQ BCTCJMNKCLR?J KMPNFMJMEGCQ), DMQQGJ PCAMPB B?R?, ?LB, KMQR PCACLRJW, KMJCASJ?P B?R?. 5FC BCR?GJQ MD RFC KMBCPL NFWJMECLCRGA RPCC AF?LEC DPCOSCLRJW ?Q LCU B?R? ?PC E?RFCPCB, ?LB KMJCASJ?P B?R? F?Q PCACLRJW AMLRPG@SRCB RM K?LW QS@QR?LRG?J KMBGDGA?RGMLQ MD RFC SLBCPQR?LBGLE MD PCJ?RGMLQFGNQ @CRUCCL ?LGK?J NFWJ?.
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5FC KMQR P?NGB BGTCPQGDGA?RGML ?LB CTMJSRGML MD ?LGK?J QNCAGCQ GL ?JJ MD FGQRMPW MAASPPCB BSPGLE RFC $?K@PG?L NCPGMB MD RFC 1?JCMXMGA &P?, ? NFCLMKCLML ILMUL ?Q RFC $?K@PG?L CVNJMQGML. 6LRGJ PCACLRJW, QAGCLRGQRQ @CJGCTCB RF?R RFCPC UCPC MLJW TCPW DCU RGLW ?LB QGKNJGQRGA ?LGK?J QNCAGCQ GL CVGQRCLAC @CDMPC
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1. Figure 27.5 *D ? H6? 13 ECLC GL ? KMSQC U?Q PCNJ?ACB UGRF ? H6? 1 ECLC, FMU KGEFR RFGQ ?JRCP ?LGK?J BCTCJMNKCLR 2. Figure 27.6 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ GQ D?JQC >. &SKCR?XM?LQ F?TC QNCAG?JGXCB RGQQSCQ ?LB N?P?XM?LQ BMLsR. ?. -MNFMRPMAFMXM? ?LB &ABWQMXM? ?PC @MRF #GJ?R?PG?. @. "AMCJ? ?LB $LGB?PG? @MRF NMQQCQQ P?BG?J QWKKCRPW. A. "PRFPMNMBQ ?PC KMPC AJMQCJW PCJ?RCB RM LCK?RMBCQ RF?L RFCW ?PC RM ?LLCJGBQ. 3. Figure 27.9 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR BGNJM@J?QRQ ?LB RPGNJM@J?QRQ GQ D?JQC >. "LGK?JQ RF?R BGQNJ?W P?BG?J QWKKCRPW ?PC BGNJM@J?QRQ. ?. "LGK?JQ RF?R BGQNJ?W @GJ?RCP?J QWKKCRPW ?PC RPGNJM@J?QRQ. @. 5FC CLBMBCPK EGTCQ PGQC RM RFC JGLGLE MD RFC BGECQRGTC RP?AR ?LB RFC PCQNGP?RMPW RP?AR. A. 5FC KCQMBCPK EGTCQ PGQC RM RFC ACLRP?J LCPTMSQ QWQRCK.
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4. 8FGAF MD RFC DMJJMUGLE GQ LMR ? DC?RSPC AMKKML RM 46:; ?LGK?JQ >. BCTCJMNKCLR GLRM ? DGVCB @MBW NJ?L ?. ?QCVS?J PCNPMBSARGML @. QNCAG?JGXCB RGQQSCQ A. FCRCPMRPMNFGA LSRPGCLR QMSPAGLE 5. %SPGLE CK@PWMLGA BCTCJMNKCLR, SLGOSC ACJJ J?WCPQ BCTCJMN ?LB BGQRGLESGQF BSPGLE ? QR?EC A?JJCB >>>>>>>>. >. RFC @J?QRSJ? QR?EC ?. RFC ECPK J?WCP QR?EC @. RFC E?QRPSJ? QR?EC A. RFC MPE?LMECLCQGQ QR?EC 6. 8FGAF MD RFC DMJJMUGLE NFCLMRWNCQ UMSJB KMQR JGICJW @C RFC PCQSJR MD ? H6? ECLC KSR?RGML >. ?. @. A. ?@LMPK?J @MBW JCLERF MP FCGEFR RUM BGDDCPCLR CWC AMJMPQ RFC AMLRP?ARGML MD ? ECLCRGA GJJLCQQ RUM DCUCP ?NNCLB?ECQ RF?L LMPK?J 9. "L ?LGK?J UFMQC BCTCJMNKCLR GQ K?PICB @W P?BG?J AJC?T?EC ?LB CLRCPMAMCJW GQ >>>>>>>>. >. ?. @. A. ? BCSRCPMQRMKC ?L ?LLCJGB MP KMJJSQI CGRFCP ?L ?AMCJMK?RC MP CSAMCJMK?RC LMLC MD RFC ?@MTC
10. $MLQSJRGLE RFC KMBCPL NFWJMECLCRGA RPCC MD ?LGK?JQ, UFGAF MD RFC DMJJMUGLE UMSJB LMR AMLQRGRSRC ? AJ?BC >. BCSRCPMQRMKCQ ?. JMNFMRPMAFMXM?LQ @. 1?P?XM? A. #GJ?RCPG? 11. 8FGAF MD RFC DMJJMUGLE GQ RFMSEFR RM @C RFC KMQR AJMQCJW PCJ?RCB RM RFC AMKKML ?LGK?J ?LACQRMP >. DSLE?J ACJJQ ?. NPMRGQR ACJJQ @. NJ?LR ACJJQ A. @?ARCPG?J ACJJQ 12. "Q UGRF RFC CKCPECLAC MD RFC "AMCJMKMPNF? NFWJSK, GR GQ AMKKML DMP >>>> B?R? RM KGQNJ?AC ?LGK?JQ GL AJMQC PCJ?RGML RM MRFCP QNCAGCQ, UFCPC?Q >>>> B?R? MDRCL PCTC?JQ ? BGDDCPCLR ?LB KMPC ?AASP?RC CTMJSRGML?PW PCJ?RGMLQFGN. >. KMJCASJ?P : KMPNFMJMEGA?J ?. KMJCASJ?P : DMQQGJ PCAMPB @. DMQQGJ PCAMPB : KMPNFMJMEGA?J A. KMPNFMJMEGA?J : KMJCASJ?P
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sQ QNCAGCQ ?PC ?OS?RGA GLTCPRC@P?RCQ RF?R F?TC LMR WCR @CCL BMASKCLRCB.
28.1 R +?PCLD +FI@=<I8

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC MPE?LGX?RGML?J DC?RSPCQ MD RFC QGKNJCQR KSJRGACJJSJ?P MPE?LGQKQ z &VNJ?GL RFC T?PGMSQ @MBW DMPKQ ?LB @MBGJW DSLARGMLQ MD QNMLECQ
752
#(!04%2 28 X ).6%24%"2!4%3
5FC GLTCPRC@P?RCQ, MP invertebrata, ?PC ?LGK?JQ RF?R BM LMR AMLR?GL @MLW QRPSARSPCQ, QSAF ?Q RFC AP?LGSK ?LB TCPRC@P?C. 5FC QGKNJCQR MD ?JJ RFC GLTCPRC@P?RCQ ?PC RFC 1?P?XM?LQ, UFGAF GLAJSBC MLJW RFC NFWJSK Porifera: RFC QNMLECQ (Figure 28.2). 1?P?XM?LQ (t@CQGBC ?LGK?JQu) BM LMR BGQNJ?W RGQQSC-JCTCJ MPE?LGX?RGML, ?JRFMSEF RFCW BM F?TC QNCAG?JGXCB ACJJQ RF?R NCPDMPK QNCAGDGA DSLARGMLQ. 4NMLEC J?PT?C ?PC ?@JC RM QUGK; FMUCTCP, ?BSJRQ ?PC LML-KMRGJC ?LB QNCLB RFCGP JGDC ?RR?AFCB RM ? QS@QRP?RSK. 4GLAC U?RCP GQ TGR?J RM QNMLECQ DMP CVAPCRGML, DCCBGLE, ?LB E?Q CVAF?LEC, RFCGP @MBW QRPSARSPC D?AGJGR?RCQ RFC KMTCKCLR MD U?RCP RFPMSEF RFC QNMLEC. 4RPSARSPCQ QSAF ?Q A?L?JQ, AF?K@CPQ, ?LB A?TGRGCQ CL?@JC U?RCP RM KMTC RFPMSEF RFC QNMLEC RM LC?PJW ?JJ @MBW ACJJQ.
!@>LI< 28.2 3MLKDBP >OB JBJ?BOP LC QEB 0EVIRJ 0LOFCBO>, TEF@E @LKQ>FKP QEB PFJMIBPQ FKSBOQB?O>QBP. (@OBAFQ: !KAOBT 4ROKBO)
(FIG?FCF>P F= .GFE><J
5FC KMPNFMJMEW MD RFC QGKNJCQR QNMLECQ R?ICQ RFC QF?NC MD ? AWJGLBCP UGRF ? J?PEC ACLRP?J A?TGRW, RFC spongocoel, MAASNWGLE RFC GLQGBC MD RFC AWJGLBCP. 8?RCP A?L CLRCP GLRM RFC QNMLEMAMCJ DPMK LSKCPMSQ NMPCQ GL RFC @MBW U?JJ. 8?RCP CLRCPGLE RFC QNMLEMAMCJ GQ CVRPSBCB TG? ? J?PEC AMKKML MNCLGLE A?JJCB RFC osculum. )MUCTCP, QNMLECQ CVFG@GR ? P?LEC MD BGTCPQGRW GL @MBW DMPKQ, GLAJSBGLE T?PG?RGMLQ GL RFC QGXC MD RFC QNMLEMAMCJ, RFC LSK@CP MD MQASJG, ?LB UFCPC RFC ACJJQ RF?R DGJRCP DMMB DPMK RFC U?RCP ?PC JMA?RCB. 8FGJC QNMLECQ (CVAJSBGLE RFC FCV?ARGLCJJGBQ) BM LMR CVFG@GR RGQQSC-J?WCP MPE?LGX?RGML, RFCW BM F?TC BGDDCPCLR ACJJ RWNCQ RF?R NCPDMPK BGQRGLAR DSLARGMLQ. Pinacocytes, UFGAF ?PC CNGRFCJG?J-JGIC ACJJQ, DMPK RFC MSRCPKMQR J?WCP MD QNMLECQ ?LB CLAJMQC ? HCJJW-JGIC QS@QR?LAC A?JJCB KCQMFWJ. Mesohyl GQ ?L CVRP?ACJJSJ?P K?RPGV AMLQGQRGLE MD ? AMJJ?ECL-JGIC ECJ UGRF QSQNCLBCB ACJJQ RF?R NCPDMPK T?PGMSQ DSLARGMLQ. 5FC ECJ-JGIC AMLQGQRCLAW MD KCQMFWJ ?ARQ JGIC ?L CLBMQICJCRML ?LB K?GLR?GLQ RFC RS@SJ?P KMPNFMJMEW MD QNMLECQ. *L ?BBGRGML RM RFC MQASJSK, QNMLECQ F?TC KSJRGNJC NMPCQ A?JJCB ostia ML RFCGP @MBGCQ RF?R ?JJMU U?RCP RM CLRCP RFC QNMLEC. *L QMKC QNMLECQ, MQRG? ?PC DMPKCB @W NMPMAWRCQ, QGLEJC RS@C-QF?NCB ACJJQ RF?R ?AR ?Q T?JTCQ RM PCESJ?RC RFC DJMU MD U?RCP GLRM RFC QNMLEMAMCJ. *L MRFCP QNMLECQ, MQRG? ?PC DMPKCB @W DMJBQ GL RFC @MBW U?JJ MD RFC QNMLEC. Choanocytes (tAMJJ?P ACJJQu) ?PC NPCQCLR ?R T?PGMSQ JMA?RGMLQ, BCNCLBGLE ML RFC RWNC MD QNMLEC, @SR RFCW ?JU?WQ JGLC RFC GLLCP NMPRGMLQ MD QMKC QN?AC RFPMSEF UFGAF U?RCP DJMUQ (RFC QNMLEMAMCJ GL QGKNJC QNMLECQ, A?L?JQ UGRFGL RFC @MBW U?JJ GL KMPC AMKNJCV QNMLECQ, ?LB AF?K@CPQ QA?RRCPCB RFPMSEFMSR RFC @MBW GL RFC KMQR AMKNJCV QNMLECQ). 8FCPC?Q NGL?AMAWRCQ JGLC RFC MSRQGBC MD RFC QNMLEC, AFM?LMAWRCQ RCLB RM JGLC ACPR?GL GLLCP NMPRGMLQ MD RFC QNMLEC @MBW RF?R QSPPMSLB RFC KCQMFWJ. 5FC QRPSARSPC MD ? AFM?LMAWRC GQ APGRGA?J RM GRQ DSLARGML, UFGAF GQ RM ECLCP?RC ? U?RCP ASPPCLR RFPMSEF RFC QNMLEC ?LB RM RP?N ?LB GLECQR DMMB N?PRGAJCQ @W NF?EMAWRMQGQ. /MRC RFC QGKGJ?PGRW GL ?NNC?P?LAC @CRUCCL RFC QNMLEC AFM?LMAWRC ?LB AFM?LMDJ?ECJJ?RCQ (1PMRGQR?). 5FGQ QGKGJ?PGRW QSEECQRQ RF?R QNMLECQ ?LB AFM?LMDJ?ECJJ?RCQ ?PC AJMQCJW PCJ?RCB ?LB JGICJW QF?PC ? PCACLR AMKKML ?LACQRPW. 5FC ACJJ @MBW GQ CK@CBBCB GL KCQMFWJ ?LB AMLR?GLQ ?JJ MPE?LCJJCQ PCOSGPCB DMP LMPK?J ACJJ DSLARGML, @SR NPMRPSBGLE GLRM RFC tMNCL QN?ACu GLQGBC MD RFC QNMLEC GQ ? KCQF-JGIC AMJJ?P AMKNMQCB MD KGAPMTGJJG UGRF ? QGLEJC DJ?ECJJSK GL RFC ACLRCP MD RFC AMJSKL. 5FC ASKSJ?RGTC CDDCAR MD RFC DJ?ECJJ? DPMK ?JJ AFM?LMAWRCQ ?GBQ RFC KMTCKCLR MD U?RCP RFPMSEF RFC QNMLEC: BP?UGLE U?RCP GLRM RFC QNMLEC RFPMSEF RFC LSKCPMSQ MQRG?, GLRM RFC QN?ACQ JGLCB @W AFM?LMAWRCQ, ?LB CTCLRS?JJW MSR RFPMSEF RFC MQASJSK (MP MQASJG). *L RFC KC?LRGKC, DMMB N?PRGAJCQ, GLAJSBGLE U?RCP@MPLC @?ARCPG? ?LB ?JE?C, ?PC RP?NNCB @W RFC QGCTC-JGIC AMJJ?P MD RFC AFM?LMAWRCQ, QJGBC BMUL GLRM RFC @MBW MD RFC ACJJ, ?PC GLECQRCB @W NF?EMAWRMQGQ, ?LB @CAMKC CLA?QCB GL ? DMMB T?ASMJC. -?QRJW, AFM?LMAWRCQ UGJJ BGDDCPCLRG?RC GLRM QNCPK DMP QCVS?J PCNPMBSARGML, UFCPC RFCW UGJJ @CAMKC BGQJMBECB DPMK RFC KCQMFWJ ?LB JC?TC RFC QNMLEC UGRF CVNCJJCB U?RCP RFPMSEF RFC MQASJSK.
#(!04%2 28 X ).6%24%"2!4%3
753
8?RAF RFGQ video (http://openstaxcollege.org/l/filter_sponges) RM QCC RFC KMTCKCLR MD U?RCP RFPMSEF RFC QNMLEC @MBW. 5FC QCAMLB APSAG?J ACJJQ GL QNMLECQ ?PC A?JJCB amoebocytes (MP ?PAF?CMAWRCQ), L?KCB DMP RFC D?AR RF?R RFCW KMTC RFPMSEFMSR RFC KCQMFWJ GL ?L ?KMC@?-JGIC D?QFGML. "KMC@MAWRCQ F?TC ? T?PGCRW MD DSLARGMLQ: BCJGTCPGLE LSRPGCLRQ DPMK AFM?LMAWRCQ RM MRFCP ACJJQ UGRFGL RFC QNMLEC, EGTGLE PGQC RM CEEQ DMP QCVS?J PCNPMBSARGML (UFGAF PCK?GL GL RFC KCQMFWJ), BCJGTCPGLE NF?EMAWRGXCB QNCPK DPMK AFM?LMAWRCQ RM CEEQ, ?LB BGDDCPCLRG?RGLE GLRM KMPC-QNCAGDGA ACJJ RWNCQ. 4MKC MD RFCQC KMPC-QNCAGDGA ACJJ RWNCQ GLAJSBC AMJJCLAWRCQ ?LB JMNFMAWRCQ, UFGAF NPMBSAC RFC AMJJ?ECL-JGIC NPMRCGL RM K?GLR?GL RFC KCQMFWJ, QAJCPMAWRCQ, UFGAF NPMBSAC QNGASJCQ GL QMKC QNMLECQ, ?LB QNMLEMAWRCQ, UFGAF NPMBSAC RFC NPMRCGL QNMLEGL GL RFC K?HMPGRW MD QNMLECQ. 5FCQC ACJJQ NPMBSAC AMJJ?ECL RM K?GLR?GL RFC AMLQGQRCLAW MD RFC KCQMFWJ. 5FC BGDDCPCLR ACJJ RWNCQ GL QNMLECQ ?PC QFMUL GL Figure 28.3.
!@>LI< 28.3 4EB PMLKDBwP (>) ?>PF@ ?LAV MI>K >KA (?) PLJB LC QEB PMB@F>IFWBA @BII QVMBP CLRKA FK PMLKDBP >OB PELTK.
7EF@E LC QEB CLIILTFKD PQ>QBJBKQP FP C>IPB >. #EL>KL@VQBP E>SB CI>DBII> QE>Q MOLMBI T>QBO QEOLRDE QEB ?LAV. ?. 0FK>@L@VQBP @>K QO>KPCLOJ FKQL >KV @BII QVMB. @. ,LMEL@VQBP PB@OBQB @LII>DBK. A. 0LOL@VQBP @LKQOLI QEB CILT LC T>QBO QEOLRDE MLOBP FK QEB PMLKDB ?LAV. *L QMKC QNMLECQ, sclerocytes QCAPCRC QK?JJ spicules GLRM RFC KCQMFWJ, UFGAF ?PC AMKNMQCB MD CGRFCP A?JAGSK A?P@ML?RC MP QGJGA?, BCNCLBGLE ML RFC RWNC MD QNMLEC. 5FCQC QNGASJCQ QCPTC RM NPMTGBC ?BBGRGML?J QRGDDLCQQ RM RFC @MBW MD RFC QNMLEC. "BBGRGML?JJW, QNGASJCQ, UFCL NPCQCLR CVRCPL?JJW, K?W U?PB MDD NPCB?RMPQ. "LMRFCP RWNC MD NPMRCGL, QNMLEGL, K?W ?JQM @C NPCQCLR GL RFC KCQMFWJ MD QMKC QNMLECQ.
754
#(!04%2 28 X ).6%24%"2!4%3
5?IC ?L SN-AJMQC tour (http://openstaxcollege.org/l/sponge_ride) RFPMSEF RFC QNMLEC ?LB GRQ ACJJQ. 5FC NPCQCLAC ?LB AMKNMQGRGML MD QNGASJCQ/QNMLEGL ?PC RFC BGDDCPCLRG?RGLE AF?P?ARCPGQRGAQ MD RFC RFPCC AJ?QQCQ MD QNMLECQ (Figure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
!@>LI< 28.4 (>) C3(;/905( *3(;/9<: ?BILKDP QL @I>PP #>I@>OB>, (?) ;(<96*(3@7;<: PMM. (@LJJLK K>JB: VBIILT 0F@>PPL PMLKDB) ?BILKDP QL @I>PP (BU>@QFKBIIFA>, >KA (@) A*(95<: ,90;/(*<: ?BILKDP QL @I>PP $BJLPMLKDF>. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V 0>OBKQ 'cOV; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V -LKQBOBV ">V !NR>OFRJ 2BPB>O@E )KPQFQRQB, ./!!; @OBAFQ @: JLAFCF@>QFLK LC TLOH ?V 3>K@QR>OV )KQBDO>QBA -LKFQLOFKD .BQTLOH, -LKQBOBV ">V .>QFLK>I ->OFKB 3>K@QR>OV, ./!!)
6QC RFC Interactive Sponge Guide (http://openstaxcollege.org/l/id_sponges) RM GBCLRGDW QNCAGCQ MD QNMLECQ @?QCB ML RFCGP CVRCPL?J DMPK, KGLCP?J QICJCRML, DG@CP, ?LB QICJCR?J ?PAFGRCARSPC.
+?PJ@FCF>@:8C +IF:<JJ<J @E .GFE><J

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
#(!04%2 28 X ).6%24%"2!4%3
755
"JJ MRFCP K?HMP @MBW DSLARGMLQ GL RFC QNMLEC (E?Q CVAF?LEC, AGPASJ?RGML, CVAPCRGML) ?PC NCPDMPKCB @W BGDDSQGML @CRUCCL RFC ACJJQ RF?R JGLC RFC MNCLGLEQ UGRFGL RFC QNMLEC ?LB RFC U?RCP RF?R GQ N?QQGLE RFPMSEF RFMQC MNCLGLEQ. "JJ ACJJ RWNCQ UGRFGL RFC QNMLEC M@R?GL MVWECL DPMK U?RCP RFPMSEF BGDDSQGML. -GICUGQC, A?P@ML BGMVGBC GQ PCJC?QCB GLRM QC?U?RCP @W BGDDSQGML. *L ?BBGRGML, LGRPMECLMSQ U?QRC NPMBSACB ?Q ? @WNPMBSAR MD NPMRCGL KCR?@MJGQK GQ CVAPCRCB TG? BGDDSQGML @W GLBGTGBS?J ACJJQ GLRM RFC U?RCP ?Q GR N?QQCQ RFPMSEF RFC QNMLEC. +574*:)9/43 4NMLECQ PCNPMBSAC @W QCVS?J ?Q UCJJ ?Q ?QCVS?J KCRFMBQ. 5FC RWNGA?J KC?LQ MD ?QCVS?J PCNPMBSARGML GQ CGRFCP DP?EKCLR?RGML (UFCPC ? NGCAC MD RFC QNMLEC @PC?IQ MDD, QCRRJCQ ML ? LCU QS@QRP?RC, ?LB BCTCJMNQ GLRM ? LCU GLBGTGBS?J) MP @SBBGLE (? ECLCRGA?JJW GBCLRGA?J MSREPMURF EPMUQ DPMK RFC N?PCLR ?LB CTCLRS?JJW BCR?AFCQ MP PCK?GLQ ?RR?AFCB RM DMPK ? AMJMLW). "L ?RWNGA?J RWNC MD ?QCVS?J PCNPMBSARGML GQ DMSLB MLJW GL DPCQFU?RCP QNMLECQ ?LB MAASPQ RFPMSEF RFC DMPK?RGML MD ECKKSJCQ. Gemmules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
8?RAF RFGQ ##$ video (http://openstaxcollege.org/l/sea_sponges) QFMUGLE RFC ?PP?W MD QNMLECQ QCCL ?JMLE RFC $?WK?L 8?JJ BSPGLE ? QS@KCPQG@JC BGTC.
28.2 R +?PCLD CE@;8I@8

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z $MKN?PC QRPSARSP?J ?LB MPE?LGX?RGML AF?P?ARCPGQRGAQ MD 1MPGDCP? ?LB $LGB?PG? z %CQAPG@C RFC NPMEPCQQGTC BCTCJMNKCLR MD RGQQSCQ ?LB RFCGP PCJCT?LAC RM ?LGK?J AMKNJCVGRW
756
#(!04%2 28 X ).6%24%"2!4%3
1FWJSK Cnidaria GLAJSBCQ ?LGK?JQ RF?R QFMU P?BG?J MP @GP?BG?J QWKKCRPW ?LB ?PC BGNJM@J?QRGA, RF?R GQ, RFCW BCTCJMN DPMK RUM CK@PWMLGA J?WCPQ. /C?PJW ?JJ (?@MSR 99 NCPACLR) ALGB?PG?LQ ?PC K?PGLC QNCAGCQ. $LGB?PG?LQ AMLR?GL QNCAG?JGXCB ACJJQ ILMUL ?Q cnidocytes (tQRGLEGLE ACJJQu) AMLR?GLGLE MPE?LCJJCQ A?JJCB nematocysts (QRGLECPQ). 5FCQC ACJJQ ?PC NPCQCLR ?PMSLB RFC KMSRF ?LB RCLR?AJCQ, ?LB QCPTC RM GKKM@GJGXC NPCW UGRF RMVGLQ AMLR?GLCB UGRFGL RFC ACJJQ. /CK?RMAWQRQ AMLR?GL AMGJCB RFPC?BQ RF?R K?W @C?P @?P@Q. 5FC MSRCP U?JJ MD RFC ACJJ F?Q F?GPJGIC NPMHCARGMLQ A?JJCB ALGBMAGJQ, UFGAF ?PC QCLQGRGTC RM RMSAF. 8FCL RMSAFCB, RFC ACJJQ ?PC ILMUL RM DGPC AMGJCB RFPC?BQ RF?R A?L CGRFCP NCLCRP?RC RFC DJCQF MD RFC NPCW MP NPCB?RMPQ MD ALGB?PG?LQ (QCC Figure 28.5) MP CLQL?PC GR. 5FCQC AMGJCB RFPC?BQ PCJC?QC RMVGLQ GLRM RFC R?PECR ?LB A?L MDRCL GKKM@GJGXC NPCW MP QA?PC ?U?W NPCB?RMPQ.
!@>LI< 28.5 !KFJ>IP COLJ QEB MEVIRJ #KFA>OF> E>SB PQFKDFKD @BIIP @>IIBA @KFAL@VQBP. #KFAL@VQBP @LKQ>FK I>ODB LOD>KBIIBP @>IIBA (>) KBJ>QL@VPQP QE>Q PQLOB > @LFIBA QEOB>A >KA ?>O?. 7EBK E>FOIFHB MOLGB@QFLKP LK QEB @BII PROC>@B >OB QLR@EBA, (?) QEB QEOB>A, ?>O?, >KA > QLUFK >OB CFOBA COLJ QEB LOD>KBIIB.
7GCU RFGQ video (http://openstaxcollege.org/l/nematocyst) ?LGK?RGML QFMUGLE RUM ?LCKMLCQ CLE?ECB GL ? @?RRJC. "LGK?JQ GL RFGQ NFWJSK BGQNJ?W RUM BGQRGLAR KMPNFMJMEGA?J @MBW NJ?LQ: polyp MP tQR?JIu ?LB medusa MP t@CJJu (Figure 28.6). "L CV?KNJC MD RFC NMJWN DMPK GQ H@+9( QNN.; NCPF?NQ RFC KMQR UCJJILMUL KCBSQMGB ?LGK?JQ ?PC RFC HCJJGCQ (HCJJWDGQF). 1MJWN DMPKQ ?PC QCQQGJC ?Q ?BSJRQ, UGRF ? QGLEJC MNCLGLE RM RFC BGECQRGTC QWQRCK (RFC KMSRF) D?AGLE SN UGRF RCLR?AJCQ QSPPMSLBGLE GR. .CBSQ? DMPKQ ?PC KMRGJC, UGRF RFC KMSRF ?LB RCLR?AJCQ F?LEGLE BMUL DPMK ?L SK@PCJJ?-QF?NCB @CJJ.
#(!04%2 28 X ).6%24%"2!4%3
757
!@>LI< 28.6 #KFA>OF>KP E>SB QTL AFPQFK@Q ?LAV MI>KP, QEB JBARP> (>) >KA QEB MLIVM (?). !II @KFA>OF>KP E>SB QTL JBJ?O>KB I>VBOP, TFQE > GBIIV-IFHB JBPLDIB> ?BQTBBK QEBJ.
4MKC ALGB?PG?LQ ?PC NMJWKMPNFGA, RF?R GQ, RFCW F?TC RUM @MBW NJ?LQ BSPGLE RFCGP JGDC AWAJC. "L CV?KNJC GQ RFC AMJMLG?J FWBPMGB A?JJCB ?L O),30(. 5FC QCQQGJC NMJWN DMPK F?Q, GL D?AR, RUM RWNCQ MD NMJWNQ, QFMUL GL Figure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
!@>LI< 28.7 4EB PBPPFIB CLOJ LC O),30( .,50*<3(;, E>P QTL QVMBP LC MLIVMP: D>PQOLWLLFAP, TEF@E >OB >A>MQBA CLO @>MQROFKD MOBV, >KA DLKLWLLFAP, TEF@E ?RA QL MOLAR@B JBARP>B >PBUR>IIV.
758
#(!04%2 28 X ).6%24%"2!4%3
$JGAI FCPC RM DMJJMU RFC life cycle (http://openstaxcollege.org/l/obelia) MD RFC O),30(. "JJ ALGB?PG?LQ QFMU RFC NPCQCLAC MD RUM KCK@P?LC J?WCPQ GL RFC @MBW RF?R ?PC BCPGTCB DPMK RFC CLBMBCPK ?LB CARMBCPK MD RFC CK@PWM. 5FC MSRCP J?WCP (DPMK CARMBCPK) GQ A?JJCB RFC epidermis ?LB JGLCQ RFC MSRQGBC MD RFC ?LGK?J, UFCPC?Q RFC GLLCP J?WCP (DPMK CLBMBCPK) GQ A?JJCB RFC gastrodermis ?LB JGLCQ RFC BGECQRGTC A?TGRW. #CRUCCL RFCQC RUM KCK@P?LC J?WCPQ GQ ? LML-JGTGLE, HCJJW-JGIC mesoglea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extracellular digestion GL UFGAF RFC DMMB GQ R?ICL GLRM RFC E?QRPMT?QASJ?P A?TGRW, CLXWKCQ ?PC QCAPCRCB GLRM RFC A?TGRW, ?LB RFC ACJJQ JGLGLE RFC A?TGRW ?@QMP@ LSRPGCLRQ. 5FC gastrovascular cavity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s 8?P.
CC8JJ AEK?FQF8
5FC AJ?QQ "LRFMXM? GLAJSBCQ ?JJ ALGB?PG?LQ RF?R CVFG@GR ? NMJWN @MBW NJ?L MLJW; GL MRFCP UMPBQ, RFCPC GQ LM KCBSQ? QR?EC UGRFGL RFCGP JGDC AWAJC. &V?KNJCQ GLAJSBC QC? ?LCKMLCQ (Figure 28.8), QC? NCLQ, ?LB AMP?JQ, UGRF ?L CQRGK?RCB LSK@CP MD 6,100 BCQAPG@CB QNCAGCQ. 4C? ?LCKMLCQ ?PC SQS?JJW @PGEFRJW AMJMPCB ?LB A?L ?RR?GL ? QGXC MD 1.8 RM 10 AK GL BG?KCRCP. 5FCQC ?LGK?JQ ?PC SQS?JJW AWJGLBPGA?J GL QF?NC ?LB ?PC ?RR?AFCB RM ? QS@QRP?RC. " KMSRF MNCLGLE GQ QSPPMSLBCB @W RCLR?AJCQ @C?PGLE ALGBMAWRCQ.
#(!04%2 28 X ).6%24%"2!4%3
759
!@>LI< 28.8 4EB PB> >KBJLKB FP PELTK (>) MELQLDO>MEBA >KA (?) FK > AF>DO>J FIIRPQO>QFKD FQP JLOMELILDV. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V "$>K@FKD 7FQE 'ELPQP"/&IF@HO; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V ./!!)
5FC KMSRF MD ? QC? ?LCKMLC GQ QSPPMSLBCB @W RCLR?AJCQ RF?R @C?P ALGBMAWRCQ. 5FC QJGR-JGIC KMSRF MNCLGLE ?LB NF?PWLV ?PC JGLCB @W ? EPMMTC A?JJCB ? siphonophore. 5FC NF?PWLV GQ RFC KSQASJ?P N?PR MD RFC BGECQRGTC QWQRCK RF?R QCPTCQ RM GLECQR ?Q UCJJ ?Q CECQR DMMB, ?LB K?W CVRCLB DMP SN RM RUMRFGPBQ RFC JCLERF MD RFC @MBW @CDMPC MNCLGLE GLRM RFC E?QRPMT?QASJ?P A?TGRW. 5FGQ A?TGRW GQ BGTGBCB GLRM QCTCP?J AF?K@CPQ @W JMLEGRSBGL?J QCNR? A?JJCB KCQCLRCPGCQ. &?AF KCQCLRCPW AMLQGQRQ MD MLC CARMBCPK?J ?LB MLC CLBMBCPK?J ACJJ J?WCP UGRF RFC KCQMEJC? Q?LBUGAFCB GL @CRUCCL. .CQCLRCPGCQ BM LMR BGTGBC RFC E?QRPMT?QASJ?P A?TGRW AMKNJCRCJW, ?LB RFC QK?JJCP A?TGRGCQ AM?JCQAC ?R RFC NF?PWLEC?J MNCLGLE. 5FC ?B?NRGTC @CLCDGR MD RFC KCQCLRCPGCQ ?NNC?PQ RM @C ?L GLAPC?QC GL QSPD?AC ?PC? DMP ?@QMPNRGML MD LSRPGCLRQ ?LB E?Q CVAF?LEC. 4C? ?LCKMLCQ DCCB ML QK?JJ DGQF ?LB QFPGKN, SQS?JJW @W GKKM@GJGXGLE RFCGP NPCW SQGLE RFC ALGBMAWRCQ. 4MKC QC? ?LCKMLCQ CQR?@JGQF ? KSRS?JGQRGA PCJ?RGMLQFGN UGRF FCPKGR AP?@Q @W ?RR?AFGLE RM RFC AP?@sQ QFCJJ. *L RFGQ PCJ?RGMLQFGN, RFC ?LCKMLC ECRQ DMMB N?PRGAJCQ DPMK NPCW A?SEFR @W RFC AP?@, ?LB RFC AP?@ GQ NPMRCARCB DPMK RFC NPCB?RMPQ @W RFC QRGLEGLE ACJJQ MD RFC ?LCKMLC. "LCKMLC DGQF, MP AJMULDGQF, ?PC ?@JC RM JGTC GL RFC ?LCKMLC QGLAC RFCW ?PC GKKSLC RM RFC RMVGLQ AMLR?GLCB UGRFGL RFC LCK?RMAWQRQ. "LRFMXM?LQ PCK?GL NMJWNMGB RFPMSEFMSR RFCGP JGTCQ ?LB A?L PCNPMBSAC ?QCVS?JJW @W @SBBGLE MP DP?EKCLR?RGML, MP QCVS?JJW @W NPMBSAGLE E?KCRCQ. #MRF E?KCRCQ ?PC NPMBSACB @W RFC NMJWN, UFGAF A?L DSQC RM EGTC PGQC RM ? DPCC-QUGKKGLE NJ?LSJ? J?PT?. 5FC J?PT? QCRRJCQ ML ? QSGR?@JC QS@QRP?RSK ?LB BCTCJMNQ GLRM ? QCQQGJC NMJWN.
CC8JJ .:PG?FQF8
$J?QQ 4AWNFMXM? GLAJSBCQ ?JJ RFC HCJJGCQ ?LB GQ CVAJSQGTCJW ? K?PGLC AJ?QQ MD ?LGK?JQ UGRF ?@MSR 200 ILMUL QNCAGCQ. 5FC BCDGLGLE AF?P?ARCPGQRGA MD RFGQ AJ?QQ GQ RF?R RFC KCBSQ? GQ RFC NPMKGLCLR QR?EC GL RFC JGDC AWAJC, ?JRFMSEF RFCPC GQ ? NMJWN QR?EC NPCQCLR. .CK@CPQ MD RFGQ QNCAGCQ P?LEC DPMK 2 RM 40 AK GL JCLERF @SR RFC J?PECQR QAWNFMXM?L QNCAGCQ, C@(5,( *(7033(;(, A?L PC?AF ? QGXC MD 2 K ?APMQQ. 4AWNFMXM?LQ BGQNJ?W ? AF?P?ARCPGQRGA @CJJ-JGIC KMPNFMJMEW (Figure 28.9).
760
#(!04%2 28 X ).6%24%"2!4%3
!@>LI< 28.9 ! GBIIV FP PELTK (>) MELQLDO>MEBA >KA (?) FK > AF>DO>J FIIRPQO>QFKD FQP JLOMELILDV. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V "*FJD944"/&IF@HO; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
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igure 28.10) MD RFCQC ?LGK?JQ A?L @C BCQAPG@CB ?Q polymorphic, @CA?SQC RFCW CVFG@GR @MRF ? KCBSQ?J ?LB NMJWNMGB @MBW NJ?L ?R QMKC NMGLR GL RFCGP JGDC AWAJC.
#(!04%2 28 X ).6%24%"2!4%3
761
!@>LI< 28.10 4EB IFCB@V@IB LC > GBIIVCFPE FK@IRABP QTL PQ>DBP: QEB JBARP> PQ>DB >KA QEB MLIVM PQ>DB. 4EB MLIVM OBMOLAR@BP >PBUR>IIV ?V ?RAAFKD, >KA QEB JBARP> OBMOLAR@BP PBUR>IIV. (@OBAFQ "JBARP>": JLAFCF@>QFLK LC TLOH ?V &O>K@BP@L #OFMM>)
*BCLRGDW RFC JGDC AWAJC QR?ECQ MD HCJJGCQ SQGLE RFGQ video animation quiz (http://openstaxcollege.org/l/ amazing_jellies) DPMK RFC /CU &LEJ?LB "OS?PGSK.
CC8JJ CL9FQF8
5FGQ AJ?QQ GLAJSBCQ HCJJGCQ RF?R F?TC ? @MV-QF?NCB KCBSQ?, MP ? @CJJ RF?R GQ QOS?PC GL APMQQ-QCARGML; FCLAC, ?PC AMJJMOSG?JJW ILMUL ?Q t@MV HCJJWDGQF.u 5FCQC QNCAGCQ K?W ?AFGCTC QGXCQ MD 15p25 AK. $S@MXM?LQ BGQNJ?W MTCP?JJ KMPNFMJMEGA?J ?LB ?L?RMKGA?J AF?P?ARCPGQRGAQ RF?R ?PC QGKGJ?P RM RFMQC MD RFC QAWNFMXM?LQ. " NPMKGLCLR BGDDCPCLAC @CRUCCL RFC RUM AJ?QQCQ GQ RFC ?PP?LECKCLR MD RCLR?AJCQ. 5FGQ GQ RFC KMQR TCLMKMSQ EPMSN MD ?JJ RFC ALGB?PG?LQ (Figure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
762
#(!04%2 28 X ).6%24%"2!4%3
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
CC8JJ #P;IFQF8
)WBPMXM? GLAJSBCQ LC?PJW 3,200 QNCAGCQ; KMQR ?PC K?PGLC, ?JRFMSEF QMKC DPCQFU?RCP QNCAGCQ ?PC ILMUL (Figure 28.12). "LGK?JQ GL RFGQ AJ?QQ ?PC NMJWKMPNFQ, ?LB KMQR CVFG@GR @MRF NMJWNMGB ?LB KCBSQMGB DMPKQ GL RFCGP JGDCAWAJC, ?JRFMSEF RFGQ GQ T?PG?@JC. 5FC NMJWN DMPK GL RFCQC ?LGK?JQ MDRCL QFMUQ ? AWJGLBPGA?J KMPNFMJMEW UGRF ? ACLRP?J E?QRPMT?QASJ?P A?TGRW JGLCB @W RFC E?QRPMBCPKGQ. 5FC E?QRPMBCPKGQ ?LB CNGBCPKGQ F?TC ? QGKNJC J?WCP MD KCQMEJC? Q?LBUGAFCB @CRUCCL RFCK. " KMSRF MNCLGLE, QSPPMSLBCB @W RCLR?AJCQ, GQ NPCQCLR ?R RFC MP?J CLB MD RFC ?LGK?J. .?LW FWBPMXM?LQ DMPK AMJMLGCQ RF?R ?PC AMKNMQCB MD ? @P?LAFCB AMJMLW MD QNCAG?JGXCB NMJWNQ RF?R QF?PC ? E?QRPMT?QASJ?P A?TGRW, QSAF ?Q GL RFC AMJMLG?J FWBPMGB O),30(. $MJMLGCQ K?W ?JQM @C DPCCDJM?RGLE ?LB AMLR?GL KCBSQMGB ?LB NMJWNMGB GLBGTGBS?JQ GL RFC AMJMLW ?Q GL P/@:(30( (RFC 1MPRSESCQC .?L 0s 8?P) MP #,3,33( (#W-RFC-UGLB Q?GJMP). &TCL MRFCP QNCAGCQ ?PC QMJGR?PW NMJWNQ (H@+9() MP QMJGR?PW KCBSQ?C (G65065,4<:). 5FC RPSC AF?P?ARCPGQRGA QF?PCB @W ?JJ MD RFCQC BGTCPQC QNCAGCQ GQ RF?R RFCGP EML?BQ DMP QCVS?J PCNPMBSARGML ?PC BCPGTCB DPMK CNGBCPK?J RGQQSC, UFCPC?Q GL ?JJ MRFCP ALGB?PG?LQ RFCW ?PC BCPGTCB DPMK E?QRPMBCPK?J RGQQSC.
#(!04%2 28 X ).6%24%"2!4%3
763
!@>LI< 28.12 (>) O),30(, (?) P/@:(30( 7/@:(30:, HKLTK >P QEB 0LOQRDRBPB ->K /v 7>O, (@) #,3,33( )(,, >KA (A) H@+9( E>SB AFCCBOBKQ ?LAV PE>MBP ?RQ >II ?BILKD QL QEB C>JFIV (VAOLWL>. (@OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V ./!!; P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
28.3 R .LG<IG?PCLD 'FG?FKIF:?FQF8

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" SLGOSC AF?P?ARCPGQRGA MD RFGQ NFWJSK GQ RFC NPCQCLAC MD ? NPM@MQAGQ CLAJMQCB GL ? rhynchocoel. 5FC NPM@MQAGQ QCPTCQ RM A?NRSPC DMMB ?LB K?W @C MPL?KCLRCB UGRF @?P@Q GL QMKC QNCAGCQ. 5FC PFWLAFMAMCJ GQ ? DJSGB-DGJJCB A?TGRW RF?R CVRCLBQ DPMK RFC FC?B RM LC?PJW RUM-RFGPBQ MD RFC JCLERF MD RFC ESR GL RFCQC ?LGK?JQ (Figure 28.19). 5FC NPM@MQAGQ K?W @C CVRCLBCB MP PCRP?ARCB @W RFC PCRP?ARMP KSQAJC ?RR?AFCB RM RFC U?JJ MD RFC PFWLAFMAMCJ.
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8?RAF RFGQ video (http://openstaxcollege.org/l/nemertean) RM QCC ? LCKCPRC?L ?RR?AI ? NMJWAF?CRC UGRF GRQ NPM@MQAGQ. D/-+89/;+ !>89+2 5FC LCKCPRGLG QFMU ? TCPW UCJJ-BCTCJMNCB BGECQRGTC QWQRCK. " KMSRF MNCLGLE RF?R GQ TCLRP?J RM RFC PFWLAFMAMCJ JC?BQ GLRM RFC DMPCESR, DMJJMUCB @W RFC GLRCQRGLC. 5FC GLRCQRGLC GQ NPCQCLR GL RFC DMPK MD BGTCPRGASJ?P NMSAFCQ ?LB CLBQ GL ? PCARSK RF?R MNCLQ TG? ?L ?LSQ. (ML?BQ ?PC GLRCPQNCPQCB UGRF RFC GLRCQRGL?J BGTCPRGASJ?P NMSAFCQ ?LB MNCL MSRU?PBQ TG? ECLGR?J NMPCQ. " AGPASJ?RMPW QWQRCK AMLQGQRQ MD ? AJMQCB JMMN MD ? N?GP MD J?RCP?J @JMMB TCQQCJQ. 5FC AGPASJ?RMPW QWQRCK GQ BCPGTCB DPMK RFC AMCJMKGA A?TGRW MD RFC CK@PWM. 4MKC ?LGK?JQ K?W ?JQM F?TC APMQQ-AMLLCARGLE TCQQCJQ GL ?BBGRGML RM J?RCP?J MLCQ. "JRFMSEF RFCQC ?PC A?JJCB @JMMB TCQQCJQ, QGLAC RFCW ?PC MD AMCJMKGA MPGEGL, RFC AGPASJ?RMPW DJSGB GQ AMJMPJCQQ. 4MKC QNCAGCQ @C?P FCKMEJM@GL ?Q UCJJ ?Q MRFCP WCJJMU MP EPCCL NGEKCLRQ. 5FC @JMMB TCQQCJQ ?PC AMLLCARCB RM RFC PFWLAFMAMCJ. 5FC DJMU MD DJSGB GL RFCQC TCQQCJQ GQ D?AGJGR?RCB @W RFC AMLRP?ARGML MD KSQAJCQ GL RFC @MBW U?JJ. " N?GP MD NPMRMLCNFPGBG?, MP NPGKGRGTC IGBLCWQ, GQ NPCQCLR GL RFCQC ?LGK?JQ RM D?AGJGR?RC MQKMPCESJ?RGML. (?QCMSQ CVAF?LEC MAASPQ RFPMSEF RFC QIGL GL RFC LCKCPRGLG.
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N+7;4:8 !>89+2 /CKCPRGLG F?TC ? E?LEJGML MP t@P?GLu QGRS?RCB ?R RFC ?LRCPGMP CLB @CRUCCL RFC KMSRF ?LB RFC DMPCESR, QSPPMSLBGLE RFC BGECQRGTC QWQRCK ?Q UCJJ ?Q RFC PFWLAFMAMCJ. " PGLE MD DMSP LCPTC K?QQCQ A?JJCB tE?LEJG?u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planuliform J?PT?. *L QMKC LCKCPRGLC QNCAGCQ, ? pilidium J?PT? K?W BCTCJMN GLQGBC RFC WMSLE UMPK, DPMK ? QCPGCQ MD GK?EGL?J BGQAQ. 5FGQ J?PT?J DMPK, AF?P?ARCPGQRGA?JJW QF?NCB JGIC ? BCCPQR?JICP A?N, BCTMSPQ RGQQSCQ DPMK RFC WMSLE UMPK DMP QSPTGT?J @CDMPC KCR?KMPNFMQGLE GLRM RFC ?BSJR-JGIC KMPNFMJMEW.
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1FWJSK Mollusca GQ RFC NPCBMKGL?LR NFWJSK GL K?PGLC CLTGPMLKCLRQ. *R GQ CQRGK?RCB RF?R 23 NCPACLR MD ?JJ ILMUL K?PGLC QNCAGCQ ?PC KMJJSQIQ; RFCPC ?PC MTCP 75,000 BCQAPG@CB QNCAGCQ, K?IGLE RFCK RFC QCAMLB KMQR BGTCPQC NFWJSK MD ?LGK?JQ. 5FC L?KC tKMJJSQA?u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igure 28.20).
!@>LI< 28.20 4EBOB >OB J>KV PMB@FBP >KA S>OF>QFLKP LC JLIIRPHP; QEFP FIIRPQO>QFLK PELTP QEB >K>QLJV LC >K >NR>QF@ D>PQOLMLA.
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$J?QQ #GT?JTG? (tRUM QFCJJQu) GLAJSBCQ AJ?KQ, MWQRCPQ, KSQQCJQ, QA?JJMNQ, ?LB ECMBSAIQ. .CK@CPQ MD RFGQ AJ?QQ ?PC DMSLB GL K?PGLC ?Q UCJJ ?Q DPCQFU?RCP F?@GR?RQ. "Q RFC L?KC QSEECQRQ, @GT?JTCQ ?PC CLAJMQCB GL ? N?GP MD QFCJJQ (T?JTCQ ?PC AMKKMLJW A?JJCB tQFCJJQu) RF?R ?PC FGLECB ?R RFC BMPQ?J CLB @W QFCJJ JGE?KCLRQ ?Q UCJJ ?Q QFCJJ RCCRF (Figure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#(!04%2 28 X ).6%24%"2!4%3
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$J?QQ $CNF?JMNMB? (tFC?B DMMRu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tHCRu NPMNSJQGML. " N?GP MD LCNFPGBG? GQ NPCQCLR UGRFGL RFC K?LRJC A?TGRW. 4CVS?J BGKMPNFGQK GQ QCCL GL RFGQ AJ?QQ MD ?LGK?JQ. .CK@CPQ MD ? QNCAGCQ K?RC, ?LB RFC DCK?JC RFCL J?WQ RFC CEEQ GL ? QCAJSBCB ?LB NPMRCARCB LGAFC. 'CK?JCQ MD QMKC QNCAGCQ A?PC DMP RFC CEEQ DMP ?L CVRCLBCB NCPGMB MD RGKC ?LB K?W CLB SN BWGLE BSPGLE RF?R RGKC NCPGMB. $CNF?JMNMBQ QSAF ?Q QOSGBQ ?LB MARMNG ?JQM NPMBSAC QCNG? MP ? B?PI GLI, UFGAF GQ QOSGPRCB SNML ? NPCB?RMP RM ?QQGQR GL ? OSGAI ECR?U?W. 3CNPMBSARGML GL ACNF?JMNMBQ GQ BGDDCPCLR DPMK MRFCP KMJJSQIQ GL RF?R RFC CEE F?RAFCQ RM NPMBSAC ? HSTCLGJC ?BSJR UGRFMSR SLBCPEMGLE RFC RPMAFMNFMPC ?LB TCJGECP J?PT?J QR?ECQ. *L RFC QFCJJ-@C?PGLE N(<;03<: QNN., RFC QNGP?J QFCJJ GQ KSJRG-AF?K@CPCB. 5FCQC AF?K@CPQ ?PC DGJJCB UGRF E?Q MP U?RCP RM PCESJ?RC @SMW?LAW. 5FC QFCJJ QRPSARSPC GL QOSGBQ ?LB ASRRJCDGQF GQ PCBSACB ?LB GQ NPCQCLR GLRCPL?JJW GL RFC DMPK MD ? QOSGB NCL ?LB ASRRJCDGQF @MLC, PCQNCARGTCJW. &V?KNJCQ ?PC QFMUL GL Figure 28.26.
!@>LI< 28.26 4EB (>) K>RQFIRP, (?) DF>KQ @RQQIBCFPE, (@) OBBC PNRFA, >KA (A) ?IRB-OFKD L@QLMRP >OB >II JBJ?BOP LC QEB @I>PP #BME>ILMLA>. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V *. ">B@HBO; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V !AOF>K -LEBA>KL; @OBAFQ @: JLAFCF@>QFLK LC TLOH ?V 3FIHB ">OLK; @OBAFQ A: JLAFCF@>QFLK LC TLOH ?V !KDBII 7FIIF>JP)
.CK@CPQ MD AJ?QQ 4A?NFMNMB? (t@M?R DCCRu) ?PC ILMUL AMJJMOSG?JJW ?Q tRSQI QFCJJQu MP tRMMRF QFCJJQ,u ?Q CTGBCLR UFCL CV?KGLGLE D,5;(30<4, MLC MD RFC DCU PCK?GLGLE QA?NFMNMB ECLCP? (Figure 28.27). 4A?NFMNMBQ ?PC SQS?JJW @SPGCB GL Q?LB UGRF RFC ?LRCPGMP MNCLGLE CVNMQCB RM U?RCP. 5FCQC ?LGK?JQ @C?P ? QGLEJC AMLGA?J QFCJJ, UFGAF F?Q @MRF CLBQ MNCL. 5FC FC?B GQ PSBGKCLR?PW ?LB NPMRPSBCQ MSR MD RFC NMQRCPGMP CLB MD RFC QFCJJ. 5FCQC ?LGK?JQ BM LMR NMQQCQQ CWCQ, @SR RFCW F?TC ? P?BSJ?, ?Q UCJJ ?Q ? DMMR KMBGDGCB GLRM RCLR?AJCQ UGRF ? @SJ@MSQ CLB, ILMUL ?Q captaculae. $?NR?ASJ?C QCPTC RM A?RAF ?LB K?LGNSJ?RC NPCW. $RCLGBG? ?PC ?@QCLR GL RFCQC ?LGK?JQ.
776
#(!04%2 28 X ).6%24%"2!4%3
!@>LI< 28.27 A5;(30: =<3.(90: PELTP QEB @I>PPF@ $BKQ>IFFA>B PE>MB QE>Q DFSBP QEBPB >KFJ>IP QEBFO @LJJLK K>JB LC "QRPH PEBII." (@OBAFQ: 'BLODBP *>KPLLKB)
+?PCLD AEE<C@;8
1FWJSK Annelida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tQCEKCLRCB UMPKQu BSC RM RFCGP ICW AF?P?ARCPGQRGA MD metamerism, MP RPSC QCEKCLR?RGML. M475.414-> "LLCJGBQ BGQNJ?W @GJ?RCP?J QWKKCRPW ?LB ?PC UMPK-JGIC GL MTCP?JJ KMPNFMJMEW. "LLCJGBQ F?TC ? QCEKCLRCB @MBW NJ?L UFCPCGL RFC GLRCPL?J ?LB CVRCPL?J KMPNFMJMEGA?J DC?RSPCQ ?PC PCNC?RCB GL C?AF @MBW QCEKCLR. .CR?KCPGQK ?JJMUQ ?LGK?JQ RM @CAMKC @GEECP @W ?BBGLE tAMKN?PRKCLRQu UFGJC K?IGLE RFCGP KMTCKCLR KMPC CDDGAGCLR. 5FGQ KCR?KCPGQK GQ RFMSEFR RM ?PGQC DPMK GBCLRGA?J RCJM@J?QR ACJJQ GL RFC CK@PWMLGA QR?EC, UFGAF EGTC PGQC RM GBCLRGA?J KCQMBCPK?J QRPSARSPCQ. 5FC MTCP?JJ @MBW A?L @C BGTGBCB GLRM FC?B, @MBW, ?LB NWEGBGSK (MP R?GJ). 5FC clitellum GQ ? PCNPMBSARGTC QRPSARSPC RF?R ECLCP?RCQ KSASQ RF?R ?GBQ GL QNCPK RP?LQDCP ?LB EGTCQ PGQC RM ? AMAMML UGRFGL UFGAF DCPRGJGX?RGML MAASPQ; GR ?NNC?PQ ?Q ? DSQCB @?LB GL RFC ?LRCPGMP RFGPB MD RFC ?LGK?J (Figure 28.28).
!@>LI< 28.28 4EB @IFQBIIRJ, PBBK EBOB >P > MOLQORAFKD PBDJBKQ TFQE AFCCBOBKQ @LILO>QFLK QE>K QEB OBPQ LC QEB ?LAV, FP > PQOR@QROB QE>Q >FAP FK >KKBIFA OBMOLAR@QFLK. (@OBAFQ: 2L? (FIIB)
A3'942> 5FC CNGBCPKGQ GQ NPMRCARCB @W ?L ?ACJJSJ?P, CVRCPL?J ASRGAJC, @SR RFGQ GQ KSAF RFGLLCP RF?L RFC ASRGAJC DMSLB GL RFC CABWQMXM?LQ ?LB BMCQ LMR PCOSGPC NCPGMBGA QFCBBGLE DMP EPMURF. $GPASJ?P ?Q UCJJ ?Q JMLEGRSBGL?J KSQAJCQ ?PC JMA?RCB GLRCPGMP RM RFC CNGBCPKGQ. $FGRGLMSQ F?GPJGIC CVRCLQGMLQ, ?LAFMPCB GL RFC CNGBCPKGQ ?LB NPMHCARGLE DPMK RFC ASRGAJC, A?JJCB setae/chaetae ?PC NPCQCLR GL CTCPW QCEKCLR. "LLCJGBQ QFMU
#(!04%2 28 X ).6%24%"2!4%3
777
RFC NPCQCLAC MD ? RPSC AMCJMK, BCPGTCB DPMK CK@PWMLGA KCQMBCPK ?LB NPMRMQRMKW. )CLAC, RFCW ?PC RFC KMQR ?BT?LACB UMPKQ. " UCJJ-BCTCJMNCB ?LB AMKNJCRC BGECQRGTC QWQRCK GQ NPCQCLR GL C?PRFUMPKQ (MJGEMAF?CRCQ) UGRF ? KMSRF, KSQASJ?P NF?PWLV, CQMNF?ESQ, APMN, ?LB EGXX?PB @CGLE NPCQCLR. 5FC EGXX?PB JC?BQ RM RFC GLRCQRGLC ?LB CLBQ GL ?L ?L?J MNCLGLE. " APMQQ-QCARGML?J TGCU MD ? @MBW QCEKCLR MD ?L C?PRFUMPK (? RCPPCQRPG?J RWNC MD ?LLCJGB) GQ QFMUL GL Figure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tIGBLCWu RF?R AMLQGQRQ MD ? AMLTMJSRCB RS@SJC ?LB ?L MNCL, AGJG?RCB DSLLCJ) RF?R GQ NPCQCLR GL CTCPW QCEKCLR RMU?PBQ RFC TCLRP?J QGBC. "LLCJGBQ QFMU UCJJ-BCTCJMNCB LCPTMSQ QWQRCKQ UGRF ? LCPTC PGLE MD DSQCB E?LEJG? NPCQCLR ?PMSLB RFC NF?PWLV. 5FC LCPTC AMPB GQ TCLRP?J GL NMQGRGML ?LB @C?PQ CLJ?PECB LMBCQ MP E?LEJG? GL C?AF QCEKCLR.
!@>LI< 28.29 4EFP P@EBJ>QF@ AO>TFKD PELTP QEB ?>PF@ >K>QLJV LC >KKBIFAP FK > @OLPP-PB@QFLK>I SFBT.
"LLCJGBQ K?W @C CGRFCP KMLMCAGMSQ UGRF NCPK?LCLR EML?BQ (?Q GL C?PRFUMPKQ ?LB JCCAFCQ) MP BGMCAGMSQ UGRF RCKNMP?PW MP QC?QML?J EML?BQ RF?R BCTCJMN (?Q GL NMJWAF?CRCQ). )MUCTCP, APMQQDCPRGJGX?RGML GQ NPCDCPPCB GL FCPK?NFPMBGRGA ?LGK?JQ. 5FCQC ?LGK?JQ K?W ?JQM QFMU QGKSJR?LCMSQ FCPK?NFPMBGRGQK ?LB N?PRGAGN?RC GL QGKSJR?LCMSQ QNCPK CVAF?LEC UFCL RFCW ?PC ?JGELCB DMP AMNSJ?RGML.
5FGQ AMK@GL?RGML video and animation (http://shapeoflife.org/video/animation/annelidanimation-body-plan) NPMTGBCQ ? AJMQC-SN JMMI ?R ?LLCJGB ?L?RMKW. C1'88/,/)'9/43 4, P.>1:2 A33+1/*' 1FWJSK "LLCJGB? AMLR?GLQ RFC AJ?QQ 1MJWAF?CR? (RFC NMJWAF?CRCQ) ?LB RFC AJ?QQ 0JGEMAF?CR? (RFC C?PRFUMPKQ, JCCAFCQ ?LB RFCGP PCJ?RGTCQ). &?PRFUMPKQ ?PC RFC KMQR ?@SLB?LR KCK@CPQ MD RFC AJ?QQ 0JGEMAF?CR?, BGQRGLESGQFCB @W RFC NPCQCLAC MD RFC AJGRCJJSK ?Q UCJJ ?Q DCU, PCBSACB AF?CR?C (tMJGEM- = tDCUu; -AF?CR?C = tF?GPQu). 5FC LSK@CP ?LB QGXC MD AF?CR?C ?PC EPC?RJW BGKGLGQFCB GL 0JGEMAF?CR? AMKN?PCB RM RFC NMJWAF?CRCQ (763@=K?LW, */(,;(, = F?GPQ). 5FC K?LW AFCR?C MD NMJWAF?CRCQ ?PC ?JQM ?PP?LECB UGRFGL DJCQFW, DJ?R, N?GPCB ?NNCLB?ECQ RF?R NPMRPSBC DPMK C?AF QCEKCLR A?JJCB parapodia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
778
#(!04%2 28 X ).6%24%"2!4%3
JCCAFCQ RM CJMLE?RC UFCL RFCW GLECQR AMNGMSQ OS?LRGRGCQ MD @JMMB DPMK FMQR TCPRC@P?RCQ. 5FC QS@AJ?QQ #P?AFGM@BCJJ? GLAJSBCQ QNCAGCQ JGIC B9(5*/06)+,33( )(3*(50*( :2,;0 ?LB B9(5*/06)+,33( (:;(*0, UMPKQ RF?R QFMU QGKGJ?PGRW UGRF JCCAFCQ ?Q UCJJ ?Q MJGEMAF?CRCQ.
!@>LI< 28.30 4EB (>) B>OQETLOJ, (?) IBB@E, >KA (@) CB>QEBOARPQBO >OB >II >KKBIFAP. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V 3. 3EBMEBOA; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V x3>O>E '...y/&IF@HO; @OBAFQ @: JLAFCF@>QFLK LC TLOH ?V #EOFP 'LQP@E>IH, ./!!)
28.4 R .LG<IG?PCLD :;PJFQF8

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC QRPSARSP?J MPE?LGX?RGML MD LCK?RMBCQ z 6LBCPQR?LB RFC GKNMPR?LAC MD $?CLMPF?@BGRGQ CJCE?LQ GL PCQC?PAF z $MKN?PC RFC GLRCPL?J QWQRCKQ ?LB ?NNCLB?EC QNCAG?JGX?RGMLQ MD NFWJSK "PRFPMNMB? z %GQASQQ RFC CLTGPMLKCLR?J GKNMPR?LAC MD ?PRFPMNMBQ z %GQASQQ RFC PC?QMLQ DMP ?PRFPMNMB QSAACQQ ?LB ?@SLB?LAC
.LG<IG?PCLD :;PJFQF8
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
+?PCLD )<D8KF;8
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ematoda GLAJSBCQ KMPC RF?L 28,000 QNCAGCQ UGRF ?L CQRGK?RCB 16,000 @CGLE N?P?QGRGA GL L?RSPC. 5FC L?KC /CK?RMB? GQ BCPGTCB DPMK RFC (PCCI UMPB t/CKMQ,u UFGAF KC?LQ tRFPC?Bu ?LB GLAJSBCQ PMSLBUMPKQ. /CK?RMBCQ ?PC NPCQCLR GL ?JJ F?@GR?RQ UGRF ? J?PEC LSK@CP MD GLBGTGBS?JQ MD C?AF QNCAGCQ NPCQCLR GL C?AF. 5FC DPCC-JGTGLE LCK?RMBC, C(,569/()+0;0: ,3,.(5: F?Q @CCL CVRCLQGTCJW SQCB ?Q ? KMBCJ QWQRCK GL J?@MP?RMPGCQ ?JJ MTCP RFC UMPJB.
1. 4RMJJ, /. 3., t5FGQ UMPKW UMPJB. 1947,u J6<95(3 6- P(9(:0;636.@ 85(3) (1999): 392-396.
#(!04%2 28 X ).6%24%"2!4%3
779
M475.414-> *L AMLRP?QR UGRF ALGB?PG?LQ, LCK?RMBCQ QFMU ? RS@SJ?P KMPNFMJMEW ?LB AGPASJ?P APMQQ-QCARGML. 5FCQC ?LGK?JQ ?PC NQCSBMAMCJMK?RCQ ?LB QFMU RFC NPCQCLAC MD ? AMKNJCRC BGECQRGTC QWQRCK UGRF ? BGQRGLAR KMSRF ?LB ?LSQ. 5FGQ GQ GL AMLRP?QR UGRF RFC ALGB?PG?LQ, UFCPC MLJW MLC MNCLGLE GQ NPCQCLR (?L GLAMKNJCRC BGECQRGTC QWQRCK). 5FC ASRGAJC MD /CK?RMBCQ GQ PGAF GL AMJJ?ECL ?LB ? A?P@MFWBP?RC-NPMRCGL NMJWKCP A?JJCB AFGRGL, ?LB DMPKQ ?L CVRCPL?J tQICJCRMLu MSRQGBC RFC CNGBCPKGQ. 5FC ASRGAJC ?JQM JGLCQ K?LW MD RFC MPE?LQ GLRCPL?JJW, GLAJSBGLE RFC NF?PWLV ?LB PCARSK. 5FC CNGBCPKGQ A?L @C CGRFCP ? QGLEJC J?WCP MD ACJJQ MP ? QWLAWRGSK, UFGAF GQ ? KSJRGLSAJC?RCB ACJJ DMPKCB DPMK RFC DSQGML MD SLGLSAJC?RCB ACJJQ. 5FC MTCP?JJ KMPNFMJMEW MD RFCQC UMPKQ GQ AWJGLBPGA?J, ?Q QCCL GL Figure 28.31. 5FC FC?B GQ P?BG?JJW QWKKCRPGA?J. " KMSRF MNCLGLE GQ NPCQCLR ?R RFC ?LRCPGMP CLB UGRF RFPCC MP QGV JGNQ ?Q UCJJ ?Q RCCRF GL QMKC QNCAGCQ GL RFC DMPK MD ASRGAJC CVRCLQGMLQ. 4MKC LCK?RMBCQ K?W NPCQCLR MRFCP CVRCPL?J KMBGDGA?RGMLQ JGIC PGLEQ, FC?B QFGCJBQ, MP U?PRQ. 3GLEQ, FMUCTCP, BM LMR PCDJCAR RPSC GLRCPL?J @MBW QCEKCLR?RGML. 5FC KMSRF JC?BQ RM ? KSQASJ?P NF?PWLV ?LB GLRCQRGLC, UFGAF JC?BQ RM ? PCARSK ?LB ?L?J MNCLGLE ?R RFC NMQRCPGMP CLB. 5FC KSQAJCQ MD LCK?RMBCQ BGDDCP DPMK RFMQC MD KMQR ?LGK?JQ: 5FCW F?TC ? JMLEGRSBGL?J J?WCP MLJW, UFGAF ?AAMSLRQ DMP RFC UFGN-JGIC KMRGML MD RFCGP KMTCKCLR.
!@>LI< 28.31 3@>KKFKD BIB@QOLK JF@OLDO>ME PELTP (>) QEB PLV?B>K @VPQ KBJ>QLAB (H,;,96+,9( .3@*05,:) >KA > KBJ>QLAB BDD. (?) ! P@EBJ>QF@ OBMOBPBKQ>QFLK PELTP QEB >K>QLJV LC > QVMF@>I KBJ>QLAB. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V 53$! !23; P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
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t@P?GLu MD RFC UMPK (UFGAF R?IC RFC DMPK MD ? PGLE ?PMSLB RFC NF?PWLV) ?Q UCJJ ?Q ?R RFC NMQRCPGMP CLB RM DMPK RFC R?GJ
780
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E?LEJG?. *L C. ,3,.(5:, RFC LCPTMSQ QWQRCK ?AAMSLRQ DMP LC?PJW MLC-RFGPB MD RFC RMR?J LSK@CP MD ACJJQ GL RFC ?LGK?J. +574*:)9/43 /CK?RMBCQ CKNJMW ? T?PGCRW MD PCNPMBSARGTC QRP?RCEGCQ RF?R P?LEC DPMK KMLMCAGMSQ RM BGMCAGMSQ RM N?PRFCLMECLGA, BCNCLBGLE SNML RFC QNCAGCQ SLBCP AMLQGBCP?RGML. C. ,3,.(5: GQ ? KMLMCAGMSQ QNCAGCQ ?LB QFMUQ BCTCJMNKCLR MD MT? AMLR?GLCB GL ? SRCPSQ ?Q UCJJ ?Q QNCPK AMLR?GLCB GL RFC QNCPK?RFCA?. 5FC SRCPSQ F?Q ?L CVRCPL?J MNCLGLE ILMUL ?Q RFC TSJT?. 5FC DCK?JC ECLGR?J NMPC GQ LC?P RFC KGBBJC MD RFC @MBW, UFCPC?Q RFC K?JCsQ GQ ?R RFC RGN. 4NCAG?JGXCB QRPSARSPCQ ?R RFC R?GJ MD RFC K?JC ICCN FGK GL NJ?AC UFGJC FC BCNMQGRQ QNCPK UGRF AMNSJ?RMPW QNGASJCQ. 'CPRGJGX?RGML GQ GLRCPL?J, ?LB CK@PWMLGA BCTCJMNKCLR QR?PRQ TCPW QMML ?DRCP DCPRGJGX?RGML. 5FC CK@PWM GQ PCJC?QCB DPMK RFC TSJT? BSPGLE RFC E?QRPSJ?RGML QR?EC. 5FC CK@PWMLGA BCTCJMNKCLR QR?EC J?QRQ DMP 14 FMSPQ; BCTCJMNKCLR RFCL AMLRGLSCQ RFPMSEF DMSP QSAACQQGTC J?PT?J QR?ECQ UGRF CABWQGQ @CRUCCL C?AF QR?ECq-1, -2, -3, ?LB -4qSJRGK?RCJW JC?BGLE RM RFC BCTCJMNKCLR MD ? WMSLE K?JC MP DCK?JC ?BSJR UMPK. "BTCPQC CLTGPMLKCLR?J AMLBGRGMLQ JGIC MTCPAPMUBGLE ?LB J?AI MD DMMB A?L PCQSJR GL RFC DMPK?RGML MD ?L GLRCPKCBG?RC J?PT?J QR?EC ILMUL ?Q RFC B?SCP J?PT?.
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xBDD QL >ARIQ QL A>RDEQBO BDD;y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xPKF@H@IRKHy/&IF@HO: @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V .)$$+, .)(; P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
782
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" LSK@CP MD AMKKML N?P?QGRGA LCK?RMBCQ QCPTC ?Q NPGKC CV?KNJCQ MD N?P?QGRGQK. 5FCQC ?LGK?JQ CVFG@GR AMKNJCV JGDCAWAJCQ RF?R GLTMJTC KSJRGNJC FMQRQ, ?LB RFCW A?L F?TC QGELGDGA?LR KCBGA?J ?LB TCRCPGL?PW GKN?ARQ. )SK?LQ K?W @CAMKC GLDCARCB @W D9(*<5*<3<: 4,+05,5:0:, ILMUL ?Q ESGLC? UMPKQ, UFCL RFCW BPGLI SLDGJRCPCB U?RCP AMLR?GLGLE AMNCNMBQ (Figure 28.33). )MMIUMPKQ, QSAF ?Q A5*@*36:;64( ?LB N,*(;69, GLDCQR RFC GLRCQRGLCQ ?LB DCCB ML RFC @JMMB MD K?KK?JQ, CQNCAG?JJW GL BMEQ, A?RQ, ?LB FSK?LQ. 5PGAFGL? UMPKQ (!90*/05,33() ?PC RFC A?SQ?J MPE?LGQK MD RPGAFGLMQGQ GL FSK?LQ, MDRCL PCQSJRGLE DPMK RFC AMLQSKNRGML MD SLBCPAMMICB NMPI; !90*/05,33( A?L GLDCAR MRFCP K?KK?JG?L FMQRQ ?Q UCJJ. A:*(90:, ? J?PEC GLRCQRGL?J PMSLBUMPK, QRC?JQ LSRPGRGML DPMK GRQ FSK?L FMQR ?LB K?W APC?RC NFWQGA?J @JMAI?EC MD RFC GLRCQRGLCQ. 5FC DGJ?PG?J UMPKQ, QSAF ?Q D096-03(90( ?LB $<*/,9,90(, ?PC AMKKMLJW TCARMPCB @W KMQOSGRMCQ, UFGAF N?QQ RFC GLDCARGTC ?ECLRQ ?KMLE K?KK?JQ RFPMSEF RFCGP @JMMB-QSAIGLE ?ARGTGRW. D096-03(90( 0440;0:, ? @JMMB-GLDCARGTC N?P?QGRC, GQ RFC LMRMPGMSQ BME FC?PRUMPK QNCAGCQ. $<*/,9,90( )(5*96-;0 GLDCARQ RFC JWKNF LMBCQ MD FSK?LQ, PCQSJRGLE GL RFC LML-JCRF?J @SR BCDMPKGLE AMLBGRGML A?JJCB CJCNF?LRG?QGQ, GL UFGAF N?PRQ MD RFC @MBW @CAMKC QUCJJCB RM EGE?LRGA NPMNMPRGMLQ BSC RM M@QRPSARGML MD JWKNF?RGA BP?GL?EC ?LB GLDJ?KK?RGML MD JWKNF?RGA RGQQSCQ.
!@>LI< 28.33 4EB DRFKB> TLOJ D9(*<5*<3<: 4,+05,5:0: FKCB@QP >?LRQ 3.5 JFIIFLK MBLMIB >KKR>IIV, JLPQIV FK !COF@>. (>) (BOB, QEB TLOJ FP TO>MMBA >OLRKA > PQF@H PL FQ @>K ?B BUQO>@QBA. (?) )KCB@QFLK L@@ROP TEBK MBLMIB @LKPRJB T>QBO @LKQ>JFK>QBA ?V FKCB@QBA @LMBMLAP, ?RQ QEFP @>K B>PFIV ?B MOBSBKQBA ?V PFJMIB CFIQO>QFLK PVPQBJP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V #$#)
#(!04%2 28 X ).6%24%"2!4%3
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+?PCLD AIK?IFGF;8
5FC L?KC t?PRFPMNMB?u KC?LQ tHMGLRCB JCEQu (GL RFC (PCCI, t?PRFPMQu KC?LQ tHMGLRu ?LB tNMBMQu KC?LQ tJCEu); GR ?NRJW BCQAPG@CQ RFC CLMPKMSQ LSK@CP MD GLTCPRC@P?RCQ GLAJSBCB GL RFGQ NFWJSK. Arthropoda BMKGL?RC RFC ?LGK?J IGLEBMK UGRF ?L CQRGK?RCB 85 NCPACLR MD ILMUL QNCAGCQ GLAJSBCB GL RFGQ NFWJSK ?LB K?LW ?PRFPMNMBQ WCR SLBMASKCLRCB. 5FC NPGLAGN?J AF?P?ARCPGQRGAQ MD ?JJ RFC ?LGK?JQ GL RFGQ NFWJSK ?PC DSLARGML?J QCEKCLR?RGML MD RFC @MBW ?LB NPCQCLAC MD HMGLRCB ?NNCLB?ECQ. "PRFPMNMBQ ?JQM QFMU RFC NPCQCLAC MD ?L CVMQICJCRML K?BC NPGLAGN?JJW MD AFGRGL, UFGAF GQ ? U?RCPNPMMD, RMSEF NMJWQ?AAF?PGBC. 1FWJSK "PRFPMNMB? GQ RFC J?PECQR NFWJSK GL RFC ?LGK?J UMPJB, ?LB GLQCARQ DMPK RFC QGLEJC J?PECQR AJ?QQ UGRFGL RFGQ NFWJSK. "PRFPMNMBQ ?PC CSAMCJMK?RC, NPMRMQRMKGA MPE?LGQKQ. 1FWJSK "PRFPMNMB? GLAJSBCQ ?LGK?JQ RF?R F?TC @CCL QSAACQQDSJ GL AMJMLGXGLE RCPPCQRPG?J, ?OS?RGA, ?LB ?CPG?J F?@GR?RQ. 5FGQ NFWJSK GQ DSPRFCP AJ?QQGDGCB GLRM DGTC QS@NFWJ?: 5PGJM@GRMKMPNF? (RPGJM@GRCQ, ?JJ CVRGLAR), )CV?NMB? (GLQCARQ ?LB PCJ?RGTCQ), .WPG?NMB? (KGJJGNCBCQ, ACLRGNCBCQ, ?LB PCJ?RGTCQ), $PSQR?AC?LQ (AP?@Q, JM@QRCPQ, AP?WDGQF, GQMNMBQ, @?PL?AJCQ, ?LB QMKC XMMNJ?LIRML), ?LB $FCJGACP?R? (FMPQCQFMC AP?@Q, ?P?AFLGBQ, QAMPNGMLQ, ?LB B?BBW JMLEJCEQ). 5PGJM@GRCQ ?PC ?L CVRGLAR EPMSN MD ?PRFPMNMBQ DMSLB AFGCDJW GL RFC NPC-$?K@PG?L &P? RF?R ?PC NPM@?@JW KMQR AJMQCJW PCJ?RCB RM RFC $FCJGACP?R?. 5FCQC ?PC GBCLRGDGCB @?QCB ML DMQQGJ PCAMPBQ (Figure 28.34).
!@>LI< 28.34 4OFIL?FQBP, IFHB QEB LKB FK QEFP CLPPFI, >OB >K BUQFK@Q DOLRM LC >OQEOLMLAP. (@OBAFQ: +BSFK 7>IPE)
M475.414-> " SLGOSC DC?RSPC MD ?LGK?JQ GL RFC ?PRFPMNMB NFWJSK GQ RFC NPCQCLAC MD ? QCEKCLRCB @MBW ?LB DSQGML MD QCRQ MD QCEKCLRQ RF?R EGTC PGQC RM DSLARGML?J @MBW PCEGMLQ A?JJCB R?EK?. 5?EK? K?W @C GL RFC DMPK MD ? FC?B, RFMP?V, ?LB ?@BMKCL, MP ? ACNF?JMRFMP?V ?LB ?@BMKCL, MP ? FC?B ?LB RPSLI. " ACLRP?J A?TGRW, A?JJCB RFC hemocoel (MP @JMMB A?TGRW), GQ NPCQCLR, ?LB RFC MNCL AGPASJ?RMPW QWQRCK GQ PCESJ?RCB @W ? RS@SJ?P MP QGLEJC-AF?K@CPCB FC?PR. 3CQNGP?RMPW QWQRCKQ T?PW BCNCLBGLE ML RFC EPMSN MD ?PRFPMNMB: GLQCARQ ?LB KWPG?NMBQ SQC ? QCPGCQ MD RS@CQ (RP?AFC?C) RF?R @P?LAF RFPMSEF RFC @MBW, MNCL RM RFC MSRQGBC RFPMSEF MNCLGLEQ A?JJCB QNGP?AJCQ, ?LB NCPDMPK E?Q CVAF?LEC BGPCARJW @CRUCCL RFC ACJJQ ?LB ?GP GL RFC RP?AFC?C, UFCPC?Q ?OS?RGA APSQR?AC?LQ SRGJGXC EGJJQ, RCPPCQRPG?J AFCJGACP?RCQ CKNJMW @MMI JSLEQ, ?LB ?OS?RGA AFCJGACP?RCQ SQC @MMI EGJJQ (Figure 28.35). 5FC @MMI JSLEQ MD ?P?AFLGBQ (QAMPNGMLQ, QNGBCPQ, RGAIQ ?LB KGRCQ) AMLR?GL ? TCPRGA?J QR?AI MD FCKMAMCJ U?JJ RGQQSC RF?R QMKCUF?R PCQCK@JCQ RFC N?ECQ MD ? @MMI. #CRUCCL C?AF MD RFC "N?ECQ" MD RGQQSC GQ ?L ?GP QN?AC. 5FGQ ?JJMUQ @MRF QGBCQ MD RFC RGQQSC RM @C GL AMLR?AR UGRF RFC ?GP ?R ?JJ RGKCQ, EPC?RJW GLAPC?QGLE RFC CDDGAGCLAW MD E?Q CVAF?LEC. 5FC EGJJQ MD APSQR?AC?LQ ?PC DGJ?KCLRMSQ QRPSARSPCQ RF?R CVAF?LEC E?QCQ UGRF RFC QSPPMSLBGLE U?RCP. (PMSNQ MD ?PRFPMNMBQ ?JQM BGDDCP GL RFC MPE?LQ SQCB DMP CVAPCRGML, UGRF APSQR?AC?LQ NMQQCQQGLE EPCCL EJ?LBQ ?LB GLQCARQ SQGLE .?JNGEFG?L RS@SJCQ, UFGAF UMPI GL AMLHSLARGML UGRF RFC FGLBESR RM PC?@QMP@ U?RCP UFGJC PGBBGLE RFC @MBW MD LGRPMECLMSQ U?QRC. 5FC ASRGAJC GQ RFC AMTCPGLE MD ?L ?PRFPMNMB. *R GQ K?BC SN MD RUM J?WCPQ: RFC CNGASRGAJC, UFGAF GQ ? RFGL, U?VW U?RCP-PCQGQR?LR MSRCP J?WCP AMLR?GLGLE LM AFGRGL, ?LB RFC J?WCP @CLC?RF GR, RFC AFGRGLMSQ NPMASRGAJC. $FGRGL GQ ? RMSEF, DJCVG@JC NMJWQ?AAF?PGBC. *L MPBCP RM EPMU, RFC ?PRFPMNMB KSQR QFCB RFC CVMQICJCRML BSPGLE ? NPMACQQ A?JJCB CABWQGQ (tRM QRPGN MDDu); RFGQ GQ ? ASK@CPQMKC KCRFMB MD EPMURF, ?LB BSPGLE RFGQ RGKC, RFC ?LGK?J GQ TSJLCP?@JC RM NPCB?RGML. 5FC AF?P?ARCPGQRGA KMPNFMJMEW MD PCNPCQCLR?RGTC ?LGK?JQ DPMK C?AF QS@NFWJSK GQ BCQAPG@CB @CJMU.
784
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!:(5.>1:2 H+='54*' 5FC L?KC )CV?NMB? BCLMRCQ RFC NPCQCLAC MD QGV JCEQ (RFPCC N?GPQ) GL RFCQC ?LGK?JQ ?Q BGDDCPCLRG?RCB DPMK RFC LSK@CP MD N?GPQ NPCQCLR GL MRFCP ?PRFPMNMBQ. )CV?NMBQ ?PC AF?P?ARCPGXCB @W RFC NPCQCLAC MD ? FC?B, RFMP?V, ?LB ?@BMKCL, AMLQRGRSRGLE RFPCC R?EK?. 5FC RFMP?V @C?PQ RFC UGLEQ ?Q UCJJ ?Q QGV JCEQ GL RFPCC N?GPQ. .?LW MD RFC AMKKML GLQCARQ UC CLAMSLRCP ML ? B?GJW @?QGQqGLAJSBGLE ?LRQ, AMAIPM?AFCQ, @SRRCPDJGCQ, ?LB DJGCQq?PC CV?KNJCQ MD )CV?NMB?. "KMLEQR RFC FCV?NMBQ, RFC GLQCARQ (Figure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785
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igure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igure 28.38).
(@) (?) !@>LI< 28.37 (>) 4EB *<;0.,9( *63,67;9(;( @BKQFMBAB E>P RM QL 15 M>FOP LC IBDP. (?) 4EFP .LOQE !JBOF@>K JFIIFMBAB (N(9*,<: (4,90*(5<:) ?B>OP J>KV IBDP, >IQELRDE KLQ > QELRP>KA, >P FQP K>JB JFDEQ PRDDBPQ. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V "OR@B ->OIFK; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V #LOV :>KHBO)
786
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!:(5.>1:2 C7:89')+' $PSQR?AC?LQ ?PC RFC KMQR BMKGL?LR ?OS?RGA ?PRFPMNMBQ, QGLAC RFC RMR?J LSK@CP MD K?PGLC APSQR?AC?L QNCAGCQ QR?LBQ ?R 67,000, @SR RFCPC ?PC ?JQM DPCQFU?RCP ?LB RCPPCQRPG?J APSQR?AC?L QNCAGCQ. ,PGJJ, QFPGKN, JM@QRCPQ, AP?@Q, ?LB AP?WDGQF ?PC CV?KNJCQ MD APSQR?AC?LQ (Figure 28.38). 5CPPCQRPG?J QNCAGCQ JGIC RFC UMMB JGAC (A94(+0330+0<4 QNN.) (?JQM A?JJCB NGJJ @SEQ, PMJJW NMJJGCQ, NMR?RM @SEQ, MP GQMNMBQ) ?PC ?JQM APSQR?AC?LQ, ?JRFMSEF RFC LSK@CP MD LML-?OS?RGA QNCAGCQ GL RFGQ QS@NFWJSK GQ PCJ?RGTCJW JMU.
!@>LI< 28.38 4EB (>) @O>? >KA (?) PEOFJM HOFII >OB ?LQE @ORPQ>@B>KP. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V 7FIIF>J 7>O?V; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V *LK 3RIIFS>K)
$PSQR?AC?LQ NMQQCQQ RUM N?GPQ MD ?LRCLL?C, K?LBG@JCQ ?Q KMSRFN?PRQ, ?LB biramous (tRUM @P?LAFCBu) ?NNCLB?ECQ, UFGAF KC?LQ RF?R RFCGP JCEQ ?PC DMPKCB GL RUM N?PRQ, ?Q BGQRGLAR DPMK RFC uniramous (tMLC @P?LAFCBu) KWPG?NMBQ ?LB FCV?NMBQ (Figure 28.39).
!@>LI< 28.39 !OQEOLMLAP J>V E>SB (>) ?FO>JLRP (QTL-?O>K@EBA) >MMBKA>DBP LO (?) RKFO>JLRP (LKB?O>K@EBA) >MMBKA>DBP. (@OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V .F@ELI>P 7. "BBPLK)
6LJGIC RF?R MD RFC )CV?NMB?, RFC FC?B ?LB RFMP?V MD KMQR APSQR?AC?LQ GQ DSQCB RM DMPK ? cephalothorax (Figure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787
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.MQR APSQR?AC?LQ ?PC BGMCAGMSQ, UFGAF KC?LQ RF?R RFC QCVCQ ?PC QCN?P?RC. 4MKC QNCAGCQ JGIC @?PL?AJCQ K?W @C hermaphrodites. 4CPG?J FCPK?NFPMBGRGQK, UFCPC RFC EML?B A?L QUGRAF DPMK NPMBSAGLE QNCPK RM MT?, K?W ?JQM @C QCCL GL QMKC QNCAGCQ. 'CPRGJGXCB CEEQ K?W @C FCJB UGRFGL RFC DCK?JC MD RFC QNCAGCQ MP K?W @C PCJC?QCB GL RFC U?RCP. 5CPPCQRPG?J APSQR?AC?LQ QCCI MSR B?KN QN?ACQ GL RFCGP F?@GR?RQ RM J?W CEEQ. -?PT?J QR?ECQq nauplius ?LB zoeaq?PC QCCL GL RFC C?PJW BCTCJMNKCLR MD APSQR?AC?LQ. " cypris J?PT? GQ ?JQM QCCL GL RFC C?PJW BCTCJMNKCLR MD @?PL?AJCQ (Figure 28.41).
!@>LI< 28.41 !II @ORPQ>@B>KP DL QEOLRDE AFCCBOBKQ I>OS>I PQ>DBP. 3ELTK >OB (>) QEB K>RMIFRP I>OS>I PQ>DB LC > Q>AMLIB PEOFJM, (?) QEB @VMOFP I>OS>I PQ>DB LC > ?>OK>@IB, >KA (@) QEB WLB> I>OS>I PQ>DB LC > DOBBK @O>?. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V 53'3; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V -Z. #. -FKDLO>K@B 2LAOdDRBW; @OBAFQ @: JLAFCF@>QFLK LC TLOH ?V ". +FJJBI ?>PBA LK LOFDFK>I TLOH ?V %OKPQ (>B@HBI)
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tFC?Bu R?EKSK GQ LMR SQS?JJW BGQACPLG@JC. 5FC NFWJSK BCPGTCQ GRQ L?KC DPMK RFC DGPQR N?GP MD ?NNCLB?ECQ: RFC chelicerae (Figure 28.42), UFGAF ?PC QNCAG?JGXCB, AJ?U-JGIC MP D?LE-JGIC KMSRFN?PRQ. 5FCQC ?LGK?JQ BM LMR NMQQCQQ ?LRCLL?C. 5FC QCAMLB N?GP MD ?NNCLB?ECQ GQ ILMUL ?Q pedipalps. *L QMKC QNCAGCQ, JGIC QC? QNGBCPQ, ?L ?BBGRGML?J N?GP MD ?NNCLB?ECQ, A?JJCB ovigers, GQ NPCQCLR @CRUCCL RFC AFCJGACP?C ?LB NCBGN?JNQ.
788
#(!04%2 28 X ).6%24%"2!4%3
!@>LI< 28.42 4EB @EBIF@BO>B (CFOPQ PBQ LC >MMBKA>DBP) >OB TBII ABSBILMBA FK QEB P@LOMFLK. (@OBAFQ: +BSFK 7>IPE)
$FCJGACP?C ?PC KMQRJW SQCB DMP DCCBGLE, @SR GL QNGBCPQ, RFCQC ?PC MDRCL KMBGDGCB GLRM D?LEQ RF?R GLHCAR TCLMK GLRM RFCGP NPCW @CDMPC DCCBGLE (Figure 28.43). .CK@CPQ MD RFGQ QS@NFWJSK F?TC ?L MNCL AGPASJ?RMPW QWQRCK UGRF ? FC?PR RF?R NSKNQ @JMMB GLRM RFC FCKMAMCJ. "OS?RGA QNCAGCQ F?TC EGJJQ, UFCPC?Q RCPPCQRPG?J QNCAGCQ F?TC CGRFCP RP?AFC? MP @MMI JSLEQ DMP E?QCMSQ CVAF?LEC.
!@>LI< 28.43 4EB QO>MALLO PMFABO, IFHB >II PMFABOP, FP > JBJ?BO LC QEB PR?MEVIRJ #EBIF@BO>Q>. (@OBAFQ: ->OPE>I (BAFK)
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
7GQGR RFGQ site (http://openstaxcollege.org/l/arthropodstory) RM AJGAI RFPMSEF ? JCQQML ML ?PRFPMNMBQ, GLAJSBGLE GLRCP?ARGTC F?@GR?R K?NQ, ?LB KMPC.
#(!04%2 28 X ).6%24%"2!4%3
789
28.5 R .LG<IG?PCLD D<LK<IFJKFD@8

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC BGQRGLESGQFGLE AF?P?ARCPGQRGAQ MD CAFGLMBCPKQ z %CQAPG@C RFC BGQRGLESGQFGLE AF?P?ARCPGQRGAQ MD AFMPB?RCQ 5FC NFWJ? &AFGLMBCPK?R? ?LB $FMPB?R? (RFC NFWJSK GL UFGAF FSK?LQ ?PC NJ?ACB) @MRF @CJMLE RM RFC QSNCPNFWJSK %CSRCPMQRMKG?. 3CA?JJ RF?R NPMRMQRMKC ?LB BCSRCPMQRMKCQ BGDDCP GL ACPR?GL ?QNCARQ MD RFCGP CK@PWMLGA BCTCJMNKCLR, ?LB RFCW ?PC L?KCB @?QCB ML UFGAF MNCLGLE MD RFC BGECQRGTC A?TGRW BCTCJMNQ DGPQR. 5FC UMPB BCSRCPMQRMKC AMKCQ DPMK RFC (PCCI UMPB KC?LGLE tKMSRF QCAMLB,u GLBGA?RGLE RF?R RFC ?LSQ GQ RFC DGPQR RM BCTCJMN. 5FCPC ?PC ? QCPGCQ MD MRFCP BCTCJMNKCLR?J AF?P?ARCPGQRGAQ RF?R BGDDCP @CRUCCL NPMRMQRMKCQ ?LB BCSRCPMQRMKCQ, GLAJSBGLE RFC KMBC MD DMPK?RGML MD RFC AMCJMK ?LB RFC C?PJW ACJJ BGTGQGML MD RFC CK@PWM. *L BCSRCPMQRMKCQ, GLRCPL?J NMAICRQ MD RFC CLBMBCPK?J JGLGLE A?JJCB RFC archenteron DSQC RM DMPK RFC AMCJMK. 5FC CLBMBCPK?J JGLGLE MD RFC ?PAFCLRCPML (MP RFC NPGKGRGTC ESR) DMPKQ KCK@P?LC NPMRPSQGMLQ RF?R @SB MDD ?LB @CAMKC RFC KCQMBCPK?J J?WCP. 5FCQC @SBQ, ILMUL ?Q AMCJMKGA NMSAFCQ, DSQC RM DMPK RFC AMCJMKGA A?TGRW, ?Q RFCW CTCLRS?JJW QCN?P?RC DPMK RFC CLBMBCPK?J J?WCP. 5FC PCQSJR?LR AMCJMK GQ RCPKCB ?L enterocoelom. 5FC ?PAFCLRCPML BCTCJMNQ GLRM RFC ?JGKCLR?PW A?L?J, ?LB ? KMSRF MNCLGLE GQ DMPKCB @W GLT?EGL?RGML MD CARMBCPK ?R RFC NMJC MNNMQGRC RFC @J?QRMNMPC MD RFC E?QRPSJ?. 5FC @J?QRMNMPC DMPKQ RFC ?LSQ MD RFC ?JGKCLR?PW QWQRCK GL RFC HSTCLGJC ?LB ?BSJR DMPKQ. 5FC D?RCQ MD CK@PWMLGA ACJJQ GL BCSRCPMQRMKCQ A?L @C ?JRCPCB GD RFCW ?PC CVNCPGKCLR?JJW KMTCB RM ? BGDDCPCLR JMA?RGML GL RFC CK@PWM BSC RM GLBCRCPKGL?LR AJC?T?EC GL C?PJW CK@PWMECLCQGQ.
+?PCLD :?@EF;<ID8K8
&AFGLMBCPK?R? ?PC QM L?KCB MUGLE RM RFCGP QNGLW QIGL (DPMK RFC (PCCI tCAFGLMQu KC?LGLE tQNGLWu ?LB tBCPKMQu KC?LGLE tQIGLu), ?LB RFGQ NFWJSK GQ ? AMJJCARGML MD ?@MSR 7,000 BCQAPG@CB JGTGLE QNCAGCQ. Echinodermata ?PC CVAJSQGTCJW K?PGLC MPE?LGQKQ. 4C? QR?PQ (Figure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water vascular system. "L GLRCPCQRGLE DC?RSPC MD RFCQC ?LGK?JQ GQ RFCGP NMUCP RM PCECLCP?RC, CTCL UFCL MTCP 75 NCPACLR MD RFCGP @MBW K?QQ GQ JMQR.
790
#(!04%2 28 X ).6%24%"2!4%3
!@>LI< 28.44 4EFP AF>DO>J PELTP QEB >K>QLJV LC > PB> PQ>O.
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madreporite. +574*:)9/43 &AFGLMBCPKQ ?PC QCVS?JJW BGKMPNFGA ?LB PCJC?QC RFCGP CEEQ ?LB QNCPK ACJJQ GLRM U?RCP; DCPRGJGX?RGML GQ CVRCPL?J. *L QMKC QNCAGCQ, RFC J?PT?C BGTGBC ?QCVS?JJW ?LB KSJRGNJW @CDMPC RFCW PC?AF QCVS?J K?RSPGRW. &AFGLMBCPKQ K?W ?JQM PCNPMBSAC ?QCVS?JJW, ?Q UCJJ ?Q PCECLCP?RC @MBW N?PRQ JMQR GL RP?SK?.
CC8JJ<J F= :?@EF;<IDJ
5FGQ NFWJSK GQ BGTGBCB GLRM DGTC CVR?LR AJ?QQCQ: "QRCPMGBC? (QC? QR?PQ), 0NFGSPMGBC? (@PGRRJC QR?PQ), &AFGLMGBC? (QC? SPAFGLQ ?LB Q?LB BMJJ?PQ), $PGLMGBC? (QC? JGJGCQ MP DC?RFCP QR?PQ), ?LB )MJMRFSPMGBC? (QC? ASASK@CPQ) (Figure 28.45). 5FC KMQR UCJJ-ILMUL CAFGLMBCPKQ ?PC KCK@CPQ MD AJ?QQ "QRCPMGBC?, MP QC? QR?PQ. 5FCW AMKC GL ? J?PEC T?PGCRW MD QF?NCQ, AMJMPQ, ?LB QGXCQ, UGRF KMPC RF?L 1,800 QNCAGCQ ILMUL QM D?P. 5FC ICW AF?P?ARCPGQRGA MD QC? QR?PQ RF?R BGQRGLESGQFCQ RFCK DPMK MRFCP CAFGLMBCPK AJ?QQCQ GLAJSBCQ RFGAI ?PKQ (?K@SJ?AP?) RF?R CVRCLB DPMK ? ACLRP?J BGQI UFCPC MPE?LQ NCLCRP?RC GLRM RFC ?PKQ. 4C? QR?PQ SQC RFCGP RS@C DCCR LMR MLJW DMP EPGNNGLE QSPD?ACQ @SR ?JQM DMP EP?QNGLE NPCW. 4C? QR?PQ F?TC RUM QRMK?AFQ, MLC MD UFGAF A?L NPMRPSBC RFPMSEF RFCGP KMSRFQ ?LB QCAPCRC BGECQRGTC HSGACQ GLRM MP MLRM NPCW, CTCL @CDMPC GLECQRGML. 5FGQ NPMACQQ A?L CQQCLRG?JJW JGOSCDW RFC NPCW ?LB K?IC BGECQRGML C?QGCP.
#(!04%2 28 X ).6%24%"2!4%3
791
&VNJMPC RFC sea stars body plan (http://openstaxcollege.org/l/sea_star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tDSLARGML?Ju @GJ?RCP?J QWKKCRPW ?Q ?BSJRQ, @CA?SQC RFC SLGOSCJW CVRCLBCB MP?J-?@MP?J ?VGQ AMKNCJQ RFC ?LGK?J RM JGC FMPGXMLR?JJW P?RFCP RF?L QR?LB TCPRGA?JJW.
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
+?PCLD C?FI;8K8
"LGK?JQ GL RFC NFWJSK Chordata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
792
#(!04%2 28 X ).6%24%"2!4%3
& 4 / -(.
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
#(!04%2 28 X ).6%24%"2!4%3
793
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sQ @MBW RFPMSEF UFGAF U?RCP JC?TCQ FJK@LD NMPC NPCQCLR ML RFC QNMLECs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
794
#(!04%2 28 X ).6%24%"2!4%3
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
C#A+/ - .0((A-4
28.1 +?PCLD +FI@=<I8
"LGK?JQ GLAJSBCB GL NFWJSK 1MPGDCP? ?PC 1?P?XM?LQ @CA?SQC RFCW BM LMR QFMU RFC DMPK?RGML MD RPSC RGQQSCQ (CVACNR GL AJ?QQ )CV?ARGLCJJGB?). 5FCQC MPE?LGQKQ QFMU TCPW QGKNJC MPE?LGX?RGML, UGRF ? PSBGKCLR?PW CLBMQICJCRML. 4NMLECQ F?TC KSJRGNJC ACJJ RWNCQ RF?R ?PC EC?PCB RMU?PB CVCASRGLE T?PGMSQ KCR?@MJGA DSLARGMLQ. "JRFMSEF RFCQC ?LGK?JQ ?PC TCPW QGKNJC, RFCW NCPDMPK QCTCP?J AMKNJCV NFWQGMJMEGA?J DSLARGMLQ.
28.2 +?PCLD CE@;8I@8

$LGB?PG?LQ PCNPCQCLR ? KMPC AMKNJCV JCTCJ MD MPE?LGX?RGML RF?L 1MPGDCP?. 5FCW NMQQCQQ MSRCP ?LB GLLCP RGQQSC J?WCPQ RF?R Q?LBUGAF ? LMLACJJSJ?P KCQMEJC?. $LGB?PG?LQ NMQQCQQ ? UCJJ-DMPKCB BGECQRGTC QWQRCK ?LB A?PPW MSR CVRP?ACJJSJ?P BGECQRGML. 5FC ALGBMAWRC GQ ? QNCAG?JGXCB ACJJ DMP BCJGTCPGLE RMVGLQ RM NPCW ?Q UCJJ ?Q U?PLGLE MDD NPCB?RMPQ. $LGB?PG?LQ F?TC QCN?P?RC QCVCQ ?LB F?TC ? JGDCAWAJC RF?R GLTMJTCQ KMPNFMJMEGA?JJW BGQRGLAR DMPKQ. 5FCQC ?LGK?JQ ?JQM QFMU RUM BGQRGLAR KMPNFMJMEGA?J DMPKQqKCBSQMGB ?LB NMJWNMGBq?R T?PGMSQ QR?ECQ GL RFCGP JGDCAWAJC.
28.3 .LG<IG?PCLD 'FG?FKIF:?FQF8

1FWJSK "LLCJGB? GLAJSBCQ TCPKGDMPK, QCEKCLRCB ?LGK?JQ. 4CEKCLR?RGML GQ QCCL GL GLRCPL?J ?L?RMKW ?Q UCJJ, UFGAF GQ A?JJCB KCR?KCPGQK. "LLCJGBQ ?PC NPMRMQRMKCQ. 5FCQC ?LGK?JQ F?TC UCJJ-BCTCJMNCB LCSPML?J ?LB BGECQRGTC QWQRCKQ. 4MKC QNCAGCQ @C?P ? QNCAG?JGXCB @?LB MD QCEKCLRQ ILMUL ?Q ? AJGRCJJSK. "LLCJGBQ QFMU RFC NPCQCLAC LSKCPMSQ AFGRGLMSQ NPMHCARGMLQ RCPKCB AF?CR?C, ?LB NMJWAF?CRCQ NMQQCQQ N?P?NMBG?. 4SAICPQ ?PC QCCL GL MPBCP )GPSBGLC?. 3CNPMBSARGTC QRP?RCEGCQ GLAJSBC QCVS?J BGKMPNFGQK, FCPK?NFPMBGRGQK, ?LB QCPG?J FCPK?NFPMBGRGQK. *LRCPL?J QCEKCLR?RGML GQ ?@QCLR GL AJ?QQ )GPSBGLC?. 'J?RUMPKQ ?PC ?AMCJMK?RC, RPGNJM@J?QRGA ?LGK?JQ. 5FCW J?AI AGPASJ?RMPW ?LB PCQNGP?RMPW QWQRCKQ, ?LB F?TC ? PSBGKCLR?PW CVAPCRMPW QWQRCK. 5FGQ BGECQRGTC QWQRCK GQ GLAMKNJCRC GL KMQR QNCAGCQ. 5FCPC ?PC DMSP RP?BGRGML?J AJ?QQCQ MD DJ?RUMPKQ, RFC J?PECJW DPCC-JGTGLE RSP@CJJ?PG?LQ, RFC CARMN?P?QGRGA KMLMECLC?LQ, ?LB RFC CLBMN?P?QGRGA RPCK?RMBCQ ?LB ACQRMBCQ. 5PCK?RMBCQ F?TC AMKNJCV JGDCAWAJCQ GLTMJTGLE ? KMJJSQA?L QCAMLB?PW FMQR ?LB ? NPGK?PW FMQR GL UFGAF QCVS?J PCNPMBSARGML R?ICQ NJ?AC. $CQRMBCQ, MP R?NCUMPKQ, GLDCAR RFC BGECQRGTC QWQRCKQ MD NPGK?PW TCPRC@P?RC FMQRQ. 5FC PMRGDCPQ ?PC KGAPMQAMNGA, KSJRGACJJSJ?P, KMQRJW ?OS?RGA MPE?LGQKQ RF?R ?PC ASPPCLRJW SLBCP R?VMLMKGA PCTGQGML. 5FC EPMSN GQ AF?P?ARCPGXCB @W RFC PMR?RGLE, AGJG?RCB, UFCCJ-JGIC QRPSARSPC, RFC AMPML?, ML RFCGP FC?B. 5FC K?QR?V MP H?UCB NF?PWLV GQ ?LMRFCP QRPSARSPC SLGOSC RM RFGQ EPMSN MD MPE?LGQKQ. 5FC LCKCPRGLG ?PC RFC QGKNJCQR CSAMCJMK?RCQ. 5FCQC PG@@ML-QF?NCB ?LGK?JQ @C?P ? QNCAG?JGXCB NPM@MQAGQ CLAJMQCB UGRFGL ? PFWLAFMAMCJ. 5FC BCTCJMNKCLR MD ? AJMQCB AGPASJ?RMPW QWQRCK BCPGTCB DPMK RFC AMCJMK GQ ? QGELGDGA?LR BGDDCPCLAC QCCL GL RFGQ QNCAGCQ AMKN?PCB RM MRFCP NQCSBMAMCJMK?RC NFWJ?. "JGKCLR?PW, LCPTMSQ, ?LB CVAPCRMPW QWQRCKQ ?PC KMPC BCTCJMNCB GL RFC LCKCPRGLG RF?L GL JCQQ ?BT?LACB NFWJ?. &K@PWMLGA BCTCJMNKCLR MD LCKCPRGLC UMPKQ NPMACCBQ TG? ? NJ?LSJGDMPK J?PT?J QR?EC. 1FWJSK .MJJSQA? GQ ? J?PEC, K?PGLC EPMSN MD GLTCPRC@P?RCQ. .MJJSQIQ QFMU ? T?PGCRW MD KMPNFMJMEGA?J T?PG?RGMLQ UGRFGL RFC NFWJSK. 5FGQ NFWJSK GQ ?JQM BGQRGLAR GL RF?R QMKC KCK@CPQ CVFG@GR ? A?JA?PCMSQ QFCJJ ?Q ?L CVRCPL?J KC?LQ MD NPMRCARGML. 4MKC KMJJSQIQ F?TC CTMJTCB ? PCBSACB QFCJJ.
#(!04%2 28 X ).6%24%"2!4%3
795
.MJJSQIQ ?PC NPMRMQRMKCQ. 5FC BMPQ?J CNGBCPKGQ GL KMJJSQIQ GQ KMBGDGCB RM DMPK RFC K?LRJC, UFGAF CLAJMQCQ RFC K?LRJC A?TGRW ?LB TGQACP?J MPE?LQ. 5FGQ A?TGRW GQ OSGRC BGQRGLAR DPMK RFC AMCJMKGA A?TGRW, UFGAF GL RFC ?BSJR ?LGK?J QSPPMSLBQ RFC FC?PR. 3CQNGP?RGML GQ D?AGJGR?RCB @W EGJJQ ILMUL ?Q ARCLGBG?. " AFGRGLMSQ-RMMRFCB RMLESC A?JJCB RFC P?BSJ? GQ NPCQCLR GL KMQR KMJJSQIQ. &?PJW BCTCJMNKCLR GL QMKC QNCAGCQ MAASPQ TG? RUM J?PT?J QR?ECQ: RPMAFMNFMPC ?LB TCJGECP. 4CVS?J BGKMPNFGQK GQ RFC NPCBMKGL?LR QCVS?J QRP?RCEW GL RFGQ NFWJSK. .MJJSQIQ A?L @C BGTGBCB GLRM QCTCL AJ?QQCQ, C?AF UGRF BGQRGLAR KMPNFMJMEGA?J AF?P?ARCPGQRGAQ.
28.4 .LG<IG?PCLD :;PJFQF8

/CK?RMBCQ ?PC NQCSBMAMCJMK?RC ?LGK?JQ ?IGL RM DJ?RUMPKQ, WCR BGQNJ?W KMPC ?BT?LACB LCSPML?J BCTCJMNKCLR, ? AMKNJCRC BGECQRGTC QWQRCK, ?LB ? @MBW A?TGRW. 5FGQ NFWJSK GLAJSBCQ DPCC-JGTGLE ?Q UCJJ ?Q N?P?QGRGA MPE?LGQKQ JGIC C(,569/()+0;0: ,3,.(5: ?LB A:*(90: QNN., PCQNCARGTCJW. 5FCW GLAJSBC BGMCACMSQ ?Q UCJJ ?Q FCPK?NFPMBGRGA QNCAGCQ. /CK?RMBCQ ?JQM NMQQCQQ ?L CVAPCRMPW QWQRCK RF?R GQ LMR OSGRC UCJJ BCTCJMNCB. &K@PWMLGA BCTCJMNKCLR GQ CVRCPL?J ?LB NPMACCBQ TG? RFPCC J?PT?J QR?ECQ. " NCASJG?P DC?RSPC MD LCK?RMBCQ GQ RFC QCAPCRGML MD ? AMJJ?ECLMSQ/AFGRGLMSQ ASRGAJC MSRQGBC RFC @MBW. "PRFPMNMBQ PCNPCQCLR RFC KMQR QSAACQQDSJ NFWJSK MD ?LGK?J ML &?PRF, GL RCPKQ MD RFC LSK@CP MD QNCAGCQ ?Q UCJJ ?Q RFC LSK@CP MD GLBGTGBS?JQ. 5FCQC ?LGK?JQ ?PC AF?P?ARCPGXCB @W ? QCEKCLRCB @MBW ?Q UCJJ ?Q RFC NPCQCLAC MD HMGLRCB ?NNCLB?ECQ. *L RFC @?QGA @MBW NJ?L, ? N?GP MD ?NNCLB?ECQ GQ NPCQCLR NCP @MBW QCEKCLR. 8GRFGL RFC NFWJSK, RP?BGRGML?J AJ?QQGDGA?RGML GQ @?QCB ML KMSRFN?PRQ, LSK@CP MD ?NNCLB?ECQ, ?LB KMBGDGA?RGMLQ MD ?NNCLB?ECQ NPCQCLR. "PRFPMNMBQ @C?P ? AFGRGLMSQ CVMQICJCRML. (GJJQ, RP?AFC?, ?LB @MMI JSLEQ D?AGJGR?RC PCQNGP?RGML. 4CVS?J BGKMPNFGQK GQ QCCL GL RFGQ NFWJSK, ?LB CK@PWMLGA BCTCJMNKCLR GLAJSBCQ KSJRGNJC J?PT?J QR?ECQ.
28.5 .LG<IG?PCLD D<LK<IFJKFD@8

&AFGLMBCPKQ ?PC BCSRCPMQRMKGA K?PGLC MPE?LGQKQ. 5FGQ NFWJSK MD ?LGK?JQ @C?PQ ? A?JA?PCMSQ CLBMQICJCRML AMKNMQCB MD MQQGAJCQ. 5FCQC ?LGK?JQ ?JQM F?TC QNGLW QIGL. &AFGLMBCPKQ NMQQCQQ U?RCP@?QCB AGPASJ?RMPW QWQRCKQ. " NMPC RCPKCB RFC K?BPCNMPGRC GQ RFC NMGLR MD CLRPW ?LB CVGR DMP U?RCP GLRM RFC U?RCP T?QASJ?P QWQRCK. 0QKMPCESJ?RGML GQ A?PPGCB MSR @W QNCAG?JGXCB ACJJQ ILMUL ?Q NMBMAWRCQ. 5FC AF?P?ARCPGQRGA DC?RSPCQ MD $FMPB?R? ?PC ? LMRMAFMPB, ? BMPQ?J FMJJMU LCPTC AMPB, NF?PWLEC?J QJGRQ, ?LB ? NMQR-?L?J R?GJ. $FMPB?R? AMLR?GLQ RUM AJ?BCQ MD GLTCPRC@P?RCQ: 6PMAFMPB?R? (RSLGA?RCQ) ?LB $CNF?JMAFMPB?R? (J?LACJCRQ), RMECRFCP UGRF RFC TCPRC@P?RCQ GL 7CPRC@P?R?. .MQR RSLGA?RCQ JGTC ML RFC MAC?L DJMMP ?LB ?PC QSQNCLQGML DCCBCPQ. -?LACJCRQ ?PC QSQNCLQGML DCCBCPQ RF?R DCCB ML NFWRMNJ?LIRML ?LB MRFCP KGAPMMPE?LGQKQ.
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1. Figure 28.3 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ GQ D?JQC >. $FM?LMAWRCQ F?TC DJ?ECJJ? RF?R NPMNCJ U?RCP RFPMSEF RFC @MBW. ?. 1GL?AMAWRCQ A?L RP?LQDMPK GLRM ?LW ACJJ RWNC. @. -MNFMAWRCQ QCAPCRC AMJJ?ECL. A. 1MPMAWRCQ AMLRPMJ RFC DJMU MD U?RCP RFPMSEF NMPCQ GL RFC QNMLEC @MBW. 2. Figure 28.20 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR RFC ?L?RMKW MD ? KMJJSQI GQ D?JQC >. .MJJSQIQ F?TC ? P?BSJ? DMP EPGLBGLE DMMB. ?. " BGECQRGTC EJ?LB GQ AMLLCARCB RM RFC QRMK?AF. @. 5FC RGQQSC @CLC?RF RFC QFCJJ GQ A?JJCB RFC K?LRJC. A. 5FC BGECQRGTC QWQRCK GLAJSBCQ ? EGXX?PB, ? QRMK?AF, ? BGECQRGTC EJ?LB, ?LB RFC GLRCQRGLC. 3. Figure 28.36 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR GLQCARQ GQ D?JQC >. *LQCARQ F?TC @MRF BMPQ?J ?LB TCLRP?J @JMMB TCQQCJQ. ?. *LQCARQ F?TC QNGP?AJCQ, MNCLGLEQ RF?R ?JJMU ?GP RM CLRCP. @. 5FC RP?AFC? GQ N?PR MD RFC BGECQRGTC QWQRCK. A. *LQCARQ F?TC ? BCTCJMNCB BGECQRGTC QWQRCK UGRF ? KMSRF, APMN, ?LB GLRCQRGLC.
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4. .CQMFWJ AMLR?GLQ: >. ? NMJWQ?AAF?PGBC ECJ ?LB BC?B ACJJQ ?. ? AMJJ?ECL-JGIC ECJ ?LB QSQNCLBCB ACJJQ DMP T?PGMSQ DSLARGMLQ @. QNGASJCQ AMKNMQCB MD QGJGA? MP A?JAGSK A?P@ML?RC A. KSJRGNJC NMPCQ 5. 5FC J?PEC ACLRP?J MNCLGLE GL RFC 1?P?XM?L @MBW GQ A?JJCB RFC:
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
13. 5FC LCK?RMBC ASRGAJC AMLR?GLQ >>>>>. >. ?. @. A. EJSAMQC QIGL ACJJQ AFGRGL LCPTC ACJJQ
14. $PSQR?AC?LQ ?PC >>>>>. >. CABWQMXM?LQ ?. LCK?RMBCQ @. ?P?AFLGBQ A. N?P?XM?LQ 15. 'JGCQ ?PC>>>>>>>. >. AFCJGACP?RCQ ?. FCV?NMBQ @. ?P?AFLGBQ A. APSQR?AC?LQ 16. &AFGLMBCPKQ F?TC >>>>>. >. RPG?LESJ?P QWKKCRPW ?. P?BG?J QWKKCRPW @. FCV?EML?J QWKKCRPW A. NCLR?P?BG?J QWKKCRPW 17. 5FC AGPASJ?RMPW DJSGB GL CAFGLMBCPKQ GQ >>>>>. >. ?. @. A. @JMMB KCQMFWJ U?RCP Q?JGLC
11. 5FC PFWLAFMAMCJ GQ ? >>>>>>>>. >. AGPASJ?RMPW QWQRCK ?. DJSGB-DGJJCB A?TGRW @. NPGKGRGTC CVAPCRMPW QWQRCK
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18. %CQAPG@C RFC BGDDCPCLR ACJJ RWNCQ ?LB RFCGP DSLARGMLQ GL QNMLECQ. 19. %CQAPG@C RFC DCCBGLE KCAF?LGQK MD QNMLECQ ?LB GBCLRGDW FMU GR GQ BGDDCPCLR DPMK MRFCP ?LGK?JQ. 20. &VNJ?GL RFC DSLARGML MD LCK?RMAWQRQ GL ALGB?PG?LQ. 21. $MKN?PC RFC QRPSARSP?J BGDDCPCLACQ @CRUCCL 1MPGDCP? ?LB $LGB?PG?. 22. %CQAPG@C RFC KMPNFMJMEW ?LB ?L?RMKW MD KMJJSQIQ. 23. 8F?R ?PC RFC ?L?RMKGA?J BGDDCPCLACQ @CRUCCL LCKCPRGLCQ ?LB KMJJSQIQ 24. &LSKCP?RC DC?RSPCQ MD C(,569/()+0;0: ,3,.(5: RF?R K?IC GR ? T?JS?@JC KMBCJ QWQRCK DMP @GMJMEGQRQ. 25. 8F?R ?PC RFC BGDDCPCLR U?WQ GL UFGAF LCK?RMBCQ A?L PCNPMBSAC 26. %CQAPG@C RFC T?PGMSQ QSNCPAJ?QQCQ RF?R NFWJSK "PRFPMNMB? A?L @C BGTGBCB GLRM. 27. $MKN?PC ?LB AMLRP?QR RFC QCEKCLR?RGML QCCL GL NFWJSK "LLCJGB? UGRF RF?R QCCL GL NFWJSK "PRFPMNMB?. 28. %CQAPG@C RFC BGDDCPCLR AJ?QQCQ MD CAFGLMBCPKQ SQGLE CV?KNJCQ.
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7CPRC@P?RCQ ?PC ?KMLE RFC KMQR PCAMELGX?@JC MPE?LGQKQ MD RFC ?LGK?J IGLEBMK. .MPC RF?L 62,000 TCPRC@P?RC QNCAGCQ F?TC @CCL GBCLRGDGCB. 5FC TCPRC@P?RC QNCAGCQ LMU JGTGLE PCNPCQCLR MLJW ? QK?JJ NMPRGML MD RFC TCPRC@P?RCQ RF?R F?TC CVGQRCB. 5FC @CQR-ILMUL CVRGLAR TCPRC@P?RCQ ?PC RFC BGLMQ?SPQ, ? SLGOSC EPMSN MD PCNRGJCQ, UFGAF PC?AFCB QGXCQ LMR QCCL @CDMPC MP ?DRCP GL RCPPCQRPG?J ?LGK?JQ. 5FCW UCPC RFC BMKGL?LR RCPPCQRPG?J ?LGK?JQ DMP 150 KGJJGML WC?PQ, SLRGJ RFCW BGCB MSR GL ? K?QQ CVRGLARGML LC?P RFC CLB MD RFC $PCR?ACMSQ NCPGMB. "JRFMSEF GR GQ LMR ILMUL UGRF ACPR?GLRW UF?R A?SQCB RFCGP CVRGLARGML, ? EPC?R BC?J GQ ILMUL ?@MSR RFC ?L?RMKW MD RFC BGLMQ?SPQ, EGTCL RFC NPCQCPT?RGML MD QICJCR?J CJCKCLRQ GL RFC DMQQGJ PCAMPB. $SPPCLRJW, ? LSK@CP MD TCPRC@P?RC QNCAGCQ D?AC CVRGLARGML NPGK?PGJW BSC RM F?@GR?R JMQQ ?LB NMJJSRGML. "AAMPBGLE RM RFC *LRCPL?RGML?J 6LGML DMP RFC $MLQCPT?RGML MD /?RSPC, KMPC RF?L 6,000 TCPRC@P?RC QNCAGCQ ?PC AJ?QQGDGCB ?Q RFPC?RCLCB. "KNFG@G?LQ ?LB K?KK?JQ ?PC RFC AJ?QQCQ UGRF RFC EPC?RCQR NCPACLR?EC MD RFPC?RCLCB QNCAGCQ, UGRF 29 NCPACLR MD ?JJ ?KNFG@G?LQ ?LB 21 NCPACLR MD ?JJ K?KK?JQ AJ?QQGDGCB ?Q RFPC?RCLCB. "RRCKNRQ ?PC @CGLE K?BC ?PMSLB RFC UMPJB RM NPCTCLR RFC CVRGLARGML MD RFPC?RCLCB QNCAGCQ. 'MP CV?KNJC, RFC #GMBGTCPQGRW "ARGML 1J?L GQ ?L GLRCPL?RGML?J NPMEP?K, P?RGDGCB @W 188 AMSLRPGCQ, UFGAF GQ BCQGELCB RM NPMRCAR QNCAGCQ ?LB F?@GR?RQ.
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC BGQRGLESGQFGLE AF?P?ARCPGQRGAQ MD AFMPB?RCQ z *BCLRGDW RFC BCPGTCB AF?P?ARCP MD AP?LG?RCQ RF?R QCRQ RFCK ?N?PR DPMK MRFCP AFMPB?RCQ z %CQAPG@C RFC BCTCJMNKCLR?J D?RC MD RFC LMRMAFMPB GL TCPRC@P?RCQ 7CPRC@P?RCQ ?PC KCK@CPQ MD RFC IGLEBMK "LGK?JG? ?LB RFC NFWJSK $FMPB?R? (Figure 29.2). 3CA?JJ RF?R ?LGK?JQ RF?R NMQQCQQ @GJ?RCP?J QWKKCRPW A?L @C BGTGBCB GLRM RUM EPMSNQqNPMRMQRMKCQ ?LB BCSRCPMQRMKCQq@?QCB ML RFCGP N?RRCPLQ MD CK@PWMLGA BCTCJMNKCLR. 5FC BCSRCPMQRMKCQ, UFMQC L?KC RP?LQJ?RCQ ?Q tQCAMLB KMSRF,u AMLQGQR MD RUM NFWJ?: $FMPB?R? ?LB &AFGLMBCPK?R?. &AFGLMBCPKQ ?PC GLTCPRC@P?RC K?PGLC ?LGK?JQ RF?R F?TC NCLR?P?BG?J QWKKCRPW ?LB ? QNGLW @MBW AMTCPGLE, ? EPMSN RF?R GLAJSBCQ QC? QR?PQ, QC? SPAFGLQ, ?LB QC? ASASK@CPQ. 5FC KMQR AMLQNGASMSQ ?LB D?KGJG?P KCK@CPQ MD $FMPB?R? ?PC TCPRC@P?RCQ, @SR RFGQ NFWJSK ?JQM GLAJSBCQ RUM EPMSNQ MD GLTCPRC@P?RC AFMPB?RCQ.
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" cranium GQ ? @MLW, A?PRGJ?EGLMSQ, MP DG@PMSQ QRPSARSPC QSPPMSLBGLE RFC @P?GL, H?U, ?LB D?AG?J @MLCQ (Figure 29.6). .MQR @GJ?RCP?JJW QWKKCRPGA?J ?LGK?JQ F?TC ? FC?B; MD RFCQC, RFMQC RF?R F?TC ? AP?LGSK AMKNMQC RFC AJ?BC Craniata. $P?LG?R? GLAJSBCQ RFC F?EDGQFCQ (.WVGLG), UFGAF F?TC ? AP?LGSK @SR J?AI ? @?AI@MLC, ?LB ?JJ MD RFC MPE?LGQKQ A?JJCB tTCPRC@P?RCQ.u
!@>LI< 29.6 #O>KF>Q>, FK@IRAFKD QEFP CFPE (D<523,6:;,<: PM.), >OB @E>O>@QBOFWBA ?V QEB MOBPBK@B LC > @O>KFRJ, J>KAF?IB, >KA LQEBO C>@F>I ?LKBP. (@OBAFQ: x3QBSBL@ 86y/7FHFJBAF> #LJJLKP)
7CPRC@P?RCQ ?PC KCK@CPQ MD RFC AJ?BC Vertebrata. 7CPRC@P?RCQ BGQNJ?W RFC DMSP AF?P?ARCPGQRGA DC?RSPCQ MD RFC AFMPB?RCQ; FMUCTCP, KCK@CPQ MD RFGQ EPMSN ?JQM QF?PC BCPGTCB AF?P?ARCPGQRGAQ RF?R BGQRGLESGQF RFCK DPMK GLTCPRC@P?RC AFMPB?RCQ. 7CPRC@P?R? GQ L?KCB DMP RFC vertebral column, AMKNMQCB MD TCPRC@P?C, ? QCPGCQ MD QCN?P?RC @MLCQ HMGLCB RMECRFCP ?Q ? @?AI@MLC (Figure 29.7). *L ?BSJR TCPRC@P?RCQ, RFC TCPRC@P?J AMJSKL PCNJ?ACQ RFC LMRMAFMPB, UFGAF GQ MLJW QCCL GL RFC CK@PWMLGA QR?EC.
!@>LI< 29.7 6BOQB?O>Q> >OB @E>O>@QBOFWBA ?V QEB MOBPBK@B LC > ?>@H?LKB, PR@E >P QEB LKB QE>Q ORKP QEOLRDE QEB JFAAIB LC QEFP CFPE. !II SBOQB?O>QBP >OB FK QEB #O>KF>Q> @I>AB >KA E>SB > @O>KFRJ. (@OBAFQ: %OKBPQ 6. -LOB; Q>HBK >Q 3JFQEPLKF>K -RPBRJ LC .>QRO>I (FPQLOV, 7>PEFKDQLK, $.#.)
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t$?K@PG?L CVNJMQGMLu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
1. $FCL, +. :., )S?LE, %. :., ?LB -G, $. 8., t"L C?PJW $?K@PG?L AP?LG?RC-JGIC AFMPB?RC,u N(;<9, 402 (1999): 518p522, BMG:10.1038/990080.
802
#(!04%2 29 X 6%24%"2!4%3
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tH?UCB KMSRF.u (L?RFMQRMKCQ GLAJSBC DGQFCQ ?LB RCRP?NMBQq?KNFG@G?LQ, PCNRGJCQ, @GPBQ, ?LB K?KK?JQ. 5CRP?NMBQ A?L @C DSPRFCP BGTGBCB GLRM RUM EPMSNQ: ?KNFG@G?LQ ?LB ?KLGMRCQ. "KLGMRCQ ?PC ?LGK?JQ UFMQC CEEQ ?PC ?B?NRCB DMP RCPPCQRPG?J JGTGLE, ?LB RFGQ EPMSN GLAJSBCQ K?KK?JQ, PCNRGJCQ, ?LB @GPBQ. "KLGMRGA CK@PWMQ, BCTCJMNGLE GL CGRFCP ?L CVRCPL?JJW QFCB CEE MP ?L CEE A?PPGCB @W RFC DCK?JC, ?PC NPMTGBCB UGRF ? U?RCP-PCR?GLGLE CLTGPMLKCLR ?LB ?PC NPMRCARCB @W ?KLGMRGA KCK@P?LCQ.
29.2 R !@J?<J
#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC BGDDCPCLAC @CRUCCL H?UJCQQ ?LB H?UCB DGQFCQ z %GQASQQ RFC BGQRGLESGQFGLE DC?RSPCQ MD QF?PIQ ?LB P?WQ AMKN?PCB RM MRFCP KMBCPL DGQFCQ .MBCPL DGQFCQ GLAJSBC ?L CQRGK?RCB 31,000 QNCAGCQ. 'GQFCQ UCPC RFC C?PJGCQR TCPRC@P?RCQ, UGRF H?UJCQQ QNCAGCQ @CGLE RFC C?PJGCQR ?LB H?UCB QNCAGCQ CTMJTGLE J?RCP. 5FCW ?PC ?ARGTC DCCBCPQ, P?RFCP RF?L QCQQGJC, QSQNCLQGML DCCBCPQ. +?UJCQQ DGQFCQqRFC F?EDGQFCQ ?LB J?KNPCWQqF?TC ? BGQRGLAR AP?LGSK ?LB AMKNJCV QCLQC MPE?LQ GLAJSBGLE CWCQ, BGQRGLESGQFGLE RFCK DPMK RFC GLTCPRC@P?RC AFMPB?RCQ.
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+?UJCQQ DGQFCQ ?PC AP?LG?RCQ RF?R PCNPCQCLR ?L ?LAGCLR TCPRC@P?RC JGLC?EC RF?R ?PMQC MTCP MLC F?JD-@GJJGML WC?PQ ?EM. *L RFC N?QR, RFC F?EDGQFCQ ?LB J?KNPCWQ UCPC AJ?QQGDGCB RMECRFCP ?Q ?EL?RF?LQ. 5MB?W, F?EDGQFCQ ?LB J?KNPCWQ ?PC PCAMELGXCB ?Q QCN?P?RC AJ?BCQ, NPGK?PGJW @CA?SQC J?KNPCWQ ?PC RPSC TCPRC@P?RCQ, UFCPC?Q F?EDGQFCQ ?PC LMR. " BCDGLGLE DC?RSPC GQ RFC J?AI MD N?GPCB J?RCP?J ?NNCLB?ECQ (DGLQ). 4MKC MD RFC C?PJGCQR H?UJCQQ DGQFCQ UCPC RFC ostracoderms (UFGAF RP?LQJ?RCQ RM tQFCJJ-QIGLu). 0QRP?AMBCPKQ UCPC TCPRC@P?RC DGQFCQ CLA?QCB GL @MLW ?PKMP, SLJGIC NPCQCLR-B?W H?UJCQQ DGQFCQ, UFGAF J?AI @MLC GL RFCGP QA?JCQ. M>=/3/: H'-,/8.+8 5FC AJ?BC Myxini GLAJSBCQ ?R JC?QR 20 QNCAGCQ MD F?EDGQFCQ. Hagfishes ?PC CCJ-JGIC QA?TCLECPQ RF?R JGTC ML RFC MAC?L DJMMP ?LB DCCB ML BC?B GLTCPRC@P?RCQ, MRFCP DGQFCQ, ?LB K?PGLC K?KK?JQ (Figure 29.8). )?EDGQFCQ ?PC CLRGPCJW K?PGLC ?LB ?PC DMSLB GL MAC?LQ ?PMSLB RFC UMPJB, CVACNR DMP RFC NMJ?P PCEGMLQ. " SLGOSC DC?RSPC MD RFCQC ?LGK?JQ GQ RFC QJGKC EJ?LBQ @CLC?RF RFC QIGL RF?R PCJC?QC KSASQ RFPMSEF QSPD?AC NMPCQ. 5FGQ KSASQ ?JJMUQ RFC F?EDGQF RM CQA?NC DPMK RFC EPGN MD NPCB?RMPQ. )?EDGQF A?L ?JQM RUGQR RFCGP @MBGCQ GL ? ILMR RM DCCB ?LB QMKCRGKCQ C?R A?PA?QQCQ DPMK RFC GLQGBC MSR.
!@>LI< 29.8 0>@FCF@ E>DCFPE >OB P@>SBKDBOP QE>Q IFSB LK QEB L@B>K CILLO. (@OBAFQ: ,FKA> 3KLLH, ./!!/ #".-3)
#(!04%2 29 X 6%24%"2!4%3
803
5FC QICJCRML MD ? F?EDGQF GQ AMKNMQCB MD A?PRGJ?EC, UFGAF GLAJSBCQ ? A?PRGJ?EGLMSQ LMRMAFMPB RF?R PSLQ RFC JCLERF MD RFC @MBW. 5FGQ LMRMAFMPB NPMTGBCQ QSNNMPR RM RFC F?EDGQFsQ @MBW. )?EDGQFCQ BM LMR PCNJ?AC RFC LMRMAFMPB UGRF ? TCPRC@P?J AMJSKL BSPGLE BCTCJMNKCLR, ?Q BM RPSC TCPRC@P?RCQ. P+9742>?439/*'+: L'257+>8 5FC AJ?BC Petromyzontidae GLAJSBCQ ?NNPMVGK?RCJW 35p40 MP KMPC QNCAGCQ MD J?KNPCWQ. Lampreys ?PC QGKGJ?P RM F?EDGQFCQ GL QGXC ?LB QF?NC; FMUCTCP, J?KNPCWQ NMQQCQQ QMKC TCPRC@P?J CJCKCLRQ. -?KNPCWQ J?AI N?GPCB ?NNCLB?ECQ ?LB @MLC, ?Q BM RFC F?EDGQFCQ. "Q ?BSJRQ, J?KNPCWQ ?PC AF?P?ARCPGXCB @W ? RMMRFCB, DSLLCJ-JGIC QSAIGLE KMSRF. .?LW QNCAGCQ F?TC ? N?P?QGRGA QR?EC MD RFCGP JGDC AWAJC BSPGLE UFGAF RFCW ?PC CARMN?P?QGRCQ MD DGQFCQ (Figure 29.9).
!@>LI< 29.9 4EBPB M>O>PFQF@ PB> I>JMOBVP >QQ>@E QL QEBFO I>HB QOLRQ ELPQ ?V PR@QFLK >KA RPB QEBFO OLRDE QLKDRBP QL O>PM >T>V CIBPE FK LOABO QL CBBA LK QEB QOLRQwP ?ILLA. (@OBAFQ: 53'3)
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
"E8K?FJKFD<J: %8N<; !@J?<J

Gnathostomes MP tH?U-KMSRFQu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igure 29.10), UFGAF ?PMQC GL RFC J?RC 4GJSPG?L NCPGMB ?LB ?PC LMU CVRGLAR. .MQR KMBCPL DGQFCQ ?PC EL?RFMQRMKCQ RF?R @CJMLE RM RFC AJ?BCQ $FMLBPGAFRFWCQ ?LB 0QRCGAFRFWCQ.
804
#(!04%2 29 X 6%24%"2!4%3
!@>LI< 29.10 D<523,6:;,6<: T>P >K BKLOJLRP MI>@LABOJ COLJ QEB $BSLKF>K MBOFLA, 380t360 JFIIFLK VB>OP >DL. )Q JB>PROBA RM QL 10 JBQBOP FK IBKDQE >KA TBFDEBA RM QL 3.6 QLKP. (@OBAFQ: .L?R 4>JRO>)
C.43*7/).9.>+8: C'79/1'-/34:8 F/8.+8 5FC AJ?BC Chondrichthyes GQ BGTCPQC, AMLQGQRGLE MD QF?PIQ (Figure 29.11), P?WQ, ?LB QI?RCQ, RMECRFCP UGRF Q?UDGQFCQ ?LB ? DCU BMXCL QNCAGCQ MD DGQFCQ A?JJCB */04(,9(:, MP tEFMQRu QF?PIQ.u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
!@>LI< 29.11 (>JJBOEB>A PE>OHP QBKA QL P@ELLI AROFKD QEB A>V >KA ERKQ MOBV >Q KFDEQ. (@OBAFQ: ->P>PEF 3RD>T>O>)
4F?PIQ F?TC UCJJ-BCTCJMNCB QCLQC MPE?LQ RF?R ?GB RFCK GL JMA?RGLE NPCW, GLAJSBGLE ? ICCL QCLQC MD QKCJJ ?LB CJCARPMPCACNRGML, UGRF RFC J?RRCP NCPF?NQ RFC KMQR QCLQGRGTC MD ?LW ?LGK?J. 0PE?LQ A?JJCB ampullae of Lorenzini ?JJMU QF?PIQ RM BCRCAR RFC CJCARPMK?ELCRGA DGCJBQ RF?R ?PC NPMBSACB @W ?JJ JGTGLE RFGLEQ, GLAJSBGLE RFCGP NPCW. &JCARPMPCACNRGML F?Q MLJW @CCL M@QCPTCB GL ?OS?RGA MP ?KNFG@GMSQ ?LGK?JQ. 4F?PIQ, RMECRFCP UGRF KMQR DGQFCQ ?LB ?OS?RGA ?LB J?PT?J ?KNFG@G?LQ, ?JQM F?TC ? QCLQC MPE?L A?JJCB RFC lateral line, UFGAF GQ SQCB RM BCRCAR KMTCKCLR ?LB TG@P?RGML GL RFC QSPPMSLBGLE U?RCP, ?LB GQ MDRCL AMLQGBCPCB FMKMJMEMSQ RM tFC?PGLEu GL RCPPCQRPG?J TCPRC@P?RCQ. 5FC J?RCP?J JGLC GQ TGQG@JC ?Q ? B?PICP QRPGNC RF?R PSLQ ?JMLE RFC JCLERF MD ? DGQFsQ @MBW. 4F?PIQ PCNPMBSAC QCVS?JJW, ?LB CEEQ ?PC DCPRGJGXCB GLRCPL?JJW. .MQR QNCAGCQ ?PC MTMTGTGN?PMSQ: 5FC DCPRGJGXCB CEE GQ PCR?GLCB GL RFC MTGBSAR MD RFC KMRFCPsQ @MBW ?LB RFC CK@PWM GQ LMSPGQFCB @W RFC CEE WMJI. 5FC CEEQ F?RAF GL RFC SRCPSQ, ?LB WMSLE ?PC @MPL ?JGTC ?LB DSJJW DSLARGML?J. 4MKC QNCAGCQ MD QF?PIQ ?PC MTGN?PMSQ: 5FCW J?W CEEQ RF?R F?RAF MSRQGBC MD RFC KMRFCPsQ @MBW. &K@PWMQ ?PC NPMRCARCB @W ? QF?PI CEE A?QC MP tKCPK?GBsQ NSPQCu (Figure 29.12) RF?R F?Q RFC AMLQGQRCLAW MD JC?RFCP. 5FC QF?PI CEE A?QC F?Q
#(!04%2 29 X 6%24%"2!4%3
805
RCLR?AJCQ RF?R QL?E GL QC?UCCB ?LB EGTC RFC LCU@MPL QF?PI AMTCP. " DCU QNCAGCQ MD QF?PIQ ?PC TGTGN?PMSQ: 5FC WMSLE BCTCJMN UGRFGL RFC KMRFCPsQ @MBW ?LB QFC EGTCQ JGTC @GPRF.
!@>LI< 29.12 3E>OH BJ?OVLP >OB @IB>OIV SFPF?IB QEOLRDE QEBPB QO>KPM>OBKQ BDD @>PBP. 4EB OLRKA PQOR@QROB FP QEB VLIH QE>Q KLROFPEBP QEB DOLTFKD BJ?OVL. (@OBAFQ: *BH ">@>OFP>P)
3?WQ ?LB QI?RCQ AMKNPGQC KMPC RF?L 500 QNCAGCQ ?LB ?PC AJMQCJW PCJ?RCB RM QF?PIQ. 5FCW A?L @C BGQRGLESGQFCB DPMK QF?PIQ @W RFCGP DJ?RRCLCB @MBGCQ, NCARMP?J DGLQ RF?R ?PC CLJ?PECB ?LB DSQCB RM RFC FC?B, ?LB EGJJ QJGRQ ML RFCGP TCLRP?J QSPD?AC (Figure 29.13). -GIC QF?PIQ, P?WQ ?LB QI?RCQ F?TC ? A?PRGJ?EGLMSQ QICJCRML. .MQR QNCAGCQ ?PC K?PGLC ?LB JGTC ML RFC QC? DJMMP, UGRF LC?PJW ? UMPJBUGBC BGQRPG@SRGML.
!@>LI< 29.13 4EFP PQFKDO>V ?IBKAP FKQL QEB P>KAV ?LQQLJ LC QEB L@B>K CILLO. (@OBAFQ: "3>FIK1"/&IF@HO)
O89+/).9.>+8: B43> F/8.+8 .CK@CPQ MD RFC AJ?BC Osteichthyes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swim bladder, ? E?Q-DGJJCB MPE?L RF?R FCJNQ RM AMLRPMJ RFC @SMW?LAW MD RFC DGQF. #MLW DGQFCQ ?PC DSPRFCP BGTGBCB GLRM RUM CVR?LR AJ?BCQ: Actinopterygii (P?W-DGLLCB DGQFCQ) ?LB Sarcopterygii (JM@C-DGLLCB DGQFCQ). "ARGLMNRCPWEGG, RFC P?W-DGLLCB DGQFCQ, GLAJSBC K?LW D?KGJG?P DGQFCQqRSL?, @?QQ, RPMSR, ?LB Q?JKML (Figure 29.14a), ?KMLE MRFCPQ. 3?W-DGLLCB DGQFCQ ?PC L?KCB DMP RFCGP DGLQ RF?R ?PC UC@Q MD QIGL QSNNMPRCB @W @MLW QNGLCQ A?JJCB P?WQ. *L AMLRP?QR, RFC DGLQ MD 4?PAMNRCPWEGG ?PC DJCQFW ?LB JM@CB, QSNNMPRCB @W @MLC (Figure 29.14b). -GTGLE KCK@CPQ MD RFGQ AJ?BC GLAJSBC RFC JCQQ-D?KGJG?P JSLEDGQFCQ ?LB AMCJ?A?LRFQ.
806
#(!04%2 29 X 6%24%"2!4%3
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8?RAF RFGQ QCPGCQ MD DGTC "LGK?J 1J?LCR TGBCMQ ML RCRP?NMB CTMJSRGML: z 1: The evolution from fish to earliest tetrapod (http://openstaxcollege.org/l/tetrapod_evol1) z 2: Fish to Earliest Tetrapod (http://openstaxcollege.org/l/tetrapod_evol2) z 3: The discovery of coelacanth and Acanthostega fossils (http://openstaxcollege.org/l/ tetrapod_evol3) z 4: The number of fingers on legs (http://openstaxcollege.org/l/tetrapod_evol4) z 5: Reconstructing the environment of early tetrapods (http://openstaxcollege.org/l/ tetrapod_evol5)
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC K?GL AF?P?ARCPGQRGAQ MD ?KLGMRCQ z &VNJ?GL RFC BGDDCPCLAC @CRUCCL ?L?NQGBQ, QWL?NQGBQ, ?LB BG?NQGBQ, ?LB EGTC ?L CV?KNJC MD C?AF z *BCLRGDW RFC AF?P?ARCPGQRGAQ MD PCNRGJCQ z %GQASQQ RFC CTMJSRGML MD PCNRGJCQ 5FC ?KLGMRCQ qPCNRGJCQ, @GPBQ, ?LB K?KK?JQq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sQ @MBW; FMUCTCP, CTCL UGRF RFGQ GLRCPL?J ECQR?RGML, ?KLGMRGA KCK@P?LCQ ?PC QRGJJ NPCQCLR.
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5FC ?KLGMRGA CEE GQ RFC ICW AF?P?ARCPGQRGA MD ?KLGMRCQ. *L ?KLGMRCQ RF?R J?W CEEQ, RFC QFCJJ MD RFC CEE NPMTGBCQ NPMRCARGML DMP RFC BCTCJMNGLE CK@PWM UFGJC @CGLE NCPKC?@JC CLMSEF RM ?JJMU DMP RFC CVAF?LEC MD A?P@ML BGMVGBC ?LB MVWECL. 5FC ?J@SKGL, MP CEE UFGRC, NPMTGBCQ RFC CK@PWM UGRF U?RCP ?LB NPMRCGL, UFCPC?Q RFC D?RRGCP CEE WMJI GQ RFC CLCPEW QSNNJW DMP RFC CK@PWM, ?Q GQ RFC A?QC UGRF RFC CEEQ MD K?LW MRFCP ?LGK?JQ, QSAF ?Q ?KNFG@G?LQ. )MUCTCP, RFC CEEQ MD ?KLGMRCQ AMLR?GL RFPCC ?BBGRGML?J CVRP?CK@PWMLGA KCK@P?LCQ: RFC AFMPGML, ?KLGML, ?LB ?JJ?LRMGQ (Figure 29.20). &VRP?-CK@PWMLGA KCK@P?LCQ ?PC KCK@P?LCQ NPCQCLR GL ?KLGMRGA CEEQ RF?R ?PC LMR ? N?PR MD RFC @MBW MD RFC BCTCJMNGLE CK@PWM. 8FGJC RFC GLLCP ?KLGMRGA KCK@P?LC QSPPMSLBQ RFC CK@PWM GRQCJD, RFC chorion QSPPMSLBQ RFC CK@PWM ?LB WMJI Q?A. 5FC AFMPGML D?AGJGR?RCQ CVAF?LEC MD MVWECL ?LB A?P@ML BGMVGBC @CRUCCL RFC CK@PWM ?LB RFC CEEsQ CVRCPL?J CLTGPMLKCLR. 5FC amnion NPMRCARQ RFC CK@PWM DPMK KCAF?LGA?J QFMAI ?LB QSNNMPRQ FWBP?RGML. 5FC allantois QRMPCQ LGRPMECLMSQ U?QRCQ NPMBSACB @W RFC CK@PWM ?LB ?JQM D?AGJGR?RCQ PCQNGP?RGML. *L K?KK?JQ, KCK@P?LCQ RF?R ?PC FMKMJMEMSQ RM RFC CVRP?-CK@PWMLGA KCK@P?LCQ GL CEEQ ?PC NPCQCLR GL RFC NJ?ACLR?.
812
#(!04%2 29 X 6%24%"2!4%3
!@>LI< 29.20 4EB HBV CB>QROBP LC >K >JKFLQF@ BDD >OB PELTK.
7EF@E LC QEB CLIILTFKD PQ>QBJBKQP >?LRQ QEB M>OQP LC >K BDD >OB C>IPB >. 4EB >II>KQLFP PQLOBP KFQOLDBKLRP T>PQB >KA C>@FIFQ>QBP OBPMFO>QFLK. ?. 4EB @ELOFLK C>@FIFQ>QBP D>P BU@E>KDB. @. 4EB VLIH MOLSFABP CLLA CLO QEB DOLTFKD BJ?OVL. A. 4EB >JKFLQF@ @>SFQV FP CFIIBA TFQE >I?RJBK. "BBGRGML?J BCPGTCB AF?P?ARCPGQRGAQ MD ?KLGMRCQ GLAJSBC U?RCPNPMMD QIGL, BSC RM RFC NPCQCLAC MD JGNGBQ, ?LB AMQR?J (PG@) TCLRGJ?RGML MD RFC JSLEQ.
MFCLK@FE F= ADE@FK<J
5FC DGPQR ?KLGMRCQ CTMJTCB DPMK ?KNFG@G?L ?LACQRMPQ ?NNPMVGK?RCJW 340 KGJJGML WC?PQ ?EM BSPGLE RFC $?P@MLGDCPMSQ NCPGMB. 5FC C?PJW ?KLGMRCQ BGTCPECB GLRM RUM K?GL JGLCQ QMML ?DRCP RFC DGPQR ?KLGMRCQ ?PMQC. 5FC GLGRG?J QNJGR U?Q GLRM QWL?NQGBQ ?LB Q?SPMNQGBQ. Synapsids GLAJSBC ?JJ K?KK?JQ, GLAJSBGLE CVRGLAR K?KK?JG?L QNCAGCQ. 4WL?NQGBQ ?JQM GLAJSBC RFCP?NQGBQ, UFGAF UCPC K?KK?J-JGIC PCNRGJCQ DPMK UFGAF K?KK?JQ CTMJTCB. Sauropsids GLAJSBC PCNRGJCQ ?LB @GPBQ, ?LB A?L @C DSPRFCP BGTGBCB GLRM ?L?NQGBQ ?LB BG?NQGBQ. 5FC ICW BGDDCPCLACQ @CRUCCL RFC QWL?NQGBQ, ?L?NQGBQ, ?LB BG?NQGBQ ?PC RFC QRPSARSPCQ MD RFC QISJJ ?LB RFC LSK@CP MD RCKNMP?J DCLCQRP?C @CFGLB C?AF CWC (Figure 29.21). Temporal fenestrae ?PC NMQR-MP@GR?J MNCLGLEQ GL RFC QISJJ RF?R ?JJMU KSQAJCQ RM CVN?LB ?LB JCLERFCL. Anapsids F?TC LM RCKNMP?J DCLCQRP?C, QWL?NQGBQ F?TC MLC, ?LB diapsids F?TC RUM. "L?NQGBQ GLAJSBC CVRGLAR MPE?LGQKQ ?LB K?W, @?QCB ML ?L?RMKW, GLAJSBC RSPRJCQ. )MUCTCP, RFGQ GQ QRGJJ AMLRPMTCPQG?J, ?LB RSPRJCQ ?PC QMKCRGKCQ AJ?QQGDGCB ?Q BG?NQGBQ @?QCB ML KMJCASJ?P CTGBCLAC. 5FC BG?NQGBQ GLAJSBC @GPBQ ?LB ?JJ MRFCP JGTGLE ?LB CVRGLAR PCNRGJCQ.
!@>LI< 29.21 #LJM>OB QEB PHRIIP >KA QBJMLO>I CBKBPQO>B LC >K>MPFAP, PVK>MPFAP, >KA AF>MPFAP. !K>MPFAP E>SB KL LMBKFKDP, PVK>MPFAP E>SB LKB LMBKFKD, >KA AF>MPFAP E>SB QTL LMBKFKDP.
5FC BG?NQGBQ BGTCPECB GLRM RUM EPMSNQ, RFC "PAFMQ?SPMKMPNF? (t?LAGCLR JGX?PB DMPKu) ?LB RFC -CNGBMQ?SPMKMPNF? (tQA?JW JGX?PB DMPKu) BSPGLE RFC .CQMXMGA NCPGMB (Figure 29.22). 5FC lepidosaurs GLAJSBC KMBCPL JGX?PBQ, QL?ICQ, ?LB RS?R?P?Q. 5FC archosaurs GLAJSBC KMBCPL APMAMBGJCQ ?LB ?JJGE?RMPQ, ?LB RFC CVRGLAR NRCPMQ?SPQ (tUGLECB JGX?PBu) ?LB BGLMQ?SPQ (tRCPPG@JC JGX?PBu). $J?BC %GLMQ?SPG? GLAJSBCQ @GPBQ, UFGAF CTMJTCB DPMK ? @P?LAF MD BGLMQ?SPQ.
#(!04%2 29 X 6%24%"2!4%3
813
!@>LI< 29.22 4EFP @E>OQ PELTP QEB BSLIRQFLK LC >JKFLQBP. 4EB MI>@BJBKQ LC 4BPQRAFKBP (QROQIBP) FP @ROOBKQIV PQFII AB?>QBA.
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
C?8I8:K<I@JK@:J F= -<GK@C<J
3CNRGJCQ ?PC RCRP?NMBQ. -GK@JCQQ PCNRGJCQqQL?ICQ ?LB MRFCP QOS?K?RCQqF?TC TCQRGEG?J JGK@Q ?LB, JGIC A?CAGJG?LQ, ?PC AJ?QQGDGCB ?Q RCRP?NMBQ @CA?SQC RFCW ?PC BCQACLBCB DPMK DMSP-JGK@CB ?LACQRMPQ. 3CNRGJCQ J?W CEEQ CLAJMQCB GL QFCJJQ ML J?LB. &TCL ?OS?RGA PCNRGJCQ PCRSPL RM RFC J?LB RM J?W CEEQ. 5FCW SQS?JJW PCNPMBSAC QCVS?JJW UGRF GLRCPL?J DCPRGJGX?RGML. 4MKC QNCAGCQ BGQNJ?W MTMTGTGN?PGRW, UGRF RFC CEEQ PCK?GLGLE GL RFC KMRFCPs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rumation GQ QGKGJ?P RM FG@CPL?RGML GL RF?R RFC ?LGK?J @CAMKCQ JCQQ ?ARGTC ?LB A?L EM DMP JMLE NCPGMBQ UGRFMSR C?RGLE, @SR BGDDCPQ DPMK FG@CPL?RGML GL RF?R @PSK?RGLE PCNRGJCQ ?PC LMR ?QJCCN MP JGTGLE MDD D?R PCQCPTCQ. 3?RFCP, RFCGP KCR?@MJGQK GQ QJMUCB GL PCQNMLQC RM AMJB RCKNCP?RSPCQ, ?LB RFC ?LGK?J GQ TCPW QJSEEGQF.
814
#(!04%2 29 X 6%24%"2!4%3
MFCLK@FE F= -<GK@C<J
3CNRGJCQ MPGEGL?RCB ?NNPMVGK?RCJW 300 KGJJGML WC?PQ ?EM BSPGLE RFC $?P@MLGDCPMSQ NCPGMB. 0LC MD RFC MJBCQR ILMUL ?KLGMRCQ GQ Casineria, UFGAF F?B @MRF ?KNFG@G?L ?LB PCNRGJG?L AF?P?ARCPGQRGAQ. 0LC MD RFC C?PJGCQR SLBGQNSRCB PCNRGJCQ U?Q Hylonomus. 4MML ?DRCP RFC DGPQR ?KLGMRCQ ?NNC?PCB, RFCW BGTCPECB GLRM RFPCC EPMSNQqQWL?NQGBQ, ?L?NQGBQ, ?LB BG?NQGBQqBSPGLE RFC 1CPKG?L NCPGMB. 5FC 1CPKG?L NCPGMB ?JQM Q?U ? QCAMLB K?HMP BGTCPECLAC MD BG?NQGB PCNRGJCQ GLRM ?PAFMQ?SPQ (NPCBCACQQMPQ MD APMAMBGJG?LQ ?LB BGLMQ?SPQ) ?LB JCNGBMQ?SPQ (NPCBCACQQMPQ MD QL?ICQ ?LB JGX?PBQ). 5FCQC EPMSNQ PCK?GLCB GLAMLQNGASMSQ SLRGJ RFC 5PG?QQGA NCPGMB, UFCL RFC ?PAFMQ?SPQ @CA?KC RFC BMKGL?LR RCPPCQRPG?J EPMSN BSC RM RFC CVRGLARGML MD J?PEC-@MBGCB ?L?NQGBQ ?LB QWL?NQGBQ BSPGLE RFC 1CPKG?L-5PG?QQGA CVRGLARGML. "@MSR 250 KGJJGML WC?PQ ?EM, ?PAFMQ?SPQ P?BG?RCB GLRM RFC BGLMQ?SPQ ?LB RFC NRCPMQ?SPQ. "JRFMSEF RFCW ?PC QMKCRGKCQ KGQR?ICLJW A?JJCB BGLMQ?SPQ, RFC NRCPMQ?SPQ UCPC BGQRGLAR DPMK RPSC BGLMQ?SPQ (Figure 29.23). 1RCPMQ?SPQ F?B ? LSK@CP MD ?B?NR?RGMLQ RF?R ?JJMUCB DMP DJGEFR, GLAJSBGLE FMJJMU @MLCQ (@GPBQ ?JQM CVFG@GR FMJJMU @MLCQ, ? A?QC MD AMLTCPECLR CTMJSRGML). 5FCGP UGLEQ UCPC DMPKCB @W KCK@P?LCQ MD QIGL RF?R ?RR?AFCB RM RFC JMLE, DMSPRF DGLECP MD C?AF ?PK ?LB CVRCLBCB ?JMLE RFC @MBW RM RFC JCEQ.
!@>LI< 29.23 0QBOLP>ROP, TEF@E BUFPQBA COLJ QEB I>QB 4OF>PPF@ QL QEB #OBQ>@BLRP MBOFLA (210 QL 65.5 JFIIFLK VB>OP >DL), MLPPBPPBA TFKDP ?RQ >OB KLQ ?BIFBSBA QL E>SB ?BBK @>M>?IB LC MLTBOBA CIFDEQ. )KPQB>A, QEBV J>V E>SB ?BBK >?IB QL PL>O >CQBO I>RK@EFKD COLJ @IFCCP. (@OBAFQ: ->OH 7FQQLK, $>OOBK .>FPE)
5FC BGLMQ?SPQ UCPC ? BGTCPQC EPMSN MD RCPPCQRPG?J PCNRGJCQ UGRF KMPC RF?L 1,000 QNCAGCQ GBCLRGDGCB RM B?RC. 1?JCMLRMJMEGQRQ AMLRGLSC RM BGQAMTCP LCU QNCAGCQ MD BGLMQ?SPQ. 4MKC BGLMQ?SPQ UCPC OS?BPSNCBQ (Figure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
!@>LI< 29.24 E+465;650( T>P >K >OJLOBA AFKLP>RO QE>Q IFSBA FK QEB I>QB #OBQ>@BLRP MBOFLA, 145.5 QL 65.6 JFIIFLK VB>OP >DL. (@OBAFQ: ->OF>K> 2RFW 6FII>OB>I)
%GLMQ?SPQ BMKGL?RCB RFC .CQMXMGA &P?, UFGAF U?Q ILMUL ?Q RFC t?EC MD PCNRGJCQ.u 5FC BMKGL?LAC MD BGLMQ?SPQ J?QRCB SLRGJ RFC CLB MD RFC $PCR?ACMSQ, RFC J?QR NCPGMB MD RFC .CQMXMGA &P?. 5FC $PCR?ACMSQ-
#(!04%2 29 X 6%24%"2!4%3
815
5CPRG?PW CVRGLARGML PCQSJRCB GL RFC JMQQ MD KMQR MD RFC J?PEC-@MBGCB ?LGK?JQ MD RFC .CQMXMGA &P?. #GPBQ ?PC RFC MLJW JGTGLE BCQACLB?LRQ MD MLC MD RFC K?HMP AJ?BCQ MD BGLMQ?SPQ.
7GQGR RFGQ QGRC RM QCC ? video (http://openstaxcollege.org/l/K-T_extinction) BGQASQQGLE RFC FWNMRFCQGQ RF?R ?L ?QRCPMGB A?SQCB RFC $PCR?ACMSQ-5PG?QQGA (,5) CVRGLARGML.
(F;<IE -<GK@C<J
$J?QQ 3CNRGJG? GLAJSBCQ K?LW BGTCPQC QNCAGCQ RF?R ?PC AJ?QQGDGCB GLRM DMSP JGTGLE AJ?BCQ. 5FCQC ?PC RFC 25 QNCAGCQ MD $PMAMBGJG?, 2 QNCAGCQ MD 4NFCLMBMLRG?, ?NNPMVGK?RCJW 9,200 4OS?K?R? QNCAGCQ, ?LB RFC 5CQRSBGLCQ, UGRF ?@MSR 325 QNCAGCQ. C74)4*/1/' Crocodilia (tQK?JJ JGX?PBu) ?PMQC UGRF ? BGQRGLAR JGLC?EC @W RFC KGBBJC 5PG?QQGA; CVR?LR QNCAGCQ GLAJSBC ?JJGE?RMPQ, APMAMBGJCQ, ?LB A?GK?LQ. $PMAMBGJG?LQ (Figure 29.25) JGTC RFPMSEFMSR RFC RPMNGAQ ?LB QS@RPMNGAQ MD "DPGA?, 4MSRF "KCPGA?, 4MSRFCPL 'JMPGB?, "QG?, ?LB "SQRP?JG?. 5FCW ?PC DMSLB GL DPCQFU?RCP, Q?JRU?RCP, ?LB @P?AIGQF F?@GR?RQ, QSAF ?Q PGTCPQ ?LB J?ICQ, ?LB QNCLB KMQR MD RFCGP RGKC GL U?RCP. 4MKC QNCAGCQ ?PC ?@JC RM KMTC ML J?LB BSC RM RFCGP QCKG-CPCAR NMQRSPC.
!@>LI< 29.25 #OL@LAFIF>KP, PR@E >P QEFP 3F>JBPB @OL@LAFIB (C96*6+@3<: :0(4,5:0:), MOLSFAB M>OBKQ>I @>OB CLO QEBFO LCCPMOFKD. (@OBAFQ: +BPE>S -RHRKA +>KAE>A>F)
!5.+34*439/' Sphenodontia (tUCBEC RMMRFu) ?PMQC GL RFC .CQMXMGA CP? ?LB GLAJSBCQ MLJW MLC JGTGLE ECLSQ, !<(;(9(, AMKNPGQGLE RUM QNCAGCQ RF?R ?PC DMSLB GL /CU ;C?J?LB (Figure 29.26). 5S?R?P?Q KC?QSPC SN RM 80 ACLRGKCRCPQ ?LB UCGEF ?@MSR 1 IGJMEP?K. "JRFMSEF OSGRC JGX?PB-JGIC GL EPMQQ ?NNC?P?LAC, QCTCP?J SLGOSC DC?RSPCQ MD RFC QISJJ ?LB H?UQ AJC?PJW BCDGLC RFCK ?LB BGQRGLESGQF RFC EPMSN DPMK RFC QOS?K?RCQ.
816
#(!04%2 29 X 6%24%"2!4%3
!@>LI< 29.26 4EFP QR>Q>O> COLJ .BT :B>I>KA J>V OBPBJ?IB > IFW>OA ?RQ ?BILKDP QL > AFPQFK@Q IFKB>DB, QEB 3MEBKLALKQFA>B C>JFIV. (@OBAFQ: 3FA -LPABII)
!6:'2'9' Squamata (tQA?JWu) ?PMQC GL RFC J?RC 1CPKG?L, ?LB CVR?LR QNCAGCQ GLAJSBC JGX?PBQ ?LB QL?ICQ. #MRF ?PC DMSLB ML ?JJ AMLRGLCLRQ CVACNR "LR?PARGA?. -GX?PBQ ?LB QL?ICQ ?PC KMQR AJMQCJW PCJ?RCB RM RS?R?P?Q, @MRF EPMSNQ F?TGLE CTMJTCB DPMK ? JCNGBMQ?SPG?L ?LACQRMP. 4OS?K?R? GQ RFC J?PECQR CVR?LR AJ?BC MD PCNRGJCQ (Figure 29.27). .MQR JGX?PBQ BGDDCP DPMK QL?ICQ @W F?TGLE DMSP JGK@Q, ?JRFMSEF RFCQC F?TC @CCL T?PGMSQJW JMQR MP QGELGDGA?LRJW PCBSACB GL ?R JC?QR 60 JGLC?ECQ. 4L?ICQ J?AI CWCJGBQ ?LB CVRCPL?J C?PQ, UFGAF ?PC NPCQCLR GL JGX?PBQ. -GX?PB QNCAGCQ P?LEC GL QGXC DPMK AF?KCJCMLQ ?LB ECAIMQ, UFGAF ?PC ? DCU ACLRGKCRCPQ GL JCLERF, RM RFC ,MKMBM BP?EML, UFGAF GQ ?@MSR 3 KCRCPQ GL JCLERF. .MQR JGX?PBQ ?PC A?PLGTMPMSQ, @SR QMKC J?PEC QNCAGCQ, QSAF ?Q GES?L?Q, ?PC FCP@GTMPCQ.
!@>LI< 29.27 4EFP *>@HPLKwP @E>JBIBLK (!906*,96: 1(*2:6500) ?IBKAP FK TFQE FQP PROOLRKAFKDP.
4L?ICQ ?PC RFMSEFR RM F?TC BCQACLBCB DPMK CGRFCP @SPPMUGLE JGX?PBQ MP ?OS?RGA JGX?PBQ MTCP 100 KGJJGML WC?PQ ?EM (Figure 29.28). 4L?ICQ AMKNPGQC ?@MSR 3,000 QNCAGCQ ?LB ?PC DMSLB ML CTCPW AMLRGLCLR CVACNR "LR?PARGA?. 5FCW P?LEC GL QGXC DPMK 10 ACLRGKCRCP-JMLE RFPC?B QL?ICQ RM 10 KCRCP-JMLE NWRFMLQ ?LB ?L?AMLB?Q. "JJ QL?ICQ ?PC A?PLGTMPMSQ ?LB C?R QK?JJ ?LGK?JQ, @GPBQ, CEEQ, DGQF, ?LB GLQCARQ. 5FC QL?IC @MBW DMPK GQ QM QNCAG?JGXCB RF?R, GL GRQ ECLCP?J KMPNFMJMEW, ? tQL?IC GQ ? QL?IC.u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q@MRF ?BT?LR?ECQ GL QU?JJMUGLE @GE GRCKQ.
#(!04%2 29 X 6%24%"2!4%3
817
!@>LI< 29.28 4EB D>OQBO PK>HB ?BILKDP QL QEB DBKRP !/(4567/0:, QEB JLPQ TFABIV AFPQOF?RQBA OBMQFIB DBKRP FK .LOQE !JBOF@>. (@OBAFQ: 3QBSB *ROSBQPLK)
"+89:*/3+8 5SPRJCQ ?PC KCK@CPQ MD RFC AJ?BC Testudines (tF?TGLE ? QFCJJu) (Figure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tRSPRJCu GQ QMKCRGKCQ SQCB RM BCQAPG@C MLJW RFMQC QNCAGCQ MD 5CQRSBGLCQ RF?R JGTC GL RFC QC?, UGRF RFC RCPKQ tRMPRMGQCu ?LB tRCPP?NGLu SQCB RM PCDCP RM QNCAGCQ RF?R JGTC ML J?LB ?LB GL DPCQF U?RCP, PCQNCARGTCJW.
!@>LI< 29.29 4EB !COF@>K PMROOBA QLOQLFPB (G,6*/,365, :<3*(;() IFSBP >Q QEB PLRQEBOK BADB LC QEB 3>E>O> $BPBOQ. )Q FP QEB QEFOA I>ODBPQ QLOQLFPB FK QEB TLOIA. (@OBAFQ: *FJ "LTBK)
29.5 R B@I;J
#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC CTMJSRGML?PW FGQRMPW MD @GPBQ z %CQAPG@C RFC BCPGTCB AF?P?ARCPGQRGAQ GL @GPBQ RF?R D?AGJGR?RC DJGEFR
818
#(!04%2 29 X 6%24%"2!4%3
5FC KMQR M@TGMSQ AF?P?ARCPGQRGA RF?R QCRQ @GPBQ ?N?PR DPMK MRFCP KMBCPL TCPRC@P?RCQ GQ RFC NPCQCLAC MD DC?RFCPQ, UFGAF ?PC KMBGDGCB QA?JCQ. 8FGJC TCPRC@P?RCQ JGIC @?RQ DJW UGRFMSR DC?RFCPQ, @GPBQ PCJW ML DC?RFCPQ ?LB UGLEQ, ?JMLE UGRF MRFCP KMBGDGA?RGMLQ MD @MBW QRPSARSPC ?LB NFWQGMJMEW, DMP DJGEFR.
C?8I8:K<I@JK@:J F= B@I;J
#GPBQ ?PC CLBMRFCPKGA, ?LB @CA?SQC RFCW DJW, RFCW PCOSGPC J?PEC ?KMSLRQ MD CLCPEW, LCACQQGR?RGLE ? FGEF KCR?@MJGA P?RC. -GIC K?KK?JQ, UFGAF ?PC ?JQM CLBMRFCPKGA, @GPBQ F?TC ?L GLQSJ?RGLE AMTCPGLE RF?R ICCNQ FC?R GL RFC @MBW: DC?RFCPQ. 4NCAG?JGXCB DC?RFCPQ A?JJCB down feathers ?PC CQNCAG?JJW GLQSJ?RGLE, RP?NNGLE ?GP GL QN?ACQ @CRUCCL C?AF DC?RFCP RM BCAPC?QC RFC P?RC MD FC?R JMQQ. $CPR?GL N?PRQ MD ? @GPBsQ @MBW ?PC AMTCPCB GL BMUL DC?RFCPQ, ?LB RFC @?QC MD MRFCP DC?RFCPQ F?TC ? BMULW NMPRGML, UFCPC?Q LCUJW F?RAFCB @GPBQ ?PC AMTCPCB GL BMUL. 'C?RFCPQ LMR MLJW ?AR ?Q GLQSJ?RGML @SR ?JQM ?JJMU DMP DJGEFR, CL?@JGLE RFC JGDR ?LB RFPSQR LCACQQ?PW RM @CAMKC ?GP@MPLC. 5FC DC?RFCPQ ML ? UGLE ?PC DJCVG@JC, QM RFC AMJJCARGTC DC?RFCPQ KMTC ?LB QCN?P?RC ?Q ?GP KMTCQ RFPMSEF RFCK, PCBSAGLE RFC BP?E ML RFC UGLE. Flight feathers ?PC ?QWKKCRPGA?J, UFGAF ?DDCARQ ?GPDJMU MTCP RFCK ?LB NPMTGBCQ QMKC MD RFC JGDRGLE ?LB RFPSQRGLE DMPAC PCOSGPCB DMP DJGEFR (Figure 29.30). 5UM RWNCQ MD DJGEFR DC?RFCPQ ?PC DMSLB ML RFC UGLEQ, NPGK?PW DC?RFCPQ ?LB QCAMLB?PW DC?RFCPQ. Primary feathers ?PC JMA?RCB ?R RFC RGN MD RFC UGLE ?LB NPMTGBC RFPSQR. Secondary feathers ?PC JMA?RCB AJMQCP RM RFC @MBW, ?RR?AF RM RFC DMPC?PK NMPRGML MD RFC UGLE ?LB NPMTGBC JGDR. Contour feathers ?PC RFC DC?RFCPQ DMSLB ML RFC @MBW, ?LB RFCW FCJN PCBSAC BP?E NPMBSACB @W UGLB PCQGQR?LAC BSPGLE DJGEFR. 5FCW APC?RC ? QKMMRF, ?CPMBWL?KGA QSPD?AC QM RF?R ?GP KMTCQ QKMMRFJW MTCP RFC @GPBsQ @MBW, ?JJMUGLE DMP CDDGAGCLR DJGEFR.
!@>LI< 29.30 0OFJ>OV CB>QEBOP >OB IL@>QBA >Q QEB TFKD QFM >KA MOLSFAB QEORPQ; PB@LKA>OV CB>QEBOP >OB IL@>QBA @ILPB QL QEB ?LAV >KA MOLSFAB IFCQ.
'J?NNGLE MD RFC CLRGPC UGLE MAASPQ NPGK?PGJW RFPMSEF RFC ?ARGMLQ MD RFC AFCQR KSQAJCQ, RFC NCARMP?JGQ ?LB RFC QSNP?AMP?AMGBCSQ. 5FCQC KSQAJCQ ?PC FGEFJW BCTCJMNCB GL @GPBQ ?LB ?AAMSLR DMP ? FGEFCP NCPACLR?EC MD @MBW K?QQ RF?L GL KMQR K?KK?JQ. 5FCQC ?RR?AF RM ? @J?BC-QF?NCB ICCJ, JGIC RF?R MD ? @M?R, JMA?RCB ML RFC QRCPLSK. 5FC QRCPLSK MD @GPBQ GQ J?PECP RF?L RF?R MD MRFCP TCPRC@P?RCQ, UFGAF ?AAMKKMB?RCQ RFC J?PEC KSQAJCQ PCOSGPCB RM ECLCP?RC CLMSEF SNU?PB DMPAC RM ECLCP?RC JGDR UGRF RFC DJ?NNGLE MD RFC UGLEQ. "LMRFCP QICJCR?J KMBGDGA?RGML DMSLB GL KMQR @GPBQ GQ RFC DSQGML MD RFC RUM AJ?TGAJCQ (AMJJ?P@MLCQ), DMPKGLE RFC furcula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neumatic bones ?PC @MLCQ RF?R ?PC FMJJMU, P?RFCP RF?L DGJJCB UGRF RGQQSC (Figure 29.31). 5FCW AMLR?GL ?GP QN?ACQ RF?R ?PC QMKCRGKCQ AMLLCARCB RM ?GP Q?AQ, ?LB RFCW F?TC QRPSRQ MD @MLC RM NPMTGBC QRPSARSP?J PCGLDMPACKCLR. 1LCSK?RGA @MLCQ ?PC LMR DMSLB GL ?JJ @GPBQ, ?LB RFCW ?PC KMPC CVRCLQGTC GL J?PEC
#(!04%2 29 X 6%24%"2!4%3
819
@GPBQ RF?L GL QK?JJ @GPBQ. /MR ?JJ @MLCQ MD RFC QICJCRML ?PC NLCSK?RGA, ?JRFMSEF RFC QISJJQ MD ?JKMQR ?JJ @GPBQ ?PC.
!@>LI< 29.31 ->KV ?FOAP E>SB ELIILT, MKBRJ>QF@ ?LKBP, TEF@E J>HB CIFDEQ B>PFBO.
0RFCP KMBGDGA?RGMLQ RF?R PCBSAC UCGEFR GLAJSBC RFC J?AI MD ? SPGL?PW @J?BBCP. #GPBQ NMQQCQQ ? AJM?A?, ? QRPSARSPC RF?R ?JJMUQ U?RCP RM @C PC?@QMP@CB DPMK U?QRC @?AI GLRM RFC @JMMBQRPC?K. 6PGA ?AGB GQ LMR CVNCJJCB ?Q ? JGOSGB @SR GQ AMLACLRP?RCB GLRM SP?RC Q?JRQ, UFGAF ?PC CVNCJJCB ?JMLE UGRF DCA?J K?RRCP. *L RFGQ U?W, U?RCP GQ LMR FCJB GL RFC SPGL?PW @J?BBCP, UFGAF UMSJB GLAPC?QC @MBW UCGEFR. .MQR @GPB QNCAGCQ MLJW NMQQCQQ MLC MT?PW P?RFCP RF?L RUM, DSPRFCP PCBSAGLE @MBW K?QQ. 5FC ?GP Q?AQ RF?R CVRCLB GLRM @MLCQ RM DMPK NLCSK?RGA @MLCQ ?JQM HMGL UGRF RFC JSLEQ ?LB DSLARGML GL PCQNGP?RGML. 6LJGIC K?KK?JG?L JSLEQ GL UFGAF ?GP DJMUQ GL RUM BGPCARGMLQ, ?Q GR GQ @PC?RFCB GL ?LB MSR, ?GPDJMU RFPMSEF @GPB JSLEQ RP?TCJQ GL MLC BGPCARGML (Figure 29.32). "GP Q?AQ ?JJMU DMP RFGQ SLGBGPCARGML?J ?GPDJMU, UFGAF ?JQM APC?RCQ ? APMQQ-ASPPCLR CVAF?LEC QWQRCK UGRF RFC @JMMB. *L ? APMQQ-ASPPCLR MP AMSLRCPASPPCLR QWQRCK, RFC ?GP DJMUQ GL MLC BGPCARGML ?LB RFC @JMMB DJMUQ GL RFC MNNMQGRC BGPCARGML, APC?RGLE ? TCPW CDDGAGCLR KC?LQ MD E?Q CVAF?LEC.
!@>LI< 29.32 !SF>K OBPMFO>QFLK FP >K BCCF@FBKQ PVPQBJ LC D>P BU@E>KDB TFQE >FO CILTFKD RKFAFOB@QFLK>IIV. $ROFKD FKE>I>QFLK, >FO M>PPBP COLJ QEB QO>@EB> FKQL MLPQBOFLO >FO P>@P, QEBK QEOLRDE QEB IRKDP QL >KQBOFLO >FO P>@P. 4EB >FO P>@P >OB @LKKB@QBA QL QEB ELIILT FKQBOFLO LC ?LKBP. $ROFKD BUE>I>QFLK, >FO COLJ >FO P>@P M>PPBP FKQL QEB IRKDP >KA LRQ QEB QO>@EB>. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ,. 3EV>J>I)
MFCLK@FE F= B@I;J
5FC CTMJSRGML?PW FGQRMPW MD @GPBQ GQ QRGJJ QMKCUF?R SLAJC?P. %SC RM RFC DP?EGJGRW MD @GPB @MLCQ, RFCW BM LMR DMQQGJGXC ?Q UCJJ ?Q MRFCP TCPRC@P?RCQ. #GPBQ ?PC BG?NQGBQ, KC?LGLE RFCW F?TC RUM DCLCQRP?RGMLQ MP MNCLGLEQ GL RFCGP QISJJQ. #GPBQ @CJMLE RM ? EPMSN MD BG?NQGBQ A?JJCB RFC ?PAFMQ?SPQ, UFGAF ?JQM GLAJSBCQ APMAMBGJCQ ?LB BGLMQ?SPQ. *R GQ AMKKMLJW ?AACNRCB RF?R @GPBQ CTMJTCB DPMK BGLMQ?SPQ. %GLMQ?SPQ (GLAJSBGLE @GPBQ) ?PC DSPRFCP QS@BGTGBCB GLRM RUM EPMSNQ, RFC 4?SPGQAFG? (tJGX?PB JGICu) ?LB RFC 0PLGRFGQAFG? (t@GPB JGICu). %CQNGRC RFC L?KCQ MD RFCQC EPMSNQ, GR U?Q LMR RFC @GPB-JGIC BGLMQ?SPQ RF?R E?TC PGQC RM KMBCPL @GPBQ. 3?RFCP, 4?SPGQAFG? BGTCPECB GLRM RUM EPMSNQ: 0LC GLAJSBCB RFC JMLELCAICB FCP@GTMPMSQ BGLMQ?SPQ, QSAF ?Q "N?RMQ?SPSQ. 5FC QCAMLB EPMSN, @GNCB?J NPCB?RMPQ A?JJCB theropods, GLAJSBCQ @GPBQ. 5FGQ AMSPQC MD CTMJSRGML GQ QSEECQRCB @W QGKGJ?PGRGCQ @CRUCCL RFCPMNMB DMQQGJQ ?LB @GPBQ, QNCAGDGA?JJW GL RFC QRPSARSPC MD RFC FGN ?LB UPGQR @MLCQ, ?Q UCJJ ?Q RFC NPCQCLAC MD RFC UGQF@MLC, DMPKCB @W RFC DSQGLE MD RFC AJ?TGAJCQ. 0LC GKNMPR?LR DMQQGJ MD ?L ?LGK?J GLRCPKCBG?RC RM BGLMQ?SPQ ?LB @GPBQ GQ Archaeopteryx, UFGAF GQ DPMK RFC +SP?QQGA NCPGMB (Figure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
820
#(!04%2 29 X 6%24%"2!4%3
!@>LI< 29.33 (>) A9*/(,67;,9@? IFSBA FK QEB I>QB *RO>PPF@ 0BOFLA >OLRKA 150 JFIIFLK VB>OP >DL. )Q E>A QBBQE IFHB > AFKLP>RO, ?RQ E>A (?) CIFDEQ CB>QEBOP IFHB JLABOK ?FOAP, TEF@E @>K ?B PBBK FK QEFP CLPPFI.
*R GQ QRGJJ SLAJC?P CV?ARJW FMU DJGEFR CTMJTCB GL @GPBQ. 5UM K?GL RFCMPGCQ CVGQR, RFC ?P@MPC?J (tRPCCu) FWNMRFCQGQ ?LB RFC RCPPCQRPG?J (tJ?LBu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qDC?RFCPQ MP CLBMRFCPKWqCTMJTCB GL PCQNMLQC RM QMKC MRFCP QCJCARGTC NPCQQSPC. %SPGLE RFC $PCR?ACMSQ NCPGMB, ? EPMSN ILMUL ?Q RFC Enantiornithes U?Q RFC BMKGL?LR @GPB RWNC (Figure 29.34). &L?LRGMPLGRFCQ KC?LQ tMNNMQGRC @GPBQ,u UFGAF PCDCPQ RM RFC D?AR RF?R ACPR?GL @MLCQ MD RFC DCCR ?PC HMGLCB BGDDCPCLRJW RF?L RFC U?W RFC @MLCQ ?PC HMGLCB GL KMBCPL @GPBQ. 5FCQC @GPBQ DMPKCB ?L CTMJSRGML?PW JGLC QCN?P?RC DPMK KMBCPL @GPBQ, ?LB RFCW BGB LMR QSPTGTC N?QR RFC $PCR?ACMSQ. "JMLE UGRF RFC &L?LRGMPLGRFCQ, 0PLGRFSP?C @GPBQ (RFC CTMJSRGML?PW JGLC RF?R GLAJSBCQ KMBCPL @GPBQ) UCPC ?JQM NPCQCLR GL RFC $PCR?ACMSQ. "DRCP RFC CVRGLARGML MD &L?LRGMPLGRFCQ, KMBCPL @GPBQ @CA?KC RFC BMKGL?LR @GPB, UGRF ? J?PEC P?BG?RGML MAASPPGLE BSPGLE RFC $CLMXMGA &P?. 3CDCPPCB RM ?Q Neornithes (tLCU @GPBQu), KMBCPL @GPBQ ?PC LMU AJ?QQGDGCB GLRM RUM EPMSNQ, RFC Paleognathae (tMJB H?Uu) MP P?RGRCQ, ? EPMSN MD DJGEFRJCQQ @GPBQ GLAJSBGLE MQRPGAFCQ, CKSQ, PFC?Q, ?LB IGUGQ, ?LB RFC Neognathae (tLCU H?Uu), UFGAF GLAJSBCQ ?JJ MRFCP @GPBQ.
!@>LI< 29.34 /(5>,050(6 *667,969<4 T>P > PMB@FBP LC %K>KQFLOKFQEBP QE>Q AFA KLQ PROSFSB M>PQ QEB #OBQ>@BLRP MBOFLA. (@OBAFQ: .L?R 4>JRO>)
#(!04%2 29 X 6%24%"2!4%3
821
1<K<I@E8I@8E
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
29.6 R (8DD8CJ
#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z /?KC ?LB BCQAPG@C RFC BGQRGLESGQFGLE DC?RSPCQ MD RFC RFPCC K?GL EPMSNQ MD K?KK?JQ z %CQAPG@C RFC NPMNMQCB JGLC MD BCQACLR RF?R NPMBSACB K?KK?JQ z -GQR QMKC BCPGTCB DC?RSPCQ RF?R K?W F?TC ?PGQCL GL PCQNMLQC RM K?KK?JQs LCCB DMP AMLQR?LR, FGEF-JCTCJ KCR?@MJGQK Mammals ?PC TCPRC@P?RCQ RF?R NMQQCQQ F?GP ?LB K?KK?PW EJ?LBQ. 4CTCP?J MRFCP AF?P?ARCPGQRGAQ ?PC BGQRGLARGTC RM K?KK?JQ, GLAJSBGLE ACPR?GL DC?RSPCQ MD RFC H?U, QICJCRML, GLRCESKCLR, ?LB GLRCPL?J ?L?RMKW. .MBCPL K?KK?JQ @CJMLE RM RFPCC AJ?BCQ: KMLMRPCKCQ, K?PQSNG?JQ, ?LB CSRFCPG?LQ (MP NJ?ACLR?J K?KK?JQ).
C?8I8:K<I@JK@:J F= (8DD8CJ
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sQ F?GP QR?LBQ tML CLB.u .?KK?JG?L GLRCESKCLR, MP QIGL, GLAJSBCQ QCAPCRMPW EJ?LBQ UGRF T?PGMSQ DSLARGMLQ. Sebaceous glands NPMBSAC ? JGNGB KGVRSPC A?JJCB QC@SK RF?R GQ QCAPCRCB MLRM RFC F?GP ?LB QIGL DMP U?RCP PCQGQR?LAC ?LB JS@PGA?RGML. 4C@?ACMSQ EJ?LBQ ?PC JMA?RCB MTCP KMQR MD RFC @MBW. Eccrine glands NPMBSAC QUC?R, MP NCPQNGP?RGML, UFGAF GQ K?GLJW AMKNMQCB MD U?RCP. *L KMQR K?KK?JQ, CAAPGLC EJ?LBQ ?PC JGKGRCB RM ACPR?GL ?PC?Q MD RFC @MBW, ?LB QMKC K?KK?JQ BM LMR NMQQCQQ RFCK ?R ?JJ. )MUCTCP, GL NPGK?RCQ, CQNCAG?JJW FSK?LQ, QUC?R DGESPCQ NPMKGLCLRJW GL RFCPKMPCESJ?RGML, PCESJ?RGLE RFC @MBW RFPMSEF CT?NMP?RGTC
822
#(!04%2 29 X 6%24%"2!4%3
AMMJGLE. 4UC?R EJ?LBQ ?PC JMA?RCB MTCP KMQR MD RFC @MBW QSPD?AC GL NPGK?RCQ. Apocrine glands, MP QACLR EJ?LBQ, QCAPCRC QS@QR?LACQ RF?R ?PC SQCB DMP AFCKGA?J AMKKSLGA?RGML, QSAF ?Q GL QISLIQ. Mammary glands NPMBSAC KGJI RF?R GQ SQCB RM DCCB LCU@MPLQ. 8FGJC K?JC KMLMRPCKCQ ?LB CSRFCPG?LQ NMQQCQQ K?KK?PW EJ?LBQ, K?JC K?PQSNG?JQ BM LMR. .?KK?PW EJ?LBQ JGICJW ?PC KMBGDGCB QC@?ACMSQ MP CAAPGLC EJ?LBQ, @SR RFCGP CTMJSRGML?PW MPGEGL GQ LMR CLRGPCJW AJC?P. 5FC QICJCR?J QWQRCK MD K?KK?JQ NMQQCQQCQ K?LW SLGOSC DC?RSPCQ. 5FC JMUCP H?U MD K?KK?JQ AMLQGQRQ MD MLJW MLC @MLC, RFC dentary. 5FC H?UQ MD MRFCP TCPRC@P?RCQ ?PC AMKNMQCB MD KMPC RF?L MLC @MLC. *L K?KK?JQ, RFC BCLR?PW @MLC HMGLQ RFC QISJJ ?R RFC QOS?KMQ?J @MLC, UFGJC GL MRFCP TCPRC@P?RCQ, RFC OS?BP?RC @MLC MD RFC H?U HMGLQ UGRF RFC ?PRGASJ?P @MLC MD RFC QISJJ. 5FCQC @MLCQ ?PC NPCQCLR GL K?KK?JQ, @SR RFCW F?TC @CCL KMBGDGCB RM DSLARGML GL FC?PGLE ?LB DMPK @MLCQ GL RFC KGBBJC C?P (Figure 29.35). 0RFCP TCPRC@P?RCQ NMQQCQQ MLJW MLC KGBBJC C?P @MLC, RFC QR?NCQ. .?KK?JQ F?TC RFPCC: RFC K?JJCSQ, GLASQ, ?LB QR?NCQ. 5FC K?JJCSQ MPGEGL?RCB DPMK RFC ?PRGASJ?P @MLC, UFCPC?Q RFC GLASQ MPGEGL?RCB DPMK RFC OS?BP?RC @MLC. 5FGQ ?PP?LECKCLR MD H?U ?LB C?P @MLCQ ?GBQ GL BGQRGLESGQFGLE DMQQGJ K?KK?JQ DPMK DMQQGJQ MD MRFCP QWL?NQGBQ.
!@>LI< 29.35 "LKBP LC QEB J>JJ>IF>K FKKBO B>O >OB JLAFCFBA COLJ ?LKBP LC QEB G>T >KA PHRII. (@OBAFQ: .#))
5FC ?BBSARMP KSQAJC RF?R AJMQCQ RFC H?U GQ AMKNMQCB MD RUM KSQAJCQ GL K?KK?JQ: RFC RCKNMP?JGQ ?LB RFC K?QQCRCP. 5FCQC ?JJMU QGBC-RM-QGBC KMTCKCLR MD RFC H?U, K?IGLE AFCUGLE NMQQG@JC, UFGAF GQ SLGOSC RM K?KK?JQ. .MQR K?KK?JQ F?TC heterodont teeth, KC?LGLE RF?R RFCW F?TC BGDDCPCLR RWNCQ ?LB QF?NCQ MD RCCRF P?RFCP RF?L HSQR MLC RWNC ?LB QF?NC MD RMMRF. .MQR K?KK?JQ ?PC diphyodonts, KC?LGLE RF?R RFCW F?TC RUM QCRQ MD RCCRF GL RFCGP JGDCRGKC: BCAGBSMSQ MP t@?@Wu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
MFCLK@FE F= (8DD8CJ
.?KK?JQ ?PC QWL?NQGBQ, KC?LGLE RFCW F?TC ? QGLEJC MNCLGLE GL RFC QISJJ. 5FCW ?PC RFC MLJW JGTGLE QWL?NQGBQ, ?Q C?PJGCP DMPKQ @CA?KC CVRGLAR @W RFC +SP?QQGA NCPGMB. 5FC C?PJW LML-K?KK?JG?L QWL?NQGBQ A?L @C BGTGBCB GLRM RUM EPMSNQ, RFC NCJWAMQ?SPQ ?LB RFC RFCP?NQGBQ. 8GRFGL RFC RFCP?NQGBQ, ? EPMSN A?JJCB RFC AWLMBMLRQ ?PC RFMSEFR RM @C RFC ?LACQRMPQ MD K?KK?JQ (Figure 29.36).
#(!04%2 29 X 6%24%"2!4%3
823
!@>LI< 29.36 #VKLALKQP, TEF@E CFOPQ >MMB>OBA FK QEB ,>QB 0BOJF>K MBOFLA 260 JFIIFLK VB>OP >DL, >OB QELRDEQ QL ?B QEB >K@BPQLOP LC JLABOK J>JJ>IP. (@OBAFQ: .L?R 4>JRO>)
" ICW AF?P?ARCPGQRGA MD QWL?NQGBQ GQ CLBMRFCPKW, P?RFCP RF?L RFC CARMRFCPKW QCCL GL KMQR MRFCP TCPRC@P?RCQ. 5FC GLAPC?QCB KCR?@MJGA P?RC PCOSGPCB RM GLRCPL?JJW KMBGDW @MBW RCKNCP?RSPC UCLR F?LB GL F?LB UGRF AF?LECQ RM ACPR?GL QICJCR?J QRPSARSPCQ. 5FC J?RCP QWL?NQGBQ, UFGAF F?B KMPC CTMJTCB AF?P?ARCPGQRGAQ SLGOSC RM K?KK?JQ, NMQQCQQ AFCCIQ DMP FMJBGLE DMMB ?LB FCRCPMBMLR RCCRF, UFGAF ?PC QNCAG?JGXCB DMP AFCUGLE, KCAF?LGA?JJW @PC?IGLE BMUL DMMB RM QNCCB BGECQRGML ?LB PCJC?QGLE RFC CLCPEW LCCBCB RM NPMBSAC FC?R. $FCUGLE ?JQM PCOSGPCQ RFC ?@GJGRW RM AFCU ?LB @PC?RFC ?R RFC Q?KC RGKC, UFGAF GQ D?AGJGR?RCB @W RFC NPCQCLAC MD ? QCAMLB?PW N?J?RC. " QCAMLB?PW N?J?RC QCN?P?RCQ RFC ?PC? MD RFC KMSRF UFCPC AFCUGLE MAASPQ DPMK RFC ?PC? ?@MTC UFCPC PCQNGP?RGML MAASPQ, ?JJMUGLE @PC?RFGLE RM NPMACCB SLGLRCPPSNRCB BSPGLE AFCUGLE. " QCAMLB?PW N?J?RC GQ LMR DMSLB GL NCJWAMQ?SPQ @SR GQ NPCQCLR GL AWLMBMLRQ ?LB K?KK?JQ. 5FC H?U@MLC ?JQM QFMUQ AF?LECQ DPMK C?PJW QWL?NQGBQ RM J?RCP MLCQ. 5FC XWEMK?RGA ?PAF, MP AFCCI@MLC, GQ NPCQCLR GL K?KK?JQ ?LB ?BT?LACB RFCP?NQGBQ QSAF ?Q AWLMBMLRQ, @SR GQ LMR NPCQCLR GL NCJWAMQ?SPQ. 5FC NPCQCLAC MD RFC XWEMK?RGA ?PAF QSEECQRQ RFC NPCQCLAC MD RFC K?QQCRCP KSQAJC, UFGAF AJMQCQ RFC H?U ?LB DSLARGMLQ GL AFCUGLE. *L RFC ?NNCLBGASJ?P QICJCRML, RFC QFMSJBCP EGPBJC MD RFCPG?L K?KK?JQ GQ KMBGDGCB DPMK RF?R MD MRFCP TCPRC@P?RCQ GL RF?R GR BMCQ LMR NMQQCQQ ? NPMAMP?AMGB @MLC MP ?L GLRCPAJ?TGAJC, ?LB RFC QA?NSJ? GQ RFC BMKGL?LR @MLC. .?KK?JQ CTMJTCB DPMK RFCP?NQGBQ GL RFC J?RC 5PG?QQGA NCPGMB, ?Q RFC C?PJGCQR ILMUL K?KK?J DMQQGJQ ?PC DPMK RFC C?PJW +SP?QQGA NCPGMB, QMKC 205 KGJJGML WC?PQ ?EM. &?PJW K?KK?JQ UCPC QK?JJ, ?@MSR RFC QGXC MD ? QK?JJ PMBCLR. .?KK?JQ DGPQR @CE?L RM BGTCPQGDW GL RFC .CQMXMGA &P?, DPMK RFC +SP?QQGA RM RFC $PCR?ACMSQ NCPGMBQ, ?JRFMSEF KMQR MD RFCQC K?KK?JQ UCPC CVRGLAR @W RFC CLB MD RFC .CQMXMGA. %SPGLE RFC $PCR?ACMSQ NCPGMB, ?LMRFCP P?BG?RGML MD K?KK?JQ @CE?L ?LB AMLRGLSCB RFPMSEF RFC $CLMXMGA &P?, ?@MSR 65 KGJJGML WC?PQ ?EM.
'@M@E> (8DD8CJ
5FC CSRFCPG?LQ, MP NJ?ACLR?J K?KK?JQ, ?LB RFC K?PQSNG?JQ RMECRFCP AMKNPGQC RFC AJ?BC MD RFCPG?L K?KK?JQ. .MLMRPCKCQ, MP KCR?RFCPG?LQ, DMPK RFCGP QGQRCP AJ?BC. 5FCPC ?PC RFPCC JGTGLE QNCAGCQ MD monotremes: RFC NJ?RWNSQ ?LB RUM QNCAGCQ MD CAFGBL?Q, MP QNGLW ?LRC?RCPQ. 5FC JC?RFCPW-@C?ICB NJ?RWNSQ @CJMLEQ RM RFC D?KGJW Ornithorhynchidae (t@GPB @C?Iu), UFCPC?Q CAFGBL?Q @CJMLE RM RFC D?KGJW Tachyglossidae (tQRGAIW RMLESCu) (Figure 29.37). 5FC NJ?RWNSQ ?LB MLC QNCAGCQ MD CAFGBL? ?PC DMSLB GL "SQRP?JG?, ?LB RFC MRFCP QNCAGCQ MD CAFGBL? GQ DMSLB GL /CU (SGLC?. .MLMRPCKCQ ?PC SLGOSC ?KMLE K?KK?JQ ?Q RFCW J?W CEEQ, P?RFCP RF?L EGTGLE @GPRF RM JGTC WMSLE. 5FC QFCJJQ MD RFCGP CEEQ ?PC LMR JGIC RFC F?PB QFCJJQ MD @GPBQ, @SR ?PC ? JC?RFCPW QFCJJ, QGKGJ?P RM RFC QFCJJQ MD PCNRGJC CEEQ. .MLMRPCKCQ F?TC LM RCCRF.
(@) (?) !@>LI< 29.37 (>) 4EB MI>QVMRP, > JLKLQOBJB, MLPPBPPBP > IB>QEBOV ?B>H >KA I>VP BDDP O>QEBO QE>K DFSFKD ?FOQE QL IFSB VLRKD. (?) 4EB B@EFAK> FP >KLQEBO JLKLQOBJB. (@OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V ">OOV 4ELJ>P)
824
#(!04%2 29 X 6%24%"2!4%3
Marsupials ?PC DMSLB NPGK?PGJW GL "SQRP?JG?, RFMSEF RFC MNMQQSK GQ DMSLB GL /MPRF "KCPGA?. "SQRP?JG?L K?PQSNG?JQ GLAJSBC RFC I?LE?PMM, IM?J?, @?LBGAMMR, 5?QK?LG?L BCTGJ (Figure 29.38), ?LB QCTCP?J MRFCP QNCAGCQ. .MQR QNCAGCQ MD K?PQSNG?JQ NMQQCQQ ? NMSAF GL UFGAF RFC TCPW NPCK?RSPC WMSLE PCQGBC ?DRCP @GPRF, PCACGTGLE KGJI ?LB AMLRGLSGLE RM BCTCJMN. .?PQSNG?JQ BGDDCP DPMK CSRFCPG?LQ GL RF?R RFCPC GQ ? JCQQ AMKNJCV NJ?ACLR?J AMLLCARGML: 5FC WMSLE ?PC @MPL ?R ?L CVRPCKCJW C?PJW ?EC ?LB J?RAF MLRM RFC LGNNJC UGRFGL RFC NMSAF.
!@>LI< 29.38 4EB 4>PJ>KF>K ABSFI FP LKB LC PBSBO>I J>OPRMF>IP K>QFSB QL !RPQO>IF>. (@OBAFQ: 7>VKB -@,B>K)
&SRFCPG?LQ ?PC RFC KMQR UGBCQNPC?B MD RFC K?KK?JQ, MAASPPGLE RFPMSEFMSR RFC UMPJB. 5FCPC ?PC 18 RM 20 MPBCPQ MD NJ?ACLR?J K?KK?JQ. 4MKC CV?KNJCQ ?PC *LQCARGTMP?, RFC GLQCAR C?RCPQ; &BCLR?R?, RFC RMMRFJCQQ ?LRC?RCPQ; 3MBCLRG?, RFC PMBCLRQ; $CR?AC?, RFC ?OS?RGA K?KK?JQ GLAJSBGLE UF?JCQ; $?PLGTMP?, A?PLGTMPMSQ K?KK?JQ GLAJSBGLE BMEQ, A?RQ, ?LB @C?PQ; ?LB 1PGK?RCQ, UFGAF GLAJSBCQ FSK?LQ. Eutherian mammals ?PC QMKCRGKCQ A?JJCB NJ?ACLR?J K?KK?JQ @CA?SQC ?JJ QNCAGCQ NMQQCQQ ? AMKNJCV NJ?ACLR? RF?R AMLLCARQ ? DCRSQ RM RFC KMRFCP, ?JJMUGLE DMP E?Q, DJSGB, ?LB LSRPGCLR CVAF?LEC. 8FGJC MRFCP K?KK?JQ NMQQCQQ ? JCQQ AMKNJCV NJ?ACLR? MP @PGCDJW F?TC ? NJ?ACLR?, ?JJ CSRFCPG?LQ NMQQCQQ ? AMKNJCV NJ?ACLR? BSPGLE ECQR?RGML.
29.7 R /?< MFCLK@FE F= +I@D8K<J

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC BCPGTCB DC?RSPCQ RF?R BGQRGLESGQF NPGK?RCQ DPMK MRFCP ?LGK?JQ z &VNJ?GL UFW QAGCLRGQRQ ?PC F?TGLE BGDDGASJRW BCRCPKGLGLE RFC RPSC JGLCQ MD BCQACLR GL FMKGLGBQ 0PBCP Primates MD AJ?QQ .?KK?JG? GLAJSBCQ JCKSPQ, R?PQGCPQ, KMLICWQ, ?NCQ, ?LB FSK?LQ. /ML-FSK?L NPGK?RCQ JGTC NPGK?PGJW GL RFC RPMNGA?J MP QS@RPMNGA?J PCEGMLQ MD 4MSRF "KCPGA?, "DPGA?, ?LB "QG?. 5FCW P?LEC GL QGXC DPMK RFC KMSQC JCKSP ?R 30 EP?KQ (1 MSLAC) RM RFC KMSLR?GL EMPGJJ? ?R 200 IGJMEP?KQ (441 NMSLBQ). 5FC AF?P?ARCPGQRGAQ ?LB CTMJSRGML MD NPGK?RCQ GQ MD N?PRGASJ?P GLRCPCQR RM SQ ?Q GR ?JJMUQ SQ RM SLBCPQR?LB RFC CTMJSRGML MD MSP MUL QNCAGCQ.
C?8I8:K<I@JK@:J F= +I@D8K<J
"JJ NPGK?RC QNCAGCQ NMQQCQQ ?B?NR?RGMLQ DMP AJGK@GLE RPCCQ, ?Q RFCW ?JJ BCQACLBCB DPMK RPCC-BUCJJCPQ. 5FGQ ?P@MPC?J FCPGR?EC MD NPGK?RCQ F?Q PCQSJRCB GL F?LBQ ?LB DCCR RF?R ?PC ?B?NRCB DMP brachiation, MP AJGK@GLE ?LB QUGLEGLE RFPMSEF RPCCQ. 5FCQC ?B?NR?RGMLQ GLAJSBC, @SR ?PC LMR JGKGRCB RM: 1) ? PMR?RGLE QFMSJBCP HMGLR, 2) ? @GE RMC RF?R GQ UGBCJW QCN?P?RCB DPMK RFC MRFCP RMCQ ?LB RFSK@Q, UFGAF ?PC UGBCJW QCN?P?RCB DPMK DGLECPQ (CVACNR FSK?LQ), UFGAF ?JJMU DMP EPGNNGLE @P?LAFCQ, 3) stereoscopic vision, RUM MTCPJ?NNGLE DGCJBQ MD TGQGML DPMK RFC CWCQ, UFGAF ?JJMUQ DMP RFC NCPACNRGML MD BCNRF ?LB E?SEGLE BGQR?LAC. 0RFCP AF?P?ARCPGQRGAQ MD NPGK?RCQ ?PC @P?GLQ RF?R ?PC J?PECP RF?L RFMQC MD KMQR MRFCP K?KK?JQ, AJ?UQ RF?R F?TC @CCL KMBGDGCB GLRM DJ?RRCLCB L?GJQ, RWNGA?JJW MLJW MLC MDDQNPGLE NCP NPCEL?LAW, ?LB ? RPCLB RMU?PB FMJBGLE RFC @MBW SNPGEFR. 0PBCP 1PGK?RCQ GQ BGTGBCB GLRM RUM EPMSNQ: NPMQGKG?LQ ?LB ?LRFPMNMGBQ. Prosimians GLAJSBC RFC @SQF @?@GCQ MD "DPGA?, RFC JCKSPQ MD .?B?E?QA?P, ?LB RFC JMPGQCQ, NMRRMQ, ?LB R?PQGCPQ MD 4MSRFC?QR "QG?. Anthropoids GLAJSBC KMLICWQ, ?NCQ, ?LB FSK?LQ. *L ECLCP?J, NPMQGKG?LQ RCLB RM @C LMARSPL?J (GL AMLRP?QR RM BGSPL?J ?LRFPMNMGBQ) ?LB CVFG@GR ? QK?JJCP QGXC ?LB QK?JJCP @P?GL RF?L ?LRFPMNMGBQ.
#(!04%2 29 X 6%24%"2!4%3
825
MFCLK@FE F= +I@D8K<J
5FC DGPQR NPGK?RC-JGIC K?KK?JQ ?PC PCDCPPCB RM ?Q NPMRM-NPGK?RCQ. 5FCW UCPC PMSEFJW QGKGJ?P RM QOSGPPCJQ ?LB RPCC QFPCUQ GL QGXC ?LB ?NNC?P?LAC. 5FC CVGQRGLE DMQQGJ CTGBCLAC (KMQRJW DPMK /MPRF "DPGA?) GQ TCPW DP?EKCLRCB. 5FCQC NPMRM-NPGK?RCQ PCK?GL J?PECJW KWQRCPGMSQ APC?RSPCQ SLRGJ KMPC DMQQGJ CTGBCLAC @CAMKCQ ?T?GJ?@JC. 5FC MJBCQR ILMUL NPGK?RC-JGIC K?KK?JQ UGRF ? PCJ?RGTCJW PM@SQR DMQQGJ PCAMPB GQ Plesiadapis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latyrrhiniq? PCDCPCLAC RM RFCGP @PM?B LMQCQ (Figure 29.39). 0JB 8MPJB KMLICWQ ?PC A?JJCB Catarrhiniq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
!@>LI< 29.39 4EB ELTIBO JLKHBV FP K>QFSB QL #BKQO>I >KA 3LRQE !JBOF@>. )Q J>HBP > @>II QE>Q PLRKAP IFHB > IFLK OL>OFKD. (@OBAFQ: 8>SF 4>IIBA>)
"NCQ CTMJTCB DPMK RFC A?R?PPFGLCQ GL "DPGA? KGBU?W RFPMSEF RFC $CLMXMGA, ?NNPMVGK?RCJW 25 KGJJGML WC?PQ ?EM. "NCQ ?PC ECLCP?JJW J?PECP RF?L KMLICWQ ?LB RFCW BM LMR NMQQCQQ ? R?GJ. "JJ ?NCQ ?PC A?N?@JC MD KMTGLE RFPMSEF RPCCQ, ?JRFMSEF K?LW QNCAGCQ QNCLB KMQR RFCGP RGKC ML RFC EPMSLB. "NCQ ?PC KMPC GLRCJJGECLR RF?L KMLICWQ, ?LB RFCW F?TC PCJ?RGTCJW J?PECP @P?GLQ NPMNMPRGML?RC RM @MBW QGXC. 5FC ?NCQ ?PC BGTGBCB GLRM RUM EPMSNQ. 5FC JCQQCP ?NCQ AMKNPGQC RFC D?KGJW Hylobatidae, GLAJSBGLE EG@@MLQ ?LB QG?K?LEQ. 5FC EPC?R ?NCQ GLAJSBC RFC ECLCP? Pan (AFGKN?LXCCQ ?LB @MLM@MQ) (Figure 29.40a), Gorilla (EMPGJJ?Q), Pongo (MP?LESR?LQ), ?LB Homo (FSK?LQ) (Figure 29.40b). 5FC TCPW ?P@MPC?J EG@@MLQ ?PC QK?JJCP RF?L RFC EPC?R ?NCQ; RFCW F?TC JMU QCVS?J BGKMPNFGQK (RF?R GQ, RFC ECLBCPQ ?PC LMR K?PICBJW BGDDCPCLR GL QGXC); ?LB RFCW F?TC PCJ?RGTCJW JMLECP ?PKQ SQCB DMP QUGLEGLE RFPMSEF RPCCQ.
826
#(!04%2 29 X 6%24%"2!4%3
!@>LI< 29.40 4EB (>) @EFJM>KWBB FP LKB LC QEB DOB>Q >MBP. )Q MLPPBPPBP > OBI>QFSBIV I>ODB ?O>FK >KA E>P KL Q>FI. (?) !II DOB>Q >MBP E>SB > PFJFI>O PHBIBQ>I PQOR@QROB. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V !>OLK ,LD>K; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V 4FJ 6F@HBOP)
#LD8E MFCLK@FE
5FC D?KGJW )MKGLGB?C MD MPBCP 1PGK?RCQ GLAJSBCQ RFC hominoids: RFC EPC?R ?NCQ (Figure 29.41). &TGBCLAC DPMK RFC DMQQGJ PCAMPB ?LB DPMK ? AMKN?PGQML MD FSK?L ?LB AFGKN?LXCC %/" QSEECQRQ RF?R FSK?LQ ?LB AFGKN?LXCCQ BGTCPECB DPMK ? AMKKML FMKGLMGB ?LACQRMP ?NNPMVGK?RCJW 6 KGJJGML WC?PQ ?EM. 4CTCP?J QNCAGCQ CTMJTCB DPMK RFC CTMJSRGML?PW @P?LAF RF?R GLAJSBCQ FSK?LQ, ?JRFMSEF MSP QNCAGCQ GQ RFC MLJW QSPTGTGLE KCK@CP. 5FC RCPK hominin GQ SQCB RM PCDCP RM RFMQC QNCAGCQ RF?R CTMJTCB ?DRCP RFGQ QNJGR MD RFC NPGK?RC JGLC, RFCPC@W BCQGEL?RGLE QNCAGCQ RF?R ?PC KMPC AJMQCJW PCJ?RCB RM FSK?LQ RF?L RM AFGKN?LXCCQ. )MKGLGLQ UCPC NPCBMKGL?LRJW @GNCB?J ?LB GLAJSBC RFMQC EPMSNQ RF?R JGICJW E?TC PGQC RM MSP QNCAGCQqGLAJSBGLE A<:;9(3670;/,*<:, H646 /()030:, ?LB H646 ,9,*;<:q?LB RFMQC LML-?LACQRP?J EPMSNQ RF?R A?L @C AMLQGBCPCB tAMSQGLQu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
#(!04%2 29 X 6%24%"2!4%3
827
!@>LI< 29.41 4EFP @E>OQ PELTP QEB BSLIRQFLK LC JLABOK ERJ>KP.
$+7> E'71> H42/3/38 5FPCC QNCAGCQ MD TCPW C?PJW FMKGLGBQ F?TC K?BC LCUQ GL RFC N?QR DCU WC?PQ. 5FC MJBCQR MD RFCQC, (/,3(5;/967<: ;*/(+,5:0:, F?Q @CCL B?RCB RM LC?PJW 7 KGJJGML WC?PQ ?EM. 5FCPC GQ ? QGLEJC QNCAGKCL MD RFGQ ECLSQ, ? QISJJ RF?R U?Q ? QSPD?AC DGLB GL $F?B. 5FC DMQQGJ, GLDMPK?JJW A?JJCB t5MSK?G,u GQ ? KMQ?GA MD NPGKGRGTC ?LB CTMJTCB AF?P?ARCPGQRGAQ, ?LB GR GQ SLAJC?P FMU RFGQ DMQQGJ DGRQ UGRF RFC NGARSPC EGTCL @W KMJCASJ?P B?R?, L?KCJW RF?R RFC JGLC JC?BGLE RM KMBCPL FSK?LQ ?LB KMBCPL AFGKN?LXCCQ ?NN?PCLRJW @GDSPA?RCB ?@MSR 6 KGJJGML WC?PQ ?EM. *R GQ LMR RFMSEFR ?R RFGQ RGKC RF?R RFGQ QNCAGCQ U?Q ?L ?LACQRMP MD KMBCPL FSK?LQ. " QCAMLB, WMSLECP QNCAGCQ, O996905 ;<.,5,5:0:, GQ ?JQM ? PCJ?RGTCJW PCACLR BGQAMTCPW, DMSLB GL 2000. 5FCPC ?PC QCTCP?J QNCAGKCLQ MD O996905. *R GQ LMR ILMUL UFCRFCP O996905 U?Q ? FSK?L ?LACQRMP, @SR RFGQ NMQQG@GJGRW F?Q LMR @CCL PSJCB MSR. 4MKC DC?RSPCQ MD O996905 ?PC KMPC QGKGJ?P RM RFMQC MD KMBCPL FSK?LQ RF?L ?PC RFC ?SQRP?JMNGRFQ, ?JRFMSEF O996905 GQ KSAF MJBCP. " RFGPB ECLSQ, A9+070;/,*<:, U?Q BGQAMTCPCB GL RFC 1990Q, ?LB RFC QAGCLRGQRQ UFM BGQAMTCPCB RFC DGPQR DMQQGJ DMSLB RF?R QMKC MRFCP QAGCLRGQRQ BGB LMR @CJGCTC RFC MPE?LGQK RM @C ? @GNCB (RFSQ, GR UMSJB LMR @C AMLQGBCPCB ? FMKGLGB). *L RFC GLRCPTCLGLE WC?PQ, QCTCP?J KMPC QNCAGKCLQ MD A9+070;/,*<:, AJ?QQGDGCB ?Q RUM BGDDCPCLR QNCAGCQ, BCKMLQRP?RCB RF?R RFC MPE?LGQK U?Q @GNCB?J. "E?GL, RFC QR?RSQ MD RFGQ ECLSQ ?Q ? FSK?L ?LACQRMP GQ SLACPR?GL. E'71> H42/3/38: G+3:8 A:897'145/9.+):8 Australopithecus (tQMSRFCPL ?NCu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
828
#(!04%2 29 X 6%24%"2!4%3
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igure 29.42) F?B QK?JJCP A?LGLCQ ?LB KMJ?PQ AMKN?PCB RM ?NCQ, @SR RFCQC UCPC J?PECP RF?L RFMQC MD KMBCPL FSK?LQ. *RQ @P?GL QGXC U?Q 380p450 AS@GA ACLRGKCRCPQ, ?NNPMVGK?RCJW RFC QGXC MD ? KMBCPL AFGKN?LXCC @P?GL. *R ?JQM F?B prognathic jaws, UFGAF GQ ? PCJ?RGTCJW JMLECP H?U RF?L RF?R MD KMBCPL FSK?LQ. *L RFC KGB-1970Q, RFC DMQQGJ MD ?L ?BSJR DCK?JC A. (-(9,5:0: U?Q DMSLB GL RFC "D?P PCEGML MD &RFGMNG? ?LB B?RCB RM 3.24 KGJJGML WC?PQ ?EM (Figure 29.43). 5FC DMQQGJ, UFGAF GQ GLDMPK?JJW A?JJCB t-SAW,u GQ QGELGDGA?LR @CA?SQC GR U?Q RFC KMQR AMKNJCRC ?SQRP?JMNGRF DMQQGJ DMSLB, UGRF 40 NCPACLR MD RFC QICJCRML PCAMTCPCB.
!@>LI< 29.42 4EB PHRII LC (>) A<:;9(3670;/,*<: (-(9,5:0:, >K B>OIV ELJFKFA QE>Q IFSBA ?BQTBBK QTL >KA QEOBB JFIIFLK VB>OP >DL, OBPBJ?IBA QE>Q LC (?) JLABOK ERJ>KP ?RQ T>P PJ>IIBO TFQE > PILMBA CLOBEB>A >KA MOLJFKBKQ G>T.
#(!04%2 29 X 6%24%"2!4%3
829
!@>LI< 29.43 4EFP >ARIQ CBJ>IB A<:;9(3670;/,*<: (-(9,5:0: PHBIBQLK, KF@HK>JBA ,R@V, T>P AFP@LSBOBA FK QEB JFA 1970P. (@OBAFQ: x120y/7FHFJBAF> #LJJLKP)
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p750 AS@GA ACLRGKCRCPQ. )MUCTCP, H. /()030: PCR?GLCB QMKC DC?RSPCQ MD MJBCP FMKGLGL QNCAGCQ, QSAF ?Q JMLE ?PKQ. 5FC L?KC H. /()030: KC?LQ tF?LBW K?L,u UFGAF GQ ? PCDCPCLAC RM RFC QRMLC RMMJQ RF?R F?TC @CCL DMSLB UGRF GRQ PCK?GLQ.
830
#(!04%2 29 X 6%24%"2!4%3
7GQGR RFGQ site (http://openstaxcollege.org/l/diet_detective) DMP ? TGBCM ?@MSR 4KGRFQMLG?L N?JCMLRMJMEGQR #PG?L? 1M@GLCP CVNJ?GLGLE RFC JGLI @CRUCCL FMKGLGL C?RGLE MD KC?R ?LB CTMJSRGML?PW RPCLBQ. H. ,9,*;<: ?NNC?PCB ?NNPMVGK?RCJW 1.8 KGJJGML WC?PQ ?EM (Figure 29.44). *R GQ @CJGCTCB RM F?TC MPGEGL?RCB GL &?QR "DPGA? ?LB U?Q RFC DGPQR FMKGLGL QNCAGCQ RM KGEP?RC MSR MD "DPGA?. 'MQQGJQ MD H. ,9,*;<: F?TC @CCL DMSLB GL *LBG?, $FGL?, +?T?, ?LB &SPMNC, ?LB UCPC ILMUL GL RFC N?QR ?Q t+?T? .?Lu MP t1CIGLE .?L.u H. ,9,*;<: F?B ? LSK@CP MD DC?RSPCQ RF?R UCPC KMPC QGKGJ?P RM KMBCPL FSK?LQ RF?L RFMQC MD H. /()030:. H. ,9,*;<: U?Q J?PECP GL QGXC RF?L C?PJGCP FMKGLGLQ, PC?AFGLE FCGEFRQ SN RM 1.85 KCRCPQ ?LB UCGEFGLE SN RM 65 IGJMEP?KQ, UFGAF ?PC QGXCQ QGKGJ?P RM RFMQC MD KMBCPL FSK?LQ. *RQ BCEPCC MD QCVS?J BGKMPNFGQK U?Q JCQQ RF?L C?PJGCP QNCAGCQ, UGRF K?JCQ @CGLE 20 RM 30 NCPACLR J?PECP RF?L DCK?JCQ, UFGAF GQ AJMQC RM RFC QGXC BGDDCPCLAC QCCL GL MSP QNCAGCQ. H. ,9,*;<: F?B ? J?PECP @P?GL RF?L C?PJGCP QNCAGCQ ?R 775p1,100 AS@GA ACLRGKCRCPQ, UFGAF AMKN?PCQ RM RFC 1,130p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
!@>LI< 29.44 H646 ,9,*;<: E>A > MOLJFKBKQ ?OLT >KA > KLPB QE>Q MLFKQBA ALTKT>OA O>QEBO QE>K CLOT>OA.
H:2'38: H424 8'5/+38 " LSK@CP MD QNCAGCQ, QMKCRGKCQ A?JJCB ?PAF?GA H646 :(70,5:, ?NN?PCLRJW CTMJTCB DPMK H. ,9,*;<: QR?PRGLE ?@MSR 500,000 WC?PQ ?EM. 5FCQC QNCAGCQ GLAJSBC H646 /,0+,3),9.,5:0:, H646 9/6+,:0,5:0:, ?LB H646 5,(5+,9;/(3,5:0:. 5FCQC ?PAF?GA H. :(70,5: F?B ? @P?GL QGXC QGKGJ?P RM RF?R MD KMBCPL FSK?LQ, ?TCP?EGLE 1,200p1,400 AS@GA ACLRGKCRCPQ. 5FCW BGDDCPCB DPMK KMBCPL FSK?LQ @W F?TGLE ? RFGAI QISJJ, ? NPMKGLCLR @PMU PGBEC, ?LB ? PCACBGLE AFGL. 4MKC MD RFCQC QNCAGCQ QSPTGTCB SLRGJ 30,000p10,000 WC?PQ ?EM, MTCPJ?NNGLE UGRF KMBCPL FSK?LQ (Figure 29.45).
#(!04%2 29 X 6%24%"2!4%3
831
!@>LI< 29.45 4EB H646 5,(5+,9;/(3,5:0: RPBA QLLIP >KA J>V E>SB TLOK @ILQEFKD.
5FCPC GQ AMLQGBCP?@JC BC@?RC ?@MSR RFC MPGEGLQ MD ?L?RMKGA?JJW KMBCPL FSK?LQ MP Homo sapiens sapiens. "Q BGQASQQCB C?PJGCP, H. ,9,*;<: KGEP?RCB MSR MD "DPGA? ?LB GLRM "QG? ?LB &SPMNC GL RFC DGPQR K?HMP U?TC MD KGEP?RGML ?@MSR 1.5 KGJJGML WC?PQ ?EM. *R GQ RFMSEFR RF?R KMBCPL FSK?LQ ?PMQC GL "DPGA? DPMK H. ,9,*;<: ?LB KGEP?RCB MSR MD "DPGA? ?@MSR 100,000 WC?PQ ?EM GL ? QCAMLB K?HMP KGEP?RGML U?TC. 5FCL, KMBCPL FSK?LQ PCNJ?ACB H. ,9,*;<: QNCAGCQ RF?R F?B KGEP?RCB GLRM "QG? ?LB &SPMNC GL RFC DGPQR U?TC. 5FGQ CTMJSRGML?PW RGKCJGLC GQ QSNNMPRCB @W KMJCASJ?P CTGBCLAC. 0LC ?NNPM?AF RM QRSBWGLE RFC MPGEGLQ MD KMBCPL FSK?LQ GQ RM CV?KGLC KGRMAFMLBPG?J %/" (KR%/") DPMK NMNSJ?RGMLQ ?PMSLB RFC UMPJB. #CA?SQC ? DCRSQ BCTCJMNQ DPMK ?L CEE AMLR?GLGLE GRQ KMRFCPsQ KGRMAFMLBPG? (UFGAF F?TC RFCGP MUL, LML-LSAJC?P %/"), KR%/" GQ N?QQCB CLRGPCJW RFPMSEF RFC K?RCPL?J JGLC. .SR?RGMLQ GL KR%/" A?L LMU @C SQCB RM CQRGK?RC RFC RGKCJGLC MD ECLCRGA BGTCPECLAC. 5FC PCQSJRGLE CTGBCLAC QSEECQRQ RF?R ?JJ KMBCPL FSK?LQ F?TC KR%/" GLFCPGRCB DPMK ? AMKKML ?LACQRMP RF?R JGTCB GL "DPGA? ?@MSR 160,000 WC?PQ ?EM. "LMRFCP ?NNPM?AF RM RFC KMJCASJ?P SLBCPQR?LBGLE MD FSK?L CTMJSRGML GQ RM CV?KGLC RFC : AFPMKMQMKC, UFGAF GQ N?QQCB DPMK D?RFCP RM QML. 5FGQ CTGBCLAC QSEECQRQ RF?R ?JJ KCL RMB?W GLFCPGRCB ? : AFPMKMQMKC DPMK ? K?JC RF?R JGTCB GL "DPGA? ?@MSR 140,000 WC?PQ ?EM.
832
#(!04%2 29 X 6%24%"2!4%3
& 4 / -(.
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
#(!04%2 29 X 6%24%"2!4%3
833
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s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
834
#(!04%2 29 X 6%24%"2!4%3
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
#(!04%2 29 X 6%24%"2!4%3
835
KLE@:8K< QCQQGJC AFMPB?RC RF?R GQ ? KCK@CP MD 6PMAFMPB?R? 0IF:?FI;8K8 AJ?BC AMKNMQCB MD RSLGA?RCQ 0IF;<C8 Q?J?K?LBCPQ 1<IK<9I8K8 KCK@CPQ MD RFC NFWJSK $FMPB?R? RF?R NMQQCQQ ? @?AI@MLC M<IK<9I8C :FCLDE QCPGCQ MD QCN?P?RC @MLCQ HMGLCB RMECRFCP ?Q ? @?AI@MLC
C#A+/ - .0((A-4
29.1 C?FI;8K<J
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
29.2 !@J?<J
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
29.3 ADG?@9@8EJ
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
29.4 -<GK@C<J
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
29.5 B@I;J
#GPBQ ?PC CLBMRFCPKGA, KC?LGLE RFCW NPMBSAC RFCGP MUL @MBW FC?R ?LB PCESJ?RC RFCGP GLRCPL?J RCKNCP?RSPC GLBCNCLBCLRJW MD RFC CVRCPL?J RCKNCP?RSPC. 'C?RFCPQ LMR MLJW ?AR ?Q GLQSJ?RGML @SR ?JQM ?JJMU DMP DJGEFR, NPMTGBGLE JGDR UGRF QCAMLB?PW DC?RFCPQ ?LB RFPSQR UGRF NPGK?PW DC?RFCPQ. 1LCSK?RGA
836
#(!04%2 29 X 6%24%"2!4%3
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
29.6 (8DD8CJ
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
29.7 /?< MFCLK@FE F= +I@D8K<J

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864
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865
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC RUM QR?ECQ MD ? NJ?LRsQ JGDCAWAJC z $MKN?PC ?LB AMLRP?QR K?JC ?LB DCK?JC E?KCRMNFWRCQ ?LB CVNJ?GL FMU RFCW DMPK GL ?LEGMQNCPKQ z %CQAPG@C RFC PCNPMBSARGTC QRPSARSPCQ MD ? NJ?LR z %CQAPG@C RFC AMKNMLCLRQ MD ? AMKNJCRC DJMUCP z %CQAPG@C RFC BCTCJMNKCLR MD KGAPMQNMP?LEGSK ?LB KCE?QNMP?LEGSK GL EWKLMQNCPKQ 4CVS?J PCNPMBSARGML R?ICQ NJ?AC UGRF QJGEFR T?PG?RGMLQ GL BGDDCPCLR EPMSNQ MD NJ?LRQ. 1J?LRQ F?TC RUM BGQRGLAR QR?ECQ GL RFCGP JGDCAWAJC: RFC E?KCRMNFWRC QR?EC ?LB RFC QNMPMNFWRC QR?EC. 5FC F?NJMGB gametophyte NPMBSACQ RFC K?JC ?LB DCK?JC E?KCRCQ @W KGRMQGQ GL BGQRGLAR KSJRGACJJSJ?P QRPSARSPCQ. 'SQGML MD RFC K?JC ?LB DCK?JCQ E?KCRCQ DMPKQ RFC BGNJMGB XWEMRC, UFGAF BCTCJMNQ GLRM RFC sporophyte. "DRCP PC?AFGLE K?RSPGRW, RFC BGNJMGB QNMPMNFWRC NPMBSACQ QNMPCQ @W KCGMQGQ, UFGAF GL RSPL BGTGBC @W
902
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KGRMQGQ RM NPMBSAC RFC F?NJMGB E?KCRMNFWRC. 5FC LCU E?KCRMNFWRC NPMBSACQ E?KCRCQ, ?LB RFC AWAJC AMLRGLSCQ. 5FGQ GQ RFC ?JRCPL?RGML MD ECLCP?RGMLQ, ?LB GQ RWNGA?J MD NJ?LR PCNPMBSARGML ( Figure 32.2).
!@>LI< 32.2 4EB >IQBOK>QFLK LC DBKBO>QFLKP FK >KDFLPMBOJP FP ABMF@QBA FK QEFP AF>DO>J. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V 0BQBO #LUEB>A)
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5FC JGDCAWAJC MD ?LEGMQNCPKQ DMJJMUQ RFC ?JRCPL?RGML MD ECLCP?RGMLQ CVNJ?GLCB NPCTGMSQJW. 5FC F?NJMGB E?KCRMNFWRC ?JRCPL?RCQ UGRF RFC BGNJMGB QNMPMNFWRC BSPGLE RFC QCVS?J PCNPMBSARGML NPMACQQ MD ?LEGMQNCPKQ. 'JMUCPQ AMLR?GL RFC NJ?LRsQ PCNPMBSARGTC QRPSARSPCQ. F14<+7 !97:)9:7+ " RWNGA?J DJMUCP F?Q DMSP K?GL N?PRQqMP UFMPJQqILMUL ?Q RFC A?JWV, AMPMJJ?, ?LBPMCAGSK, ?LB EWLMCAGSK (Figure 32.3). 5FC MSRCPKMQR UFMPJ MD RFC DJMUCP F?Q EPCCL, JC?DW QRPSARSPCQ ILMUL ?Q QCN?JQ. 5FC QCN?JQ, AMJJCARGTCJW A?JJCB RFC A?JWV, FCJN RM NPMRCAR RFC SLMNCLCB @SB. 5FC QCAMLB UFMPJ GQ AMKNPGQCB MD NCR?JQqSQS?JJW, @PGEFRJW AMJMPCBqAMJJCARGTCJW A?JJCB RFC AMPMJJ?. 5FC LSK@CP MD QCN?JQ ?LB NCR?JQ T?PGCQ BCNCLBGLE ML UFCRFCP RFC NJ?LR GQ ? KMLMAMR MP BGAMR. *L KMLMAMRQ, NCR?JQ SQS?JJW LSK@CP RFPCC MP KSJRGNJCQ MD RFPCC; GL BGAMRQ, RFC LSK@CP MD NCR?JQ GQ DMSP MP DGTC, MP KSJRGNJCQ MD DMSP ?LB DGTC. 5MECRFCP, RFC A?JWV ?LB AMPMJJ? ?PC ILMUL ?Q RFC perianth. 5FC RFGPB UFMPJ AMLR?GLQ RFC K?JC PCNPMBSARGTC QRPSARSPCQ ?LB GQ ILMUL ?Q RFC ?LBPMCAGSK. 5FC androecium F?Q QR?KCLQ UGRF ?LRFCPQ RF?R AMLR?GL RFC KGAPMQNMP?LEG?. 5FC GLLCPKMQR EPMSN MD QRPSARSPCQ GL RFC DJMUCP GQ RFC gynoecium, MP RFC DCK?JC PCNPMBSARGTC AMKNMLCLR(Q). 5FC A?PNCJ GQ RFC GLBGTGBS?J SLGR MD RFC EWLMCAGSK ?LB F?Q ? QRGEK?, QRWJC, ?LB MT?PW. " DJMUCP K?W F?TC MLC MP KSJRGNJC A?PNCJQ.
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igure 32.4).
904
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!@>LI< 32.4 4EB @LOK MI>KQ E>P ?LQE PQ>JFK>QB (J>IB) >KA @>OMBII>QB (CBJ>IB) CILTBOP. 3Q>JFK>QB CILTBOP, TEF@E >OB @IRPQBOBA FK QEB Q>PPBI >Q QEB QFM LC QEB PQBJ, MOLAR@B MLIIBK DO>FKP. #>OMBII>QB CILTBO >OB @IRPQBOBA FK QEB FJJ>QROB B>OP. %>@E PQO>KA LC PFIH FP > PQFDJ>. 4EB @LOK HBOKBIP >OB PBBAP QE>Q ABSBILM LK QEB B>O >CQBO CBOQFIFW>QFLK. !IPL PELTK FP QEB ILTBO PQBJ >KA OLLQ.
*D @MRF K?JC ?LB DCK?JC DJMUCPQ ?PC @MPLC ML RFC Q?KC NJ?LR, RFC QNCAGCQ GQ A?JJCB KMLMCAGMSQ (KC?LGLE tMLC FMKCu): CV?KNJCQ ?PC AMPL ?LB NC?. 4NCAGCQ UGRF K?JC ?LB DCK?JC DJMUCPQ @MPLC ML QCN?P?RC NJ?LRQ ?PC RCPKCB BGMCAGMSQ, MP tRUM FMKCQ,u CV?KNJCQ MD UFGAF ?PC C. 7(7(@( ?LB C(55()0:. 5FC MT?PW, UFGAF K?W AMLR?GL MLC MP KSJRGNJC MTSJCQ, K?W @C NJ?ACB ?@MTC MRFCP DJMUCP N?PRQ, UFGAF GQ PCDCPPCB RM ?Q QSNCPGMP; MP, GR K?W @C NJ?ACB @CJMU RFC MRFCP DJMUCP N?PRQ, PCDCPPCB RM ?Q GLDCPGMP (Figure 32.5).
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905
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M'1+ G'2+945.>9+ (".+ P411+3 G7'/3) 5FC K?JC E?KCRMNFWRC BCTCJMNQ ?LB PC?AFCQ K?RSPGRW GL ?L GKK?RSPC ?LRFCP. *L ? NJ?LRsQ K?JC PCNPMBSARGTC MPE?LQ, BCTCJMNKCLR MD NMJJCL R?ICQ NJ?AC GL ? QRPSARSPC ILMUL ?Q RFC microsporangium (Figure 32.6). 5FC KGAPMQNMP?LEG?, UFGAF ?PC SQS?JJW @G-JM@CB, ?PC NMJJCL Q?AQ GL UFGAF RFC KGAPMQNMPCQ BCTCJMN GLRM NMJJCL EP?GLQ. 5FCQC ?PC DMSLB GL RFC ?LRFCP, UFGAF GQ ?R RFC CLB MD RFC QR?KCLqRFC JMLE DGJ?KCLR RF?R QSNNMPRQ RFC ?LRFCP.
906
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
8GRFGL RFC KGAPMQNMP?LEGSK, RFC KGAPMQNMPC KMRFCP ACJJ BGTGBCQ @W KCGMQGQ RM EGTC PGQC RM DMSP KGAPMQNMPCQ, C?AF MD UFGAF UGJJ SJRGK?RCJW DMPK ? NMJJCL EP?GL (Figure 32.7). "L GLLCP J?WCP MD ACJJQ, ILMUL ?Q RFC R?NCRSK, NPMTGBCQ LSRPGRGML RM RFC BCTCJMNGLE KGAPMQNMPCQ ?LB AMLRPG@SRCQ ICW AMKNMLCLRQ RM RFC NMJJCL U?JJ. .?RSPC NMJJCL EP?GLQ AMLR?GL RUM ACJJQ: ? ECLCP?RGTC ACJJ ?LB ? NMJJCL RS@C ACJJ. 5FC ECLCP?RGTC ACJJ GQ AMLR?GLCB UGRFGL RFC J?PECP NMJJCL RS@C ACJJ. 6NML ECPKGL?RGML, RFC RS@C ACJJ DMPKQ RFC NMJJCL RS@C RFPMSEF UFGAF RFC ECLCP?RGTC ACJJ KGEP?RCQ RM CLRCP RFC MT?PW. %SPGLE GRQ RP?LQGR GLQGBC RFC NMJJCL RS@C, RFC ECLCP?RGTC ACJJ BGTGBCQ RM DMPK RUM K?JC E?KCRCQ (QNCPK ACJJQ). 6NML K?RSPGRW, RFC KGAPMQNMP?LEG? @SPQR, PCJC?QGLE RFC NMJJCL EP?GLQ DPMK RFC ?LRFCP.
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907
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&?AF NMJJCL EP?GL F?Q RUM AMTCPGLEQ: RFC exine (RFGAICP, MSRCP J?WCP) ?LB RFC intine (Figure 32.7). 5FC CVGLC AMLR?GLQ QNMPMNMJJCLGL, ? AMKNJCV U?RCPNPMMDGLE QS@QR?LAC QSNNJGCB @W RFC R?NCR?J ACJJQ. 4NMPMNMJJCLGL ?JJMUQ RFC NMJJCL RM QSPTGTC SLBCP SLD?TMP?@JC AMLBGRGMLQ ?LB RM @C A?PPGCB @W UGLB, U?RCP, MP @GMJMEGA?J ?ECLRQ UGRFMSR SLBCPEMGLE B?K?EC. F+2'1+ G'2+945.>9+ (".+ E2(7>4 !')) 8FGJC RFC BCR?GJQ K?W T?PW @CRUCCL QNCAGCQ, RFC MTCP?JJ BCTCJMNKCLR MD RFC DCK?JC E?KCRMNFWRC F?Q RUM BGQRGLAR NF?QCQ. 'GPQR, GL RFC NPMACQQ MD megasporogenesis, ? QGLEJC ACJJ GL RFC BGNJMGB megasporangiumq?L ?PC? MD RGQQSC GL RFC MTSJCQqSLBCPEMCQ KCGMQGQ RM NPMBSAC DMSP KCE?QNMPCQ, MLJW MLC MD UFGAF QSPTGTCQ. %SPGLE RFC QCAMLB NF?QC, megagametogenesis, RFC QSPTGTGLE F?NJMGB KCE?QNMPC SLBCPEMCQ KGRMQGQ RM NPMBSAC ?L CGEFR-LSAJC?RC, QCTCL-ACJJ DCK?JC E?KCRMNFWRC, ?JQM ILMUL ?Q RFC KCE?E?KCRMNFWRC MP CK@PWM Q?A. 5UM MD RFC LSAJCGqRFC polar nucleiqKMTC RM RFC COS?RMP ?LB DSQC, DMPKGLE ? QGLEJC, BGNJMGB ACLRP?J ACJJ. 5FGQ ACLRP?J ACJJ J?RCP DSQCQ UGRF ? QNCPK RM DMPK RFC RPGNJMGB CLBMQNCPK. 5FPCC LSAJCG NMQGRGML RFCKQCJTCQ ML RFC CLB MD RFC CK@PWM Q?A MNNMQGRC RFC KGAPMNWJC ?LB BCTCJMN GLRM RFC antipodal ACJJQ, UFGAF J?RCP BCECLCP?RC. 5FC LSAJCSQ AJMQCQR RM RFC KGAPMNWJC @CAMKCQ RFC DCK?JC E?KCRC, MP CEE ACJJ, ?LB RFC RUM ?BH?ACLR LSAJCG BCTCJMN GLRM synergid ACJJQ (Figure 32.8). 5FC QWLCPEGBQ FCJN ESGBC RFC NMJJCL RS@C DMP QSAACQQDSJ DCPRGJGX?RGML, ?DRCP UFGAF RFCW BGQGLRCEP?RC. 0LAC DCPRGJGX?RGML GQ AMKNJCRC, RFC PCQSJRGLE BGNJMGB XWEMRC BCTCJMNQ GLRM RFC CK@PWM, ?LB RFC DCPRGJGXCB MTSJC DMPKQ RFC MRFCP RGQQSCQ MD RFC QCCB.
908
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915
QACLR. *L UGLB-NMJJGL?RCB QNCAGCQ, RFC KGAPMQNMP?LEG? F?LE MSR MD RFC DJMUCP, ?LB, ?Q RFC UGLB @JMUQ, RFC JGEFRUCGEFR NMJJCL GQ A?PPGCB UGRF GR (Figure 32.15). 5FC DJMUCPQ SQS?JJW CKCPEC C?PJW GL RFC QNPGLE, @CDMPC RFC JC?TCQ, QM RF?R RFC JC?TCQ BM LMR @JMAI RFC KMTCKCLR MD RFC UGLB. 5FC NMJJCL GQ BCNMQGRCB ML RFC CVNMQCB DC?RFCPW QRGEK? MD RFC DJMUCP (Figure 32.16).
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916
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"DRCP NMJJCL GQ BCNMQGRCB ML RFC QRGEK?, GR KSQR ECPKGL?RC ?LB EPMU RFPMSEF RFC QRWJC RM PC?AF RFC MTSJC. 5FC KGAPMQNMPCQ, MP RFC NMJJCL, AMLR?GL RUM ACJJQ: RFC NMJJCL RS@C ACJJ ?LB RFC ECLCP?RGTC ACJJ. 5FC NMJJCL RS@C ACJJ EPMUQ GLRM ? NMJJCL RS@C RFPMSEF UFGAF RFC ECLCP?RGTC ACJJ RP?TCJQ. 5FC ECPKGL?RGML MD RFC NMJJCL RS@C PCOSGPCQ U?RCP, MVWECL, ?LB ACPR?GL AFCKGA?J QGEL?JQ. "Q GR RP?TCJQ RFPMSEF RFC QRWJC RM PC?AF RFC CK@PWM Q?A, RFC NMJJCL RS@CsQ EPMURF GQ QSNNMPRCB @W RFC RGQQSCQ MD RFC QRWJC. *L RFC KC?LRGKC, GD RFC ECLCP?RGTC ACJJ F?Q LMR ?JPC?BW QNJGR GLRM RUM ACJJQ, GR LMU BGTGBCQ RM DMPK RUM QNCPK ACJJQ. 5FC NMJJCL RS@C GQ ESGBCB @W RFC AFCKGA?JQ QCAPCRCB @W RFC QWLCPEGBQ NPCQCLR GL RFC CK@PWM Q?A, ?LB GR CLRCPQ RFC MTSJC Q?A RFPMSEF RFC KGAPMNWJC. 0D RFC RUM QNCPK ACJJQ, MLC QNCPK DCPRGJGXCQ RFC CEE ACJJ, DMPKGLE ? BGNJMGB XWEMRC; RFC MRFCP QNCPK DSQCQ UGRF RFC RUM NMJ?P LSAJCG, DMPKGLE ? RPGNJMGB ACJJ RF?R BCTCJMNQ GLRM RFC endosperm. 5MECRFCP, RFCQC RUM DCPRGJGX?RGML CTCLRQ GL ?LEGMQNCPKQ ?PC ILMUL ?Q double fertilization (Figure 32.18). "DRCP DCPRGJGX?RGML GQ AMKNJCRC, LM MRFCP QNCPK A?L CLRCP. 5FC DCPRGJGXCB MTSJC DMPKQ RFC QCCB, UFCPC?Q RFC RGQQSCQ MD RFC MT?PW @CAMKC RFC DPSGR, SQS?JJW CLTCJMNGLE RFC QCCB.
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917
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"DRCP DCPRGJGX?RGML, RFC XWEMRC BGTGBCQ RM DMPK RUM ACJJQ: RFC SNNCP ACJJ, MP RCPKGL?J ACJJ, ?LB RFC JMUCP, MP @?Q?J, ACJJ. 5FC BGTGQGML MD RFC @?Q?J ACJJ EGTCQ PGQC RM RFC suspensor, UFGAF CTCLRS?JJW K?ICQ AMLLCARGML UGRF RFC K?RCPL?J RGQQSC. 5FC QSQNCLQMP NPMTGBCQ ? PMSRC DMP LSRPGRGML RM @C RP?LQNMPRCB DPMK RFC KMRFCP NJ?LR RM RFC EPMUGLE CK@PWM. 5FC RCPKGL?J ACJJ ?JQM BGTGBCQ, EGTGLE PGQC RM ? EJM@SJ?PQF?NCB NPMCK@PWM (Figure 32.19a). *L BGAMRQ (CSBGAMRQ), RFC BCTCJMNGLE CK@PWM F?Q ? FC?PR QF?NC, BSC RM RFC NPCQCLAC MD RFC RUM PSBGKCLR?PW cotyledons (Figure 32.19b). *L LML-CLBMQNCPKGA BGAMRQ, QSAF ?Q C(7:,33( )<9:(, RFC CLBMQNCPK BCTCJMNQ GLGRG?JJW, @SR GQ RFCL BGECQRCB, ?LB RFC DMMB PCQCPTCQ ?PC KMTCB GLRM RFC RUM AMRWJCBMLQ. "Q RFC CK@PWM ?LB AMRWJCBMLQ CLJ?PEC, RFCW PSL MSR MD PMMK GLQGBC RFC BCTCJMNGLE QCCB, ?LB ?PC DMPACB RM @CLB (Figure 32.19c). 6JRGK?RCJW, RFC CK@PWM ?LB AMRWJCBMLQ DGJJ RFC QCCB (Figure 32.19d), ?LB RFC QCCB GQ PC?BW DMP BGQNCPQ?J. &K@PWMLGA BCTCJMNKCLR GQ QSQNCLBCB ?DRCP QMKC RGKC, ?LB EPMURF GQ PCQSKCB MLJW UFCL RFC QCCB ECPKGL?RCQ. 5FC BCTCJMNGLE QCCBJGLE UGJJ PCJW ML RFC DMMB PCQCPTCQ QRMPCB GL RFC AMRWJCBMLQ SLRGJ RFC DGPQR QCR MD JC?TCQ @CEGL NFMRMQWLRFCQGQ.
918
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919
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7E>Q FP LC QEB CLIILTFKD PQ>QBJBKQP FP QORB >. "LQE JLKL@LQP >KA AF@LQP E>SB >K BKALPMBOJ. ?. 4EB O>AF@IB ABSBILMP FKQL QEB OLLQ. @. 4EB MIRJRIB FP M>OQ LC QEB BMF@LQVI A. 4EB BKALPMBOJ FP M>OQ LC QEB BJ?OVL. 5FC QRMP?EC MD DMMB PCQCPTCQ GL ?LEGMQNCPK QCCBQ BGDDCPQ @CRUCCL KMLMAMRQ ?LB BGAMRQ. *L KMLMAMRQ, QSAF ?Q AMPL ?LB UFC?R, RFC QGLEJC AMRWJCBML GQ A?JJCB ? scutellum; RFC QASRCJJSK GQ AMLLCARCB BGPCARJW RM RFC CK@PWM TG? T?QASJ?P RGQQSC (VWJCK ?LB NFJMCK). 'MMB PCQCPTCQ ?PC QRMPCB GL RFC J?PEC CLBMQNCPK. 6NML ECPKGL?RGML, CLXWKCQ ?PC QCAPCRCB @W RFC aleurone, ? QGLEJC J?WCP MD ACJJQ HSQR GLQGBC RFC QCCB AM?R RF?R QSPPMSLBQ RFC CLBMQNCPK ?LB CK@PWM. 5FC CLXWKCQ BCEP?BC RFC QRMPCB A?P@MFWBP?RCQ, NPMRCGLQ ?LB JGNGBQ, RFC NPMBSARQ MD UFGAF ?PC ?@QMP@CB @W RFC QASRCJJSK ?LB RP?LQNMPRCB TG? ? T?QASJ?RSPC QRP?LB RM RFC BCTCJMNGLE CK@PWM. 5FCPCDMPC, RFC QASRCJJSK A?L @C QCCL RM @C ?L ?@QMPNRGTC MPE?L, LMR ? QRMP?EC MPE?L. 5FC RUM AMRWJCBMLQ GL RFC BGAMR QCCB ?JQM F?TC T?QASJ?P AMLLCARGMLQ RM RFC CK@PWM. *L endospermic dicots, RFC DMMB PCQCPTCQ ?PC QRMPCB GL RFC CLBMQNCPK. %SPGLE ECPKGL?RGML, RFC RUM AMRWJCBMLQ RFCPCDMPC ?AR ?Q ?@QMPNRGTC MPE?LQ RM R?IC SN RFC CLXWK?RGA?JJW PCJC?QCB DMMB PCQCPTCQ, KSAF JGIC GL KMLMAMRQ (KMLMAMRQ, @W BCDGLGRGML, ?JQM F?TC CLBMQNCPKGA QCCBQ). 5M@?AAM (N0*6;0(5( ;()(**<4), RMK?RM ( 63(5<4 3@*67,9:0*<4), ?LB NCNNCP (C(7:0*<4 (55<<4) ?PC CV?KNJCQ MD CLBMQNCPKGA BGAMRQ. *L nonendospermic dicots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testa ?LB GLLCP AM?R ILMUL ?Q RFC tegmen. 5FC CK@PWMLGA ?VGQ AMLQGQRQ MD RFPCC N?PRQ: RFC NJSKSJC, RFC P?BGAJC, ?LB RFC FWNMAMRWJ. 5FC NMPRGML MD RFC CK@PWM @CRUCCL RFC AMRWJCBML ?RR?AFKCLR NMGLR ?LB RFC P?BGAJC GQ ILMUL ?Q RFC hypocotyl (FWNMAMRWJ KC?LQ t@CJMU RFC AMRWJCBMLQu). 5FC CK@PWMLGA ?VGQ RCPKGL?RCQ GL ? radicle (RFC CK@PWMLGA PMMR), UFGAF GQ RFC PCEGML DPMK UFGAF RFC PMMR UGJJ BCTCJMN. *L BGAMRQ, RFC FWNMAMRWJQ CVRCLB ?@MTC EPMSLB, EGTGLE PGQC RM RFC QRCK MD RFC NJ?LR. *L KMLMAMRQ, RFC FWNMAMRWJ BMCQ LMR QFMU ?@MTC EPMSLB @CA?SQC KMLMAMRQ BM LMR CVFG@GR QRCK CJMLE?RGML. 5FC N?PR MD RFC CK@PWMLGA ?VGQ RF?R NPMHCARQ ?@MTC RFC AMRWJCBMLQ GQ ILMUL ?Q RFC epicotyl. 5FC plumule GQ AMKNMQCB MD RFC CNGAMRWJ, WMSLE JC?TCQ, ?LB RFC QFMMR ?NGA?J KCPGQRCK. 6NML ECPKGL?RGML GL BGAMR QCCBQ, RFC CNGAMRWJ GQ QF?NCB JGIC ? FMMI UGRF RFC NJSKSJC NMGLRGLE BMULU?PBQ. 5FGQ QF?NC GQ A?JJCB RFC NJSKSJC FMMI, ?LB GR NCPQGQRQ ?Q JMLE ?Q ECPKGL?RGML NPMACCBQ GL RFC B?PI. 5FCPCDMPC, ?Q RFC CNGAMRWJ NSQFCQ RFPMSEF RFC RMSEF ?LB ?@P?QGTC QMGJ, RFC NJSKSJC GQ NPMRCARCB DPMK B?K?EC. 6NML CVNMQSPC RM JGEFR, RFC FWNMAMRWJ FMMI QRP?GEFRCLQ MSR, RFC WMSLE DMJG?EC JC?TCQ D?AC RFC QSL ?LB CVN?LB, ?LB RFC CNGAMRWJ AMLRGLSCQ RM CJMLE?RC. %SPGLE RFGQ RGKC, RFC P?BGAJC GQ ?JQM EPMUGLE
920
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?LB NPMBSAGLE RFC NPGK?PW PMMR. "Q GR EPMUQ BMULU?PB RM DMPK RFC R?N PMMR, J?RCP?J PMMRQ @P?LAF MDD RM ?JJ QGBCQ, NPMBSAGLE RFC RWNGA?J BGAMR R?N PMMR QWQRCK. *L KMLMAMR QCCBQ (Figure 32.21), RFC RCQR? ?LB RCEKCL MD RFC QCCB AM?R ?PC DSQCB. "Q RFC QCCB ECPKGL?RCQ, RFC NPGK?PW PMMR CKCPECQ, NPMRCARCB @W RFC PMMR-RGN AMTCPGLE: RFC coleorhiza. /CVR, RFC NPGK?PW QFMMR CKCPECQ, NPMRCARCB @W RFC coleoptile: RFC AMTCPGLE MD RFC QFMMR RGN. 6NML CVNMQSPC RM JGEFR (G.C. UFCL RFC NJSKSJC F?Q CVGRCB RFC QMGJ ?LB RFC NPMRCARGTC AMJCMNRGJC GQ LM JMLECP LCCBCB), CJMLE?RGML MD RFC AMJCMNRGJC AC?QCQ ?LB RFC JC?TCQ CVN?LB ?LB SLDMJB. "R RFC MRFCP CLB MD RFC CK@PWMLGA ?VGQ, RFC NPGK?PW PMMR QMML BGCQ, UFGJC MRFCP, ?BTCLRGRGMSQ PMMRQ (PMMRQ RF?R BM LMR ?PGQC DPMK RFC SQS?J NJ?AC p G.C. RFC PMMR) CKCPEC DPMK RFC @?QC MD RFC QRCK. 5FGQ EGTCQ RFC KMLMAMR ? DG@PMSQ PMMR QWQRCK.
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921
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928
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929
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936
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E3+7-> +6:/7+2+398 +1'9+* 94 L+;+18 4, A)9/;/9> 5FC KMPC ?ARGTC ?L ?LGK?J GQ, RFC KMPC CLCPEW GQ LCCBCB RM K?GLR?GL RF?R ?ARGTGRW, ?LB RFC FGEFCP GRQ #.3 MP 4.3. 5FC ?TCP?EC B?GJW P?RC MD CLCPEW AMLQSKNRGML GQ ?@MSR RUM RM DMSP RGKCQ ?L ?LGK?Js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orpor GQ ? NPMACQQ RF?R JC?BQ RM ? BCAPC?QC GL ?ARGTGRW ?LB KCR?@MJGQK ?LB ?JJMUQ ?LGK?JQ RM QSPTGTC ?BTCPQC AMLBGRGMLQ. 5MPNMP A?L @C SQCB @W ?LGK?JQ DMP JMLE NCPGMBQ, QSAF ?Q CLRCPGLE ? QR?RC MD hibernation BSPGLE RFC UGLRCP KMLRFQ, GL UFGAF A?QC GR CL?@JCQ RFCK RM K?GLR?GL ? PCBSACB @MBW RCKNCP?RSPC. %SPGLE FG@CPL?RGML, EPMSLB QOSGPPCJQ A?L ?AFGCTC ?L ?@BMKGL?J RCKNCP?RSPC MD 0\ $ (32\ '), UFGJC ? @C?PsQ GLRCPL?J RCKNCP?RSPC GQ K?GLR?GLCB FGEFCP ?R ?@MSR 37\ $ (99\ '). *D RMPNMP MAASPQ BSPGLE RFC QSKKCP KMLRFQ UGRF FGEF RCKNCP?RSPCQ ?LB JGRRJC U?RCP, GR GQ A?JJCB estivation. 4MKC BCQCPR ?LGK?JQ SQC RFGQ RM QSPTGTC RFC F?PQFCQR KMLRFQ MD RFC WC?P. 5MPNMP A?L MAASP ML ? B?GJW @?QGQ; RFGQ GQ QCCL GL @?RQ ?LB FSKKGLE@GPBQ. 8FGJC CLBMRFCPKW GQ JGKGRCB GL QK?JJCP ?LGK?JQ @W QSPD?AC RM TMJSKC P?RGM, QMKC MPE?LGQKQ A?L @C QK?JJCP ?LB QRGJJ @C CLBMRFCPKQ @CA?SQC RFCW CKNJMW B?GJW RMPNMP BSPGLE RFC N?PR MD RFC B?W RF?R GQ AMJBCQR. 5FGQ ?JJMUQ RFCK RM AMLQCPTC CLCPEW BSPGLE RFC AMJBCP N?PRQ MD RFC B?W, UFCL RFCW AMLQSKC KMPC CLCPEW RM K?GLR?GL RFCGP @MBW RCKNCP?RSPC.
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" QR?LBGLE TCPRC@P?RC ?LGK?J A?L @C BGTGBCB @W QCTCP?J NJ?LCQ. " sagittal plane BGTGBCQ RFC @MBW GLRM PGEFR ?LB JCDR NMPRGMLQ. " midsagittal plane BGTGBCQ RFC @MBW CV?ARJW GL RFC KGBBJC, K?IGLE RUM COS?J PGEFR ?LB JCDR F?JTCQ. " frontal plane (?JQM A?JJCB ? AMPML?J NJ?LC) QCN?P?RCQ RFC DPMLR DPMK RFC @?AI. " transverse plane (MP, FMPGXMLR?J NJ?LC) BGTGBCQ RFC ?LGK?J GLRM SNNCP ?LB JMUCP NMPRGMLQ. 5FGQ GQ QMKCRGKCQ A?JJCB ? APMQQ QCARGML, ?LB, GD RFC RP?LQTCPQC ASR GQ ?R ?L ?LEJC, GR GQ A?JJCB ?L M@JGOSC NJ?LC. Figure 33.5 GJJSQRP?RCQ RFCQC NJ?LCQ ML ? EM?R (? DMSP-JCEECB ?LGK?J) ?LB ? FSK?L @CGLE.
938
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!@>LI< 33.5 3ELTK >OB QEB MI>KBP LC > NR>AORMBA DL>Q >KA > ?FMBA>I ERJ>K. 4EB JFAP>DFQQ>I MI>KB AFSFABP QEB ?LAV BU>@QIV FK E>IC, FKQL OFDEQ >KA IBCQ MLOQFLKP. 4EB COLKQ>I MI>KB AFSFABP QEB COLKQ >KA ?>@H, >KA QEB QO>KPSBOPB MI>KB AFSFABP QEB ?LAV FKQL RMMBO >KA ILTBO MLOQFLKP.
7CPRC@P?RC ?LGK?JQ F?TC ? LSK@CP MD BCDGLCB @MBW A?TGRGCQ, ?Q GJJSQRP?RCB GL Figure 33.6. 5UM MD RFCQC ?PC K?HMP A?TGRGCQ RF?R AMLR?GL QK?JJCP A?TGRGCQ UGRFGL RFCK. 5FC dorsal cavity AMLR?GLQ RFC AP?LG?J ?LB RFC TCPRC@P?J (MP QNGL?J) A?TGRGCQ. 5FC ventral cavity AMLR?GLQ RFC RFMP?AGA A?TGRW, UFGAF GL RSPL AMLR?GLQ RFC NJCSP?J A?TGRW ?PMSLB RFC JSLEQ ?LB RFC NCPGA?PBG?J A?TGRW, UFGAF QSPPMSLBQ RFC FC?PR. 5FC TCLRP?J A?TGRW ?JQM AMLR?GLQ RFC ?@BMKGLMNCJTGA A?TGRW, UFGAF A?L @C QCN?P?RCB GLRM RFC ?@BMKGL?J ?LB RFC NCJTGA A?TGRGCQ.
!@>LI< 33.6 6BOQB?O>QB >KFJ>IP E>SB QTL J>GLO ?LAV @>SFQFBP. 4EB ALOP>I @>SFQV, FKAF@>QBA FK DOBBK, @LKQ>FKP QEB @O>KF>I >KA QEB PMFK>I @>SFQV. 4EB SBKQO>I @>SFQV, FKAF@>QBA FK VBIILT, @LKQ>FKP QEB QELO>@F@ @>SFQV >KA QEB >?ALJFKLMBISF@ @>SFQV. 4EB QELO>@F@ @>SFQV FP PBM>O>QBA COLJ QEB >?ALJFKLMBISF@ @>SFQV ?V QEB AF>MEO>DJ. 4EB QELO>@F@ @>SFQV FP PBM>O>QBA FKQL QEB >?ALJFK>I @>SFQV >KA QEB MBISF@ @>SFQV ?V >K FJ>DFK>OV IFKB M>O>IIBI QL QEB MBISFP ?LKBP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .#))
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C CNGRFCJG?J RGQQSCQ z %GQASQQ RFC BGDDCPCLR RWNCQ MD AMLLCARGTC RGQQSCQ GL ?LGK?JQ z %CQAPG@C RFPCC RWNCQ MD KSQAJC RGQQSCQ z %CQAPG@C LCPTMSQ RGQQSC 5FC RGQQSCQ MD KSJRGACJJSJ?P, AMKNJCV ?LGK?JQ ?PC DMSP NPGK?PW RWNCQ: CNGRFCJG?J, AMLLCARGTC, KSQAJC, ?LB LCPTMSQ. 3CA?JJ RF?R RGQQSCQ ?PC EPMSNQ MD QGKGJ?P ACJJQ EPMSN MD QGKGJ?P ACJJQ A?PPWGLE MSR PCJ?RCB DSLARGMLQ. 5FCQC RGQQSCQ AMK@GLC RM DMPK MPE?LQqJGIC RFC QIGL MP IGBLCWqRF?R F?TC QNCAGDGA, QNCAG?JGXCB DSLARGMLQ UGRFGL RFC @MBW. 0PE?LQ ?PC MPE?LGXCB GLRM MPE?L QWQRCKQ RM NCPDMPK DSLARGMLQ; CV?KNJCQ GLAJSBC RFC AGPASJ?RMPW QWQRCK, UFGAF AMLQGQRQ MD RFC FC?PR ?LB @JMMB TCQQCJQ, ?LB RFC BGECQRGTC QWQRCK, AMLQGQRGLE MD QCTCP?J MPE?LQ, GLAJSBGLE RFC QRMK?AF, GLRCQRGLCQ, JGTCP, ?LB N?LAPC?Q. 0PE?L QWQRCKQ AMKC RMECRFCP RM APC?RC ?L CLRGPC MPE?LGQK.
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Epithelial tissues AMTCP RFC MSRQGBC MD MPE?LQ ?LB QRPSARSPCQ GL RFC @MBW ?LB JGLC RFC JSKCLQ MD MPE?LQ GL ? QGLEJC J?WCP MP KSJRGNJC J?WCPQ MD ACJJQ. 5FC RWNCQ MD CNGRFCJG? ?PC AJ?QQGDGCB @W RFC QF?NCQ MD ACJJQ NPCQCLR ?LB RFC LSK@CP MD J?WCPQ MD ACJJQ. &NGRFCJG? AMKNMQCB MD ? QGLEJC J?WCP MD ACJJQ GQ A?JJCB simple epithelia; CNGRFCJG?J RGQQSC AMKNMQCB MD KSJRGNJC J?WCPQ GQ A?JJCB stratified epithelia. Table 33.2 QSKK?PGXCQ RFC BGDDCPCLR RWNCQ MD CNGRFCJG?J RGQQSCQ.
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!@>LI< 33.7 3NR>JLRP BMFQEBIF> @BIIP (>) E>SB > PIFDEQIV FOOBDRI>O PE>MB, >KA > PJ>II, @BKQO>IIV IL@>QBA KR@IBRP. 4EBPB @BIIP @>K ?B PQO>QFCFBA FKQL I>VBOP, >P FK (?) QEFP ERJ>K @BOSFU PMB@FJBK. (@OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V %A 5QEJ>K; P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
Figure 33.7a GJJSQRP?RCQ ? J?WCP MD QOS?KMSQ ACJJQ UGRF RFCGP KCK@P?LCQ HMGLCB RMECRFCP RM DMPK ?L CNGRFCJGSK. *K?EC Figure 33.7b GJJSQRP?RCQ QOS?KMSQ CNGRFCJG?J ACJJQ ?PP?LECB GL QRP?RGDGCB J?WCPQ, UFCPC NPMRCARGML GQ LCCBCB ML RFC @MBW DPMK MSRQGBC ?@P?QGML ?LB B?K?EC. 5FGQ GQ A?JJCB ? QRP?RGDGCB QOS?KMSQ CNGRFCJGSK ?LB MAASPQ GL RFC QIGL ?LB GL RGQQSCQ JGLGLE RFC KMSRF ?LB T?EGL?. C:(4/*'1 E5/9.+1/' Cuboidal epithelial ACJJQ, QFMUL GL Figure 33.8, ?PC AS@C-QF?NCB UGRF ? QGLEJC, ACLRP?J LSAJCSQ. 5FCW ?PC KMQR AMKKMLJW DMSLB GL ? QGLEJC J?WCP PCNPCQCLRGLE ? QGKNJC CNGRFCJG? GL EJ?LBSJ?P RGQQSCQ RFPMSEFMSR RFC @MBW UFCPC RFCW NPCN?PC ?LB QCAPCRC EJ?LBSJ?P K?RCPG?J. 5FCW ?PC ?JQM DMSLB GL RFC U?JJQ MD RS@SJCQ ?LB GL RFC BSARQ MD RFC IGBLCW ?LB JGTCP.
!@>LI< 33.8 3FJMIB @R?LFA>I BMFQEBIF>I @BIIP IFKB QR?RIBP FK QEB J>JJ>IF>K HFAKBV, TEBOB QEBV >OB FKSLISBA FK CFIQBOFKD QEB ?ILLA.
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941
C41:23'7 E5/9.+1/' Columnar epithelial ACJJQ ?PC R?JJCP RF?L RFCW ?PC UGBC: RFCW PCQCK@JC ? QR?AI MD AMJSKLQ GL ?L CNGRFCJG?J J?WCP, ?LB ?PC KMQR AMKKMLJW DMSLB GL ? QGLEJC-J?WCP ?PP?LECKCLR. 5FC LSAJCG MD AMJSKL?P CNGRFCJG?J ACJJQ GL RFC BGECQRGTC RP?AR ?NNC?P RM @C JGLCB SN ?R RFC @?QC MD RFC ACJJQ, ?Q GJJSQRP?RCB GL Figure 33.9. 5FCQC ACJJQ ?@QMP@ K?RCPG?J DPMK RFC JSKCL MD RFC BGECQRGTC RP?AR ?LB NPCN?PC GR DMP CLRPW GLRM RFC @MBW RFPMSEF RFC AGPASJ?RMPW ?LB JWKNF?RGA QWQRCKQ.
!@>LI< 33.9 3FJMIB @LIRJK>O BMFQEBIF>I @BIIP >?PLO? J>QBOF>I COLJ QEB AFDBPQFSB QO>@Q. 'L?IBQ @BIIP PB@OBQ JR@LRP FKQL QEB AFDBPQFSB QO>@Q IRJBK.
$MJSKL?P CNGRFCJG?J ACJJQ JGLGLE RFC PCQNGP?RMPW RP?AR ?NNC?P RM @C QRP?RGDGCB. )MUCTCP, C?AF ACJJ GQ ?RR?AFCB RM RFC @?QC KCK@P?LC MD RFC RGQQSC ?LB, RFCPCDMPC, RFCW ?PC QGKNJC RGQQSCQ. 5FC LSAJCG ?PC ?PP?LECB ?R BGDDCPCLR JCTCJQ GL RFC J?WCP MD ACJJQ, K?IGLE GR ?NNC?P ?Q RFMSEF RFCPC GQ KMPC RF?L MLC J?WCP, ?Q QCCL GL Figure 33.10. 5FGQ GQ A?JJCB pseudostratified, AMJSKL?P CNGRFCJG?. 5FGQ ACJJSJ?P AMTCPGLE F?Q AGJG? ?R RFC ?NGA?J, MP DPCC, QSPD?AC MD RFC ACJJQ. 5FC AGJG? CLF?LAC RFC KMTCKCLR MD KSAMSQ ?LB RP?NNCB N?PRGAJCQ MSR MD RFC PCQNGP?RMPW RP?AR, FCJNGLE RM NPMRCAR RFC QWQRCK DPMK GLT?QGTC KGAPMMPE?LGQKQ ?LB F?PKDSJ K?RCPG?J RF?R F?Q @CCL @PC?RFCB GLRM RFC @MBW. (M@JCR ACJJQ ?PC GLRCPQNCPQCB GL QMKC RGQQSCQ (QSAF ?Q RFC JGLGLE MD RFC RP?AFC?). 5FC EM@JCR ACJJQ AMLR?GL KSAMSQ RF?R RP?NQ GPPGR?LRQ, UFGAF GL RFC A?QC MD RFC RP?AFC? ICCN RFCQC GPPGR?LRQ DPMK ECRRGLE GLRM RFC JSLEQ.
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942
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Connective tissues ?PC K?BC SN MD ? K?RPGV AMLQGQRGLE MD JGTGLE ACJJQ ?LB ? LML-JGTGLE QS@QR?LAC, A?JJCB RFC EPMSLB QS@QR?LAC. 5FC EPMSLB QS@QR?LAC GQ K?BC MD ?L MPE?LGA QS@QR?LAC (SQS?JJW ? NPMRCGL) ?LB ?L GLMPE?LGA QS@QR?LAC (SQS?JJW ? KGLCP?J MP U?RCP). 5FC NPGLAGN?J ACJJ MD AMLLCARGTC RGQQSCQ GQ RFC DG@PM@J?QR. 5FGQ ACJJ K?ICQ RFC DG@CPQ DMSLB GL LC?PJW ?JJ MD RFC AMLLCARGTC RGQQSCQ. 'G@PM@J?QRQ ?PC KMRGJC, ?@JC RM A?PPW MSR KGRMQGQ, ?LB A?L QWLRFCQGXC UFGAFCTCP AMLLCARGTC RGQQSC GQ LCCBCB. .?APMNF?ECQ, JWKNFMAWRCQ, ?LB, MAA?QGML?JJW, JCSIMAWRCQ A?L @C DMSLB GL QMKC MD RFC RGQQSCQ. 4MKC RGQQSCQ F?TC QNCAG?JGXCB ACJJQ RF?R ?PC LMR DMSLB GL RFC MRFCPQ. 5FC matrix GL AMLLCARGTC RGQQSCQ EGTCQ RFC RGQQSC GRQ BCLQGRW. 8FCL ? AMLLCARGTC RGQQSC F?Q ? FGEF AMLACLRP?RGML MD ACJJQ MP DG@CPQ, GR F?Q NPMNMPRGML?JJW ? JCQQ BCLQC K?RPGV. 5FC MPE?LGA NMPRGML MP NPMRCGL DG@CPQ DMSLB GL AMLLCARGTC RGQQSCQ ?PC CGRFCP AMJJ?ECL, CJ?QRGA, MP PCRGASJ?P DG@CPQ. $MJJ?ECL DG@CPQ NPMTGBC QRPCLERF RM RFC RGQQSC, NPCTCLRGLE GR DPMK @CGLE RMPL MP QCN?P?RCB DPMK RFC QSPPMSLBGLE RGQQSCQ. &J?QRGA DG@CPQ ?PC K?BC MD RFC NPMRCGL CJ?QRGL; RFGQ DG@CP A?L QRPCRAF RM MLC ?LB MLC F?JD MD GRQ JCLERF ?LB PCRSPL RM GRQ MPGEGL?J QGXC ?LB QF?NC. &J?QRGA DG@CPQ NPMTGBC DJCVG@GJGRW RM RFC RGQQSCQ. 3CRGASJ?P DG@CPQ ?PC RFC RFGPB RWNC MD NPMRCGL DG@CP DMSLB GL AMLLCARGTC RGQQSCQ. 5FGQ DG@CP AMLQGQRQ MD RFGL QRP?LBQ MD AMJJ?ECL RF?R DMPK ? LCRUMPI MD DG@CPQ RM QSNNMPR RFC RGQQSC ?LB MRFCP MPE?LQ RM UFGAF GR GQ AMLLCARCB. 5FC T?PGMSQ RWNCQ MD AMLLCARGTC RGQQSCQ, RFC RWNCQ MD ACJJQ ?LB DG@CPQ RFCW ?PC K?BC MD, ?LB Q?KNJC JMA?RGMLQ MD RFC RGQQSCQ GQ QSKK?PGXCB GL Table 33.3.
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943
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>OLRKA ?ILLA SBPPBIP; >K@ELOP BMFQEBIF> FOOBDRI>O: PHFK OBDRI>O: QBKALKP, IFD>JBKQP PE>OH PHBIBQLK, CBQ>I ?LKBP, ERJ>K B>OP, FKQBOSBOQB?O>I AFP@P SBOQB?O>QB PHBIBQLKP >AFMLPB (C>Q) ?ILLA
CF?OL?I>PQP, J>@OLME>DBP, CBT: @LII>DBK, BI>PQF@, PLJB IVJMEL@VQBP, PLJB OBQF@RI>O KBRQOLMEFIP CF?OL?I>PQP, J>@OLME>DBP, JLPQIV @LII>DBK EV>IFKB: CBT @LII>DBK CF?OL@>OQFI>DB: I>ODB >JLRKQ LC @LII>DBK PLJB: @LII>DBK, BI>PQF@ CBT KLKB
@ELKAOL@VQBP, @ELKAOL?I>PQP LPQBL?I>PQP, LPQBL@VQBP, LPQBL@I>PQP >AFML@VQBP OBA ?ILLA @BIIP, TEFQB ?ILLA @BIIP
L448+/A7+41'7 C433+)9/;+ "/88:+ Loose connective tissue, ?JQM A?JJCB ?PCMJ?P AMLLCARGTC RGQQSC, F?Q ? Q?KNJGLE MD ?JJ MD RFC AMKNMLCLRQ MD ? AMLLCARGTC RGQQSC. "Q GJJSQRP?RCB GL Figure 33.12, JMMQC AMLLCARGTC RGQQSC F?Q QMKC DG@PM@J?QRQ; K?APMNF?ECQ ?PC NPCQCLR ?Q UCJJ. $MJJ?ECL DG@CPQ ?PC PCJ?RGTCJW UGBC ?LB QR?GL ? JGEFR NGLI, UFGJC CJ?QRGA DG@CPQ ?PC RFGL ?LB QR?GL B?PI @JSC RM @J?AI. 5FC QN?AC @CRUCCL RFC DMPKCB CJCKCLRQ MD RFC RGQQSC GQ DGJJCB UGRF RFC K?RPGV. 5FC K?RCPG?J GL RFC AMLLCARGTC RGQQSC EGTCQ GR ? JMMQC AMLQGQRCLAW QGKGJ?P RM ? AMRRML @?JJ RF?R F?Q @CCL NSJJCB ?N?PR. -MMQC AMLLCARGTC RGQQSC GQ DMSLB ?PMSLB CTCPW @JMMB TCQQCJ ?LB FCJNQ RM ICCN RFC TCQQCJ GL NJ?AC. 5FC RGQQSC GQ ?JQM DMSLB ?PMSLB ?LB @CRUCCL KMQR @MBW MPE?LQ. *L QSKK?PW, ?PCMJ?P RGQQSC GQ RMSEF, WCR DJCVG@JC, ?LB AMKNPGQCQ KCK@P?LCQ.
!@>LI< 33.12 ,LLPB @LKKB@QFSB QFPPRB FP @LJMLPBA LC ILLPBIV TLSBK @LII>DBK >KA BI>PQF@ CF?BOP. 4EB CF?BOP >KA LQEBO @LJMLKBKQP LC QEB @LKKB@QFSB QFPPRB J>QOFU >OB PB@OBQBA ?V CF?OL?I>PQP.
F/(74:8 C433+)9/;+ "/88:+ Fibrous connective tissues AMLR?GL J?PEC ?KMSLRQ MD AMJJ?ECL DG@CPQ ?LB DCU ACJJQ MP K?RPGV K?RCPG?J. 5FC DG@CPQ A?L @C ?PP?LECB GPPCESJ?PJW MP PCESJ?PJW UGRF RFC QRP?LBQ JGLCB SN GL N?P?JJCJ. *PPCESJ?PJW ?PP?LECB DG@PMSQ AMLLCARGTC RGQQSCQ ?PC DMSLB GL ?PC?Q MD RFC @MBW UFCPC QRPCQQ MAASPQ DPMK ?JJ BGPCARGMLQ, QSAF ?Q RFC BCPKGQ MD RFC QIGL. 3CESJ?P DG@PMSQ AMLLCARGTC RGQQSC, QFMUL GL Figure 33.13, GQ DMSLB GL RCLBMLQ (UFGAF AMLLCAR KSQAJCQ RM @MLCQ) ?LB JGE?KCLRQ (UFGAF AMLLCAR @MLCQ RM @MLCQ).
944
#(!04%2 33 X 4(% !.)-!, "/$9: "!3)# &/2- !.$ &5.#4)/.
!@>LI< 33.13 &F?OLRP @LKKB@QFSB QFPPRB COLJ QEB QBKALK E>P PQO>KAP LC @LII>DBK CF?BOP IFKBA RM FK M>O>IIBI.
C'79/1'-+ Cartilage GQ ? AMLLCARGTC RGQQSC UGRF ? J?PEC ?KMSLR MD RFC K?RPGV ?LB T?PG?@JC ?KMSLRQ MD DG@CPQ. 5FC ACJJQ, A?JJCB chondrocytes, K?IC RFC K?RPGV ?LB DG@CPQ MD RFC RGQQSC. $FMLBPMAWRCQ ?PC DMSLB GL QN?ACQ UGRFGL RFC RGQQSC A?JJCB lacunae. " A?PRGJ?EC UGRF DCU AMJJ?ECL ?LB CJ?QRGA DG@CPQ GQ FW?JGLC A?PRGJ?EC, GJJSQRP?RCB GL Figure 33.14. 5FC J?ASL?C ?PC P?LBMKJW QA?RRCPCB RFPMSEFMSR RFC RGQQSC ?LB RFC K?RPGV R?ICQ ML ? KGJIW MP QAPS@@CB ?NNC?P?LAC UGRF PMSRGLC FGQRMJMEGA?J QR?GLQ. 4F?PIQ F?TC A?PRGJ?EGLMSQ QICJCRMLQ, ?Q BMCQ LC?PJW RFC CLRGPC FSK?L QICJCRML BSPGLE ? QNCAGDGA NPC-@GPRF BCTCJMNKCLR?J QR?EC. " PCKL?LR MD RFGQ A?PRGJ?EC NCPQGQRQ GL RFC MSRCP NMPRGML MD RFC FSK?L LMQC. )W?JGLC A?PRGJ?EC GQ ?JQM DMSLB ?R RFC CLBQ MD JMLE @MLCQ, PCBSAGLE DPGARGML ?LB ASQFGMLGLE RFC ?PRGASJ?RGMLQ MD RFCQC @MLCQ.
!@>LI< 33.14 (V>IFKB @>OQFI>DB @LKPFPQP LC > J>QOFU TFQE @BIIP @>IIBA @ELKAOL@VQBP BJ?BAABA FK FQ. 4EB @ELKAOL@VQBP BUFPQ FK @>SFQFBP FK QEB J>QOFU @>IIBA I>@RK>B.
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tQRGDD.u B43+ #MLC, MP MQQCMSQ RGQQSC, GQ ? AMLLCARGTC RGQQSC RF?R F?Q ? J?PEC ?KMSLR MD RUM BGDDCPCLR RWNCQ MD K?RPGV K?RCPG?J. 5FC MPE?LGA K?RPGV GQ QGKGJ?P RM RFC K?RPGV K?RCPG?J DMSLB GL MRFCP AMLLCARGTC RGQQSCQ, GLAJSBGLE QMKC ?KMSLR MD AMJJ?ECL ?LB CJ?QRGA DG@CPQ. 5FGQ EGTCQ QRPCLERF ?LB DJCVG@GJGRW RM RFC RGQQSC. 5FC GLMPE?LGA K?RPGV AMLQGQRQ MD KGLCP?J Q?JRQqKMQRJW A?JAGSK Q?JRQqRF?R EGTC RFC RGQQSC F?PBLCQQ. 8GRFMSR ?BCOS?RC
#(!04%2 33 X 4(% !.)-!, "/$9: "!3)# &/2- !.$ &5.#4)/.
945
MPE?LGA K?RCPG?J GL RFC K?RPGV, RFC RGQQSC @PC?IQ; UGRFMSR ?BCOS?RC GLMPE?LGA K?RCPG?J GL RFC K?RPGV, RFC RGQQSC @CLBQ. 5FCPC ?PC RFPCC RWNCQ MD ACJJQ GL @MLC: MQRCM@J?QRQ, MQRCMAWRCQ, ?LB MQRCMAJ?QRQ. 0QRCM@J?QRQ ?PC ?ARGTC GL K?IGLE @MLC DMP EPMURF ?LB PCKMBCJGLE. 0QRCM@J?QRQ BCNMQGR @MLC K?RCPG?J GLRM RFC K?RPGV ?LB, ?DRCP RFC K?RPGV QSPPMSLBQ RFCK, RFCW AMLRGLSC RM JGTC, @SR GL ? PCBSACB KCR?@MJGA QR?RC ?Q MQRCMAWRCQ. 0QRCMAWRCQ ?PC DMSLB GL J?ASL?C MD RFC @MLC. 0QRCMAJ?QRQ ?PC ?ARGTC GL @PC?IGLE BMUL @MLC DMP @MLC PCKMBCJGLE, ?LB RFCW NPMTGBC ?AACQQ RM A?JAGSK QRMPCB GL RGQQSCQ. 0QRCMAJ?QRQ ?PC SQS?JJW DMSLB ML RFC QSPD?AC MD RFC RGQQSC. #MLC A?L @C BGTGBCB GLRM RUM RWNCQ: AMKN?AR ?LB QNMLEW. $MKN?AR @MLC GQ DMSLB GL RFC QF?DR (MP BG?NFWQGQ) MD ? JMLE @MLC ?LB RFC QSPD?AC MD RFC DJ?R @MLCQ, UFGJC QNMLEW @MLC GQ DMSLB GL RFC CLB (MP CNGNFWQGQ) MD ? JMLE @MLC. $MKN?AR @MLC GQ MPE?LGXCB GLRM QS@SLGRQ A?JJCB osteons, ?Q GJJSQRP?RCB GL Figure 33.15. " @JMMB TCQQCJ ?LB ? LCPTC ?PC DMSLB GL RFC ACLRCP MD RFC QRPSARSPC UGRFGL RFC )?TCPQG?L A?L?J, UGRF P?BG?RGLE AGPAJCQ MD J?ASL?C ?PMSLB GR ILMUL ?Q J?KCJJ?C. 5FC U?TW JGLCQ QCCL @CRUCCL RFC J?ASL?C ?PC KGAPMAF?LLCJQ A?JJCB canaliculi; RFCW AMLLCAR RFC J?ASL?C RM ?GB BGDDSQGML @CRUCCL RFC ACJJQ. 4NMLEW @MLC GQ K?BC MD RGLW NJ?RCQ A?JJCB trabeculae RFCQC NJ?RCQ QCPTC ?Q QRPSRQ RM EGTC RFC QNMLEW @MLC QRPCLERF. 0TCP RGKC, RFCQC NJ?RCQ A?L @PC?I A?SQGLE RFC @MLC RM @CAMKC JCQQ PCQGJGCLR. #MLC RGQQSC DMPKQ RFC GLRCPL?J QICJCRML MD TCPRC@P?RC ?LGK?JQ, NPMTGBGLE QRPSARSPC RM RFC ?LGK?J ?LB NMGLRQ MD ?RR?AFKCLR DMP RCLBMLQ.
946
#(!04%2 33 X 4(% !.)-!, "/$9: "!3)# &/2- !.$ &5.#4)/.
!@>LI< 33.15 (>) #LJM>@Q ?LKB FP > ABKPB J>QOFU LK QEB LRQBO PROC>@B LC ?LKB. 3MLKDV ?LKB, FKPFAB QEB @LJM>@Q ?LKB, FP MLOLRP TFQE TB?-IFHB QO>?B@RI>B. (?) #LJM>@Q ?LKB FP LOD>KFWBA FKQL OFKDP @>IIBA LPQBLKP. "ILLA SBPPBIP, KBOSBP, >KA IVJME>QF@ SBPPBIP >OB CLRKA FK QEB @BKQO>I (>SBOPF>K @>K>I. 2FKDP LC I>JBII>B PROOLRKA QEB (>SBOPF>K @>K>I. "BQTBBK QEB I>JBII>B >OB @>SFQFBP @>IIBA I>@RK>B. #>K>IF@RIF >OB JF@OL@E>KKBIP @LKKB@QFKD QEB I>@RK>B QLDBQEBO. (@) /PQBL?I>PQP PROOLRKA QEB BUQBOFLO LC QEB ?LKB. /PQBL@I>PQP ?LOB QRKKBIP FKQL QEB ?LKB >KA LPQBL@VQBP >OB CLRKA FK QEB I>@RK>B.
A*/548+ "/88:+ "BGNMQC RGQQSC, MP D?R RGQQSC, GQ AMLQGBCPCB ? AMLLCARGTC RGQQSC CTCL RFMSEF GR BMCQ LMR F?TC DG@PM@J?QRQ MP ? PC?J K?RPGV ?LB MLJW F?Q ? DCU DG@CPQ. "BGNMQC RGQQSC GQ K?BC SN MD ACJJQ A?JJCB ?BGNMAWRCQ RF?R AMJJCAR ?LB QRMPC D?R GL RFC DMPK MD RPGEJWACPGBCQ, DMP CLCPEW KCR?@MJGQK. "BGNMQC RGQQSCQ ?BBGRGML?JJW QCPTC ?Q GLQSJ?RGML RM FCJN K?GLR?GL @MBW RCKNCP?RSPCQ, ?JJMUGLE ?LGK?JQ RM @C CLBMRFCPKGA, ?LB RFCW DSLARGML ?Q ASQFGMLGLE ?E?GLQR B?K?EC RM @MBW MPE?LQ. 6LBCP ? KGAPMQAMNC, ?BGNMQC RGQQSC ACJJQ ?NNC?P CKNRW BSC RM RFC CVRP?ARGML MD D?R BSPGLE RFC NPMACQQGLE MD RFC K?RCPG?J DMP TGCUGLE, ?Q QCCL GL Figure 33.16. 5FC RFGL JGLCQ GL RFC GK?EC ?PC RFC ACJJ KCK@P?LCQ, ?LB RFC LSAJCG ?PC RFC QK?JJ, @J?AI BMRQ ?R RFC CBECQ MD RFC ACJJQ.
#(!04%2 33 X 4(% !.)-!, "/$9: "!3)# &/2- !.$ &5.#4)/.
947
!@>LI< 33.16 !AFMLPB FP > @LKKB@QFSB QFPPRB FP J>AB RM LC @BIIP @>IIBA >AFML@VQBP. !AFML@VQBP E>SB PJ>II KR@IBF IL@>IFWBA >Q QEB @BII BADB.
B144* #JMMB GQ AMLQGBCPCB ? AMLLCARGTC RGQQSC @CA?SQC GR F?Q ? K?RPGV, ?Q QFMUL GL Figure 33.17. 5FC JGTGLE ACJJ RWNCQ ?PC PCB @JMMB ACJJQ (3#$), ?JQM A?JJCB CPWRFPMAWRCQ, ?LB UFGRC @JMMB ACJJQ (8#$), ?JQM A?JJCB JCSIMAWRCQ. 5FC DJSGB NMPRGML MD UFMJC @JMMB, GRQ K?RPGV, GQ AMKKMLJW A?JJCB NJ?QK?.
!@>LI< 33.17 "ILLA FP > @LKKB@QFSB QFPPRB QE>Q E>P > CIRFA J>QOFU, @>IIBA MI>PJ>, >KA KL CF?BOP. %OVQEOL@VQBP (OBA ?ILLA @BIIP), QEB MOBALJFK>KQ @BII QVMB, >OB FKSLISBA FK QEB QO>KPMLOQ LC LUVDBK >KA @>O?LK AFLUFAB. !IPL MOBPBKQ >OB S>OFLRP IBRHL@VQBP (TEFQB ?ILLA @BIIP) FKSLISBA FK FJJRKB OBPMLKPB.
5FC ACJJ DMSLB GL EPC?RCQR ?@SLB?LAC GL @JMMB GQ RFC CPWRFPMAWRC. &PWRFPMAWRCQ ?PC AMSLRCB GL KGJJGMLQ GL ? @JMMB Q?KNJC: RFC ?TCP?EC LSK@CP MD PCB @JMMB ACJJQ GL NPGK?RCQ GQ 4.7 RM 5.5 KGJJGML ACJJQ NCP KGAPMJGRCP. &PWRFPMAWRCQ ?PC AMLQGQRCLRJW RFC Q?KC QGXC GL ? QNCAGCQ, @SR T?PW GL QGXC @CRUCCL QNCAGCQ. 'MP CV?KNJC, RFC ?TCP?EC BG?KCRCP MD ? NPGK?RC PCB @JMMB ACJJ GQ 7.5 ]J, ? BME GQ AJMQC ?R 7.0 ]J, @SR ? A?RsQ 3#$ BG?KCRCP GQ 5.9 ]J. 4FCCN CPWRFPMAWRCQ ?PC CTCL QK?JJCP ?R 4.6 ]J. .?KK?JG?L CPWRFPMAWRCQ JMQC RFCGP LSAJCG ?LB KGRMAFMLBPG? UFCL RFCW ?PC PCJC?QCB DPMK RFC @MLC K?PPMU UFCPC RFCW ?PC K?BC. 'GQF, ?KNFG@G?L, ?LB ?TG?L PCB @JMMB ACJJQ K?GLR?GL RFCGP LSAJCG ?LB KGRMAFMLBPG? RFPMSEFMSR RFC ACJJsQ JGDC. 5FC NPGLAGN?J HM@ MD ?L CPWRFPMAWRC GQ RM A?PPW ?LB BCJGTCP MVWECL RM RFC RGQQSCQ. -CSIMAWRCQ ?PC RFC NPCBMKGL?LR UFGRC @JMMB ACJJQ DMSLB GL RFC NCPGNFCP?J @JMMB. -CSIMAWRCQ ?PC AMSLRCB GL RFC RFMSQ?LBQ GL RFC @JMMB UGRF KC?QSPCKCLRQ CVNPCQQCB ?Q P?LECQ: NPGK?RC AMSLRQ P?LEC DPMK 4,800 RM 10,800 ACJJQ NCP ]J, BMEQ DPMK 5,600 RM 19,200 ACJJQ NCP ]J, A?RQ DPMK 8,000 RM 25,000 ACJJQ NCP ]J, A?RRJC DPMK 4,000 RM 12,000 ACJJQ NCP ]J, ?LB NGEQ DPMK 11,000 RM 22,000 ACJJQ NCP ]J. -WKNFMAWRCQ DSLARGML NPGK?PGJW GL RFC GKKSLC PCQNMLQC RM DMPCGEL ?LRGECLQ MP K?RCPG?J. %GDDCPCLR RWNCQ MD JWKNFMAWRCQ K?IC ?LRG@MBGCQ R?GJMPCB RM RFC DMPCGEL ?LRGECLQ ?LB AMLRPMJ RFC NPMBSARGML MD RFMQC ?LRG@MBGCQ. /CSRPMNFGJQ ?PC NF?EMAWRGA ACJJQ ?LB RFCW N?PRGAGN?RC GL MLC MD RFC C?PJW JGLCQ MD BCDCLQC ?E?GLQR KGAPM@G?J GLT?BCPQ, ?GBGLE GL RFC PCKMT?J MD @?ARCPG? RF?R F?Q CLRCPCB RFC @MBW. "LMRFCP JCSIMAWRC RF?R GQ DMSLB GL RFC NCPGNFCP?J @JMMB GQ RFC KMLMAWRC. .MLMAWRCQ EGTC PGQC RM NF?EMAWRGA K?APMNF?ECQ RF?R AJC?L SN BC?B ?LB B?K?ECB ACJJQ GL RFC @MBW, UFCRFCP RFCW ?PC DMPCGEL MP DPMK RFC FMQR
948
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?LGK?J. 5UM ?BBGRGML?J JCSIMAWRCQ GL RFC @JMMB ?PC CMQGLMNFGJQ ?LB @?QMNFGJQq@MRF FCJN RM D?AGJGR?RC RFC GLDJ?KK?RMPW PCQNMLQC. 5FC QJGEFRJW EP?LSJ?P K?RCPG?J ?KMLE RFC ACJJQ GQ ? AWRMNJ?QKGA DP?EKCLR MD ? ACJJ GL RFC @MLC K?PPMU. 5FGQ GQ A?JJCB ? NJ?RCJCR MP RFPMK@MAWRC. 1J?RCJCRQ N?PRGAGN?RC GL RFC QR?ECQ JC?BGLE SN RM AM?ESJ?RGML MD RFC @JMMB RM QRMN @JCCBGLE RFPMSEF B?K?ECB @JMMB TCQQCJQ. #JMMB F?Q ? LSK@CP MD DSLARGMLQ, @SR NPGK?PGJW GR RP?LQNMPRQ K?RCPG?J RFPMSEF RFC @MBW RM @PGLE LSRPGCLRQ RM ACJJQ ?LB PCKMTC U?QRC K?RCPG?J DPMK RFCK.
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5FCPC ?PC RFPCC RWNCQ MD KSQAJC GL ?LGK?J @MBGCQ: QKMMRF, QICJCR?J, ?LB A?PBG?A. 5FCW BGDDCP @W RFC NPCQCLAC MP ?@QCLAC MD QRPG?RGMLQ MP @?LBQ, RFC LSK@CP ?LB JMA?RGML MD LSAJCG, UFCRFCP RFCW ?PC TMJSLR?PGJW MP GLTMJSLR?PGJW AMLRPMJJCB, ?LB RFCGP JMA?RGML UGRFGL RFC @MBW. Table 33.4 QSKK?PGXCQ RFCQC BGDDCPCLACQ.
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!2449. M:8)1+ 4KMMRF KSQAJC BMCQ LMR F?TC QRPG?RGMLQ GL GRQ ACJJQ. *R F?Q ? QGLEJC, ACLRP?JJW JMA?RCB LSAJCSQ, ?Q QFMUL GL Figure 33.18. $MLQRPGARGML MD QKMMRF KSQAJC MAASPQ SLBCP GLTMJSLR?PW, ?SRMLMKGA LCPTMSQ AMLRPMJ ?LB GL PCQNMLQC RM JMA?J AMLBGRGMLQ GL RFC RGQQSCQ. 4KMMRF KSQAJC RGQQSC GQ ?JQM A?JJCB LML-QRPG?RCB ?Q GR J?AIQ RFC @?LBCB ?NNC?P?LAC MD QICJCR?J ?LB A?PBG?A KSQAJC. 5FC U?JJQ MD @JMMB TCQQCJQ, RFC RS@CQ MD RFC BGECQRGTC QWQRCK, ?LB RFC RS@CQ MD RFC PCNPMBSARGTC QWQRCKQ ?PC AMKNMQCB MD KMQRJW QKMMRF KSQAJC.
!@>LI< 33.18 3JLLQE JRP@IB @BIIP AL KLQ E>SB PQOF>QFLKP, TEFIB PHBIBQ>I JRP@IB @BIIP AL. #>OAF>@ JRP@IB @BIIP E>SB PQOF>QFLKP, ?RQ, RKIFHB QEB JRIQFKR@IB>QB PHBIBQ>I @BIIP, QEBV E>SB LKIV LKB KR@IBRP. #>OAF>@ JRP@IB QFPPRB >IPL E>P FKQBO@>I>QBA AFP@P, PMB@F>IFWBA OBDFLKP ORKKFKD >ILKD QEB MI>PJ> JBJ?O>KB QE>Q GLFK >AG>@BKQ @>OAF>@ JRP@IB @BIIP >KA >PPFPQ FK M>PPFKD >K BIB@QOF@>I FJMRIPB COLJ @BII QL @BII.
!0+1+9'1 M:8)1+ 4ICJCR?J KSQAJC F?Q QRPG?RGMLQ ?APMQQ GRQ ACJJQ A?SQCB @W RFC ?PP?LECKCLR MD RFC AMLRP?ARGJC NPMRCGLQ ?ARGL ?LB KWMQGL. 5FCQC KSQAJC ACJJQ ?PC PCJ?RGTCJW JMLE ?LB F?TC KSJRGNJC LSAJCG ?JMLE RFC CBEC MD RFC ACJJ. 4ICJCR?J KSQAJC GQ SLBCP TMJSLR?PW, QMK?RGA LCPTMSQ QWQRCK AMLRPMJ ?LB GQ DMSLB GL RFC KSQAJCQ RF?R KMTC @MLCQ. Figure 33.18 GJJSQRP?RCQ RFC FGQRMJMEW MD QICJCR?J KSQAJC. C'7*/') M:8)1+ $?PBG?A KSQAJC, QFMUL GL Figure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/CPTMSQ RGQQSCQ ?PC K?BC MD ACJJQ QNCAG?JGXCB RM PCACGTC ?LB RP?LQKGR CJCARPGA?J GKNSJQCQ DPMK QNCAGDGA ?PC?Q MD RFC @MBW ?LB RM QCLB RFCK RM QNCAGDGA JMA?RGMLQ GL RFC @MBW. 5FC K?GL ACJJ MD RFC LCPTMSQ QWQRCK
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949
GQ RFC LCSPML, GJJSQRP?RCB GL Figure 33.19. 5FC J?PEC QRPSARSPC UGRF ? ACLRP?J LSAJCSQ GQ RFC ACJJ @MBW MD RFC LCSPML. 1PMHCARGMLQ DPMK RFC ACJJ @MBW ?PC CGRFCP BCLBPGRCQ QNCAG?JGXCB GL PCACGTGLE GLNSR MP ? QGLEJC ?VML QNCAG?JGXCB GL RP?LQKGRRGLE GKNSJQCQ. 4MKC EJG?J ACJJQ ?PC ?JQM QFMUL. "QRPMAWRCQ PCESJ?RC RFC AFCKGA?J CLTGPMLKCLR MD RFC LCPTC ACJJ, ?LB MJGEMBCLBPMAWRCQ GLQSJ?RC RFC ?VML QM RFC CJCARPGA?J LCPTC GKNSJQC GQ RP?LQDCPPCB KMPC CDDGAGCLRJW. 0RFCP EJG?J ACJJQ RF?R ?PC LMR QFMUL QSNNMPR RFC LSRPGRGML?J ?LB U?QRC PCOSGPCKCLRQ MD RFC LCSPML. 4MKC MD RFC EJG?J ACJJQ ?PC NF?EMAWRGA ?LB PCKMTC BC@PGQ MP B?K?ECB ACJJQ DPMK RFC RGQQSC. " LCPTC AMLQGQRQ MD LCSPMLQ ?LB EJG?J ACJJQ.
!@>LI< 33.19 4EB KBROLK E>P MOLGB@QFLKP @>IIBA ABKAOFQBP QE>Q OB@BFSB PFDK>IP >KA MOLGB@QFLKP @>IIBA >ULKP QE>Q PBKA PFDK>IP. !IPL PELTK >OB QTL QVMBP LC DIF>I @BIIP: >PQOL@VQBP OBDRI>QB QEB @EBJF@>I BKSFOLKJBKQ LC QEB KBOSB @BII, >KA LIFDLABKAOL@VQBP FKPRI>QB QEB >ULK PL QEB BIB@QOF@>I KBOSB FJMRIPB FP QO>KPCBOOBA JLOB BCCF@FBKQIV.
$JGAI RFPMSEF RFC interactive review (http://openstaxcollege.org/l/tissues) RM JC?PL KMPC ?@MSR CNGRFCJG?J RGQQSCQ.
+8K?FCF>@JK
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
950
#(!04%2 33 X 4(% !.)-!, "/$9: "!3)# &/2- !.$ &5.#4)/.
33.3 R #FD<FJK8J@J
#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CDGLC FMKCMQR?QGQ z %CQAPG@C RFC D?ARMPQ ?DDCARGLE FMKCMQR?QGQ z %GQASQQ NMQGRGTC ?LB LCE?RGTC DCCB@?AI KCAF?LGQKQ SQCB GL FMKCMQR?QGQ z %CQAPG@C RFCPKMPCESJ?RGML MD CLBMRFCPKGA ?LB CARMRFCPKGA ?LGK?JQ "LGK?J MPE?LQ ?LB MPE?L QWQRCKQ AMLQR?LRJW ?BHSQR RM GLRCPL?J ?LB CVRCPL?J AF?LECQ RFPMSEF ? NPMACQQ A?JJCB FMKCMQR?QGQ (tQRC?BW QR?RCu). 5FCQC AF?LECQ KGEFR @C GL RFC JCTCJ MD EJSAMQC MP A?JAGSK GL @JMMB MP GL CVRCPL?J RCKNCP?RSPCQ. Homeostasis KC?LQ RM K?GLR?GL BWL?KGA COSGJG@PGSK GL RFC @MBW. *R GQ BWL?KGA @CA?SQC GR GQ AMLQR?LRJW ?BHSQRGLE RM RFC AF?LECQ RF?R RFC @MBWsQ QWQRCKQ CLAMSLRCP. *R GQ COSGJG@PGSK @CA?SQC @MBW DSLARGMLQ ?PC ICNR UGRFGL QNCAGDGA P?LECQ. &TCL ?L ?LGK?J RF?R GQ ?NN?PCLRJW GL?ARGTC GQ K?GLR?GLGLE RFGQ FMKCMQR?RGA COSGJG@PGSK.
#FD<FJK8K@: +IF:<JJ
5FC EM?J MD FMKCMQR?QGQ GQ RFC K?GLRCL?LAC MD COSGJG@PGSK ?PMSLB ? NMGLR MP T?JSC A?JJCB ? set point. 8FGJC RFCPC ?PC LMPK?J DJSARS?RGMLQ DPMK RFC QCR NMGLR, RFC @MBWsQ QWQRCKQ UGJJ SQS?JJW ?RRCKNR RM EM @?AI RM RFGQ NMGLR. " AF?LEC GL RFC GLRCPL?J MP CVRCPL?J CLTGPMLKCLR GQ A?JJCB ? QRGKSJSQ ?LB GQ BCRCARCB @W ? PCACNRMP; RFC PCQNMLQC MD RFC QWQRCK GQ RM ?BHSQR RFC BCTG?RGML N?P?KCRCP RMU?PB RFC QCR NMGLR. 'MP GLQR?LAC, GD RFC @MBW @CAMKCQ RMM U?PK, ?BHSQRKCLRQ ?PC K?BC RM AMMJ RFC ?LGK?J. *D RFC @JMMBsQ EJSAMQC PGQCQ ?DRCP ? KC?J, ?BHSQRKCLRQ ?PC K?BC RM JMUCP RFC @JMMB EJSAMQC JCTCJ @W ECRRGLE RFC LSRPGCLR GLRM RGQQSCQ RF?R LCCB GR MP RM QRMPC GR DMP J?RCP SQC.
CFEKIFC F= #FD<FJK8J@J
8FCL ? AF?LEC MAASPQ GL ?L ?LGK?Js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negative feedback loop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igure 33.20. )MUCTCP, GD ?L ?LGK?J F?Q LMR C?RCL ?LB @JMMB EJSAMQC JCTCJQ BCAPC?QC, RFGQ GQ QCLQCB GL ?LMRFCP EPMSN MD ACJJQ GL RFC N?LAPC?Q, ?LB RFC FMPKMLC EJSA?EML GQ PCJC?QCB A?SQGLE EJSAMQC JCTCJQ RM GLAPC?QC. 5FGQ GQ QRGJJ ? LCE?RGTC DCCB@?AI JMMN, @SR LMR GL RFC BGPCARGML CVNCARCB @W RFC SQC MD RFC RCPK tLCE?RGTC.u "LMRFCP CV?KNJC MD ?L GLAPC?QC ?Q ? PCQSJR MD RFC DCCB@?AI JMMN GQ RFC AMLRPMJ MD @JMMB A?JAGSK. *D A?JAGSK JCTCJQ BCAPC?QC, QNCAG?JGXCB ACJJQ GL RFC N?P?RFWPMGB EJ?LB QCLQC RFGQ ?LB PCJC?QC N?P?RFWPMGB FMPKMLC (15)), A?SQGLE ?L GLAPC?QCB ?@QMPNRGML MD A?JAGSK RFPMSEF RFC GLRCQRGLCQ ?LB IGBLCWQ ?LB, NMQQG@JW, RFC @PC?IBMUL MD @MLC GL MPBCP RM JG@CP?RC A?JAGSK. 5FC CDDCARQ MD 15) ?PC RM P?GQC @JMMB JCTCJQ MD RFC CJCKCLR. /CE?RGTC DCCB@?AI JMMNQ ?PC RFC NPCBMKGL?LR KCAF?LGQK SQCB GL FMKCMQR?QGQ.
#(!04%2 33 X 4(% !.)-!, "/$9: "!3)# &/2- !.$ &5.#4)/.
951
!@>LI< 33.20 "ILLA PRD>O IBSBIP >OB @LKQOLIIBA ?V > KBD>QFSB CBBA?>@H ILLM. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V *LK 3RIIFS>K)
P48/9/;+ F++*(')0 L445 " positive feedback loop K?GLR?GLQ RFC BGPCARGML MD RFC QRGKSJSQ, NMQQG@JW ?AACJCP?RGLE GR. 'CU CV?KNJCQ MD NMQGRGTC DCCB@?AI JMMNQ CVGQR GL ?LGK?J @MBGCQ, @SR MLC GQ DMSLB GL RFC A?QA?BC MD AFCKGA?J PC?ARGMLQ RF?R PCQSJR GL @JMMB AJMRRGLE, MP AM?ESJ?RGML. "Q MLC AJMRRGLE D?ARMP GQ ?ARGT?RCB, GR ?ARGT?RCQ RFC LCVR D?ARMP GL QCOSCLAC SLRGJ ? DG@PGL AJMR GQ ?AFGCTCB. 5FC BGPCARGML GQ K?GLR?GLCB, LMR AF?LECB, QM RFGQ GQ NMQGRGTC DCCB@?AI. "LMRFCP CV?KNJC MD NMQGRGTC DCCB@?AI GQ SRCPGLC AMLRP?ARGMLQ BSPGLE AFGJB@GPRF, ?Q GJJSQRP?RCB GL Figure 33.21. 5FC FMPKMLC MVWRMAGL, K?BC @W RFC CLBMAPGLC QWQRCK, QRGKSJ?RCQ RFC AMLRP?ARGML MD RFC SRCPSQ. 5FGQ NPMBSACQ N?GL QCLQCB @W RFC LCPTMSQ QWQRCK. *LQRC?B MD JMUCPGLE RFC MVWRMAGL ?LB A?SQGLE RFC N?GL RM QS@QGBC, KMPC MVWRMAGL GQ NPMBSACB SLRGJ RFC AMLRP?ARGMLQ ?PC NMUCPDSJ CLMSEF RM NPMBSAC AFGJB@GPRF.
!@>LI< 33.21 4EB ?FOQE LC > ERJ>K FKC>KQ FP QEB OBPRIQ LC MLPFQFSB CBBA?>@H.
3Q>QB TEBQEBO B>@E LC QEB CLIILTFKD MOL@BPPBP FP OBDRI>QBA ?V > MLPFQFSB CBBA?>@H ILLM LO > KBD>QFSB CBBA?>@H ILLM. >. ! MBOPLK CBBIP P>QF>QBA >CQBO B>QFKD > I>ODB JB>I. ?. 4EB ?ILLA E>P MIBKQV LC OBA ?ILLA @BIIP. !P > OBPRIQ, BOVQEOLMLFBQFK, > ELOJLKB QE>Q PQFJRI>QBP QEB MOLAR@QFLK LC KBT OBA ?ILLA @BIIP, FP KL ILKDBO OBIB>PBA COLJ QEB HFAKBV.
952
#(!04%2 33 X 4(% !.)-!, "/$9: "!3)# &/2- !.$ &5.#4)/.
!+9 P4/39 *R GQ NMQQG@JC RM ?BHSQR ? QWQRCKsQ QCR NMGLR. 8FCL RFGQ F?NNCLQ, RFC DCCB@?AI JMMN UMPIQ RM K?GLR?GL RFC LCU QCRRGLE. "L CV?KNJC MD RFGQ GQ @JMMB NPCQQSPC: MTCP RGKC, RFC LMPK?J MP QCR NMGLR DMP @JMMB NPCQQSPC A?L GLAPC?QC ?Q ? PCQSJR MD AMLRGLSCB GLAPC?QCQ GL @JMMB NPCQQSPC. 5FC @MBW LM JMLECP PCAMELGXCQ RFC CJCT?RGML ?Q ?@LMPK?J ?LB LM ?RRCKNR GQ K?BC RM PCRSPL RM RFC JMUCP QCR NMGLR. 5FC PCQSJR GQ RFC K?GLRCL?LAC MD ?L CJCT?RCB @JMMB NPCQQSPC RF?R A?L F?TC F?PKDSJ CDDCARQ ML RFC @MBW. .CBGA?RGML A?L JMUCP @JMMB NPCQQSPC ?LB JMUCP RFC QCR NMGLR GL RFC QWQRCK RM ? KMPC FC?JRFW JCTCJ. 5FGQ GQ A?JJCB ? NPMACQQ MD alteration MD RFC QCR NMGLR GL ? DCCB@?AI JMMN. $F?LECQ A?L @C K?BC GL ? EPMSN MD @MBW MPE?L QWQRCKQ GL MPBCP RM K?GLR?GL ? QCR NMGLR GL ?LMRFCP QWQRCK. 5FGQ GQ A?JJCB acclimatization. 5FGQ MAASPQ, DMP GLQR?LAC, UFCL ?L ?LGK?J KGEP?RCQ RM ? FGEFCP ?JRGRSBC RF?L GR GQ ?AASQRMKCB RM. *L MPBCP RM ?BHSQR RM RFC JMUCP MVWECL JCTCJQ ?R RFC LCU ?JRGRSBC, RFC @MBW GLAPC?QCQ RFC LSK@CP MD PCB @JMMB ACJJQ AGPASJ?RGLE GL RFC @JMMB RM CLQSPC ?BCOS?RC MVWECL BCJGTCPW RM RFC RGQQSCQ. "LMRFCP CV?KNJC MD ?AAJGK?RGX?RGML GQ ?LGK?JQ RF?R F?TC QC?QML?J AF?LECQ GL RFCGP AM?RQ: ? FC?TGCP AM?R GL RFC UGLRCP CLQSPCQ ?BCOS?RC FC?R PCRCLRGML, ?LB ? JGEFR AM?R GL QSKKCP ?QQGQRQ GL ICCNGLE @MBW RCKNCP?RSPC DPMK PGQGLE RM F?PKDSJ JCTCJQ.
'CCB@?AI KCAF?LGQKQ A?L @C SLBCPQRMMB GL RCPKQ MD BPGTGLE ? P?AC A?P ?JMLE ? RP?AI: U?RAF ? QFMPR video lesson (http://openstaxcollege.org/l/feedback_loops) ML NMQGRGTC ?LB LCE?RGTC DCCB@?AI JMMNQ.
#FD<FJK8J@J: /?<IDFI<>LC8K@FE
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
8?RAF RFGQ Discovery Channel video (http://openstaxcollege.org/l/thermoregulate) RFCPKMPCESJ?RGML RM QCC GJJSQRP?RGMLQ MD RFGQ NPMACQQ GL ? T?PGCRW MD ?LGK?JQ.
ML
E;FK?<IDJ 8E; :KFK?<IDJ

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
#(!04%2 33 X 4(% !.)-!, "/$9: "!3)# &/2- !.$ &5.#4)/.
953
)C?R A?L @C CVAF?LECB @CRUCCL ?L ?LGK?J ?LB GRQ CLTGPMLKCLR RFPMSEF DMSP KCAF?LGQKQ: P?BG?RGML, CT?NMP?RGML, AMLTCARGML, ?LB AMLBSARGML (Figure 33.22). 3?BG?RGML GQ RFC CKGQQGML MD CJCARPMK?ELCRGA tFC?Ru U?TCQ. )C?R AMKCQ DPMK RFC QSL GL RFGQ K?LLCP ?LB P?BG?RCQ DPMK BPW QIGL RFC Q?KC U?W. )C?R A?L @C PCKMTCB UGRF JGOSGB DPMK ? QSPD?AC BSPGLE CT?NMP?RGML. 5FGQ MAASPQ UFCL ? K?KK?J QUC?RQ. $MLTCARGML ASPPCLRQ MD ?GP PCKMTC FC?R DPMK RFC QSPD?AC MD BPW QIGL ?Q RFC ?GP N?QQCQ MTCP GR. )C?R UGJJ @C AMLBSARCB DPMK MLC QSPD?AC RM ?LMRFCP BSPGLE BGPCAR AMLR?AR UGRF RFC QSPD?ACQ, QSAF ?Q ?L ?LGK?J PCQRGLE ML ? U?PK PMAI.
!@>LI< 33.22 (B>Q @>K ?B BU@E>KDBA ?V CLRO JB@E>KFPJP: (>) O>AF>QFLK, (?) BS>MLO>QFLK, (@) @LKSB@QFLK, LO (A) @LKAR@QFLK. (@OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V x+RIIBWy/&IF@HO; @OBAFQ @: JLAFCF@>QFLK LC TLOH ?V #E>A 2LPBKQE>I; @OBAFQ A: JLAFCF@>QFLK LC TLOH ?V xPQ>@BV.Ay/&IF@HO)
#<8K CFEJ<IM8K@FE 8E; D@JJ@G8K@FE

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tEMMQC @SKNQ,u A?SQGLE QK?JJ F?GPQ RM QR?LB SN UFCL RFC GLBGTGBS?J GQ AMJB; RFGQ F?Q RFC GLRCLBCB CDDCAR MD GLAPC?QGLE @MBW RCKNCP?RSPC. .?KK?JQ SQC J?WCPQ MD D?R RM ?AFGCTC RFC Q?KC CLB. -MQQ MD QGELGDGA?LR ?KMSLRQ MD @MBW D?R UGJJ AMKNPMKGQC ?L GLBGTGBS?JsQ ?@GJGRW RM AMLQCPTC FC?R. &LBMRFCPKQ SQC RFCGP AGPASJ?RMPW QWQRCKQ RM FCJN K?GLR?GL @MBW RCKNCP?RSPC. 7?QMBGJ?RGML @PGLEQ KMPC @JMMB ?LB FC?R RM RFC @MBW QSPD?AC, D?AGJGR?RGLE P?BG?RGML ?LB CT?NMP?RGTC FC?R JMQQ, UFGAF FCJNQ RM AMMJ RFC @MBW. 7?QMAMLQRPGARGML PCBSACQ @JMMB DJMU GL NCPGNFCP?J @JMMB TCQQCJQ, DMPAGLE @JMMB RMU?PB RFC AMPC ?LB RFC TGR?J MPE?LQ DMSLB RFCPC, ?LB AMLQCPTGLE FC?R. 4MKC ?LGK?JQ F?TC ?B?NRGMLQ RM RFCGP AGPASJ?RMPW QWQRCK RF?R CL?@JC RFCK RM RP?LQDCP FC?R DPMK ?PRCPGCQ RM TCGLQ, U?PKGLE @JMMB PCRSPLGLE RM RFC FC?PR. 5FGQ GQ A?JJCB ? AMSLRCPASPPCLR FC?R CVAF?LEC; GR NPCTCLRQ RFC AMJB TCLMSQ @JMMB DPMK AMMJGLE RFC FC?PR ?LB MRFCP GLRCPL?J MPE?LQ. 5FGQ ?B?NRGML A?L @C QFSR BMUL GL QMKC ?LGK?JQ RM NPCTCLR MTCPFC?RGLE RFC GLRCPL?J MPE?LQ. 5FC AMSLRCPASPPCLR ?B?NRGML GQ DMSLB GL K?LW ?LGK?JQ, GLAJSBGLE BMJNFGLQ, QF?PIQ, @MLW DGQF, @CCQ, ?LB FSKKGLE@GPBQ. *L AMLRP?QR, QGKGJ?P ?B?NR?RGMLQ A?L FCJN AMMJ CLBMRFCPKQ UFCL LCCBCB, QSAF ?Q BMJNFGL DJSICQ ?LB CJCNF?LR C?PQ. 4MKC CARMRFCPKGA ?LGK?JQ SQC AF?LECQ GL RFCGP @CF?TGMP RM FCJN PCESJ?RC @MBW RCKNCP?RSPC. 'MP CV?KNJC, ? BCQCPR CARMRFCPKGA ?LGK?J K?W QGKNJW QCCI AMMJCP ?PC?Q BSPGLE RFC FMRRCQR N?PR MD RFC B?W GL RFC BCQCPR RM ICCN DPMK ECRRGLE RMM U?PK. 5FC Q?KC ?LGK?JQ K?W AJGK@ MLRM PMAIQ RM A?NRSPC FC?R BSPGLE ? AMJB BCQCPR LGEFR. 4MKC ?LGK?JQ QCCI U?RCP RM ?GB CT?NMP?RGML GL AMMJGLE RFCK, ?Q QCCL UGRF PCNRGJCQ. 0RFCP CARMRFCPKQ SQC EPMSN ?ARGTGRW QSAF ?Q RFC ?ARGTGRW MD @CCQ RM U?PK ? FGTC RM QSPTGTC UGLRCP.
954
#(!04%2 33 X 4(% !.)-!, "/$9: "!3)# &/2- !.$ &5.#4)/.
.?LW ?LGK?JQ, CQNCAG?JJW K?KK?JQ, SQC KCR?@MJGA U?QRC FC?R ?Q ? FC?R QMSPAC. 8FCL KSQAJCQ ?PC AMLRP?ARCB, KMQR MD RFC CLCPEW DPMK RFC "51 SQCB GL KSQAJC ?ARGMLQ GQ U?QRCB CLCPEW RF?R RP?LQJ?RCQ GLRM FC?R. 4CTCPC AMJB CJGAGRQ ? QFGTCPGLE PCDJCV RF?R ECLCP?RCQ FC?R DMP RFC @MBW. .?LW QNCAGCQ ?JQM F?TC ? RWNC MD ?BGNMQC RGQQSC A?JJCB @PMUL D?R RF?R QNCAG?JGXCQ GL ECLCP?RGLE FC?R.
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5FC LCPTMSQ QWQRCK GQ GKNMPR?LR RM thermoregulation, ?Q GJJSQRP?RCB GL Figure 33.22. 5FC NPMACQQCQ MD FMKCMQR?QGQ ?LB RCKNCP?RSPC AMLRPMJ ?PC ACLRCPCB GL RFC FWNMRF?J?KSQ MD RFC ?BT?LACB ?LGK?J @P?GL.
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8::C@D8K@Q8K@FE ?JRCP?RGML GL ? @MBW QWQRCK GL PCQNMLQC RM CLTGPMLKCLR?J AF?LEC 8CK<I8K@FE AF?LEC MD RFC QCR NMGLR GL ? FMKCMQR?RGA QWQRCK 8GF;<D< GLEPMURF MD ?L ?LGK?Js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
956
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1. Figure 33.11 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR RWNCQ MD CNGRFCJG?J ACJJQ GQ D?JQC >. 4GKNJC AMJSKL?P CNGRFCJG?J ACJJQ JGLC RFC RGQQSC MD RFC JSLE. ?. 4GKNJC AS@MGB?J CNGRFCJG?J ACJJQ ?PC GLTMJTCB GL RFC DGJRCPGLE MD @JMMB GL RFC IGBLCW. @. 1QCSBMQRP?RGQDGCB AMJSKL?P CNGRFGJG? MAASP GL ? QGLEJC J?WCP, @SR RFC ?PP?LECKCLR MD LSAJCG K?ICQ GR ?NNC?P RF?R KMPC RF?L MLC J?WCP GQ NPCQCLR. A. 5P?LQGRGML?J CNGRFCJG? AF?LEC GL RFGAILCQQ BCNCLBGLE ML FMU DSJJ RFC @J?BBCP GQ. 2. Figure 33.21 4R?RC UFCRFCP C?AF MD RFC DMJJMUGLE NPMACQQCQ ?PC PCESJ?RCB @W ? NMQGRGTC DCCB@?AI JMMN MP ? LCE?RGTC DCCB@?AI JMMN. >. " NCPQML DCCJQ Q?RG?RCB ?DRCP C?RGLE ? J?PEC KC?J. ?. 5FC @JMMB F?Q NJCLRW MD PCB @JMMB ACJJQ. "Q ? PCQSJR, CPWRFPMNMGCRGL, ? FMPKMLC
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RF?R QRGKSJ?RCQ RFC NPMBSARGML MD LCU PCB @JMMB ACJJQ, GQ LM JMLECP PCJC?QCB DPMK RFC IGBLCW. 3. Figure 33.22 8FCL @?ARCPG? ?PC BCQRPMWCB @W JCSAIMAWRCQ, NWPMECLQ ?PC PCJC?QCB GLRM RFC @JMMB.
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4. 8FGAF RWNC MD ?LGK?J K?GLR?GLQ ? AMLQR?LR GLRCPL?J @MBW RCKNCP?RSPC >. CLBMRFCPK ?. CARMRFCPK @. AMCJMK?RC A. KCQMBCPK 5. 5FC QWKKCRPW DMSLB GL ?LGK?JQ RF?R KMTC QUGDRJW GQ >>>>>>>>. >. P?BG?J ?. @GJ?RCP?J @. QCOSCLRG?J A. GLRCPPSNRCB 6. 8F?R RCPK BCQAPG@CQ RFC AMLBGRGML MD ? BCQCPR KMSQC RF?R JMUCPQ GRQ KCR?@MJGA P?RC ?LB tQJCCNQu BSPGLE RFC FMR B?W >. RSPEGB ?. FG@CPL?RGML @. CQRGT?RGML A. LMPK?J QJCCN N?RRCPL 7. " NJ?LC RF?R BGTGBCQ ?L ?LGK?J GLRM COS?J PGEFR ?LB JCDR NMPRGMLQ GQ >>>>>>>>. >. BG?EML?J ?. KGBQ?EGRR?J @. AMPML?J A. RP?LQTCPQC 8. " NJ?LC RF?R BGTGBCQ ?L ?LGK?J GLRM BMPQ?J ?LB TCLRP?J NMPRGMLQ GQ >>>>>>>>. >. Q?EGRR?J ?. KGBQ?EGRR?J @. AMPML?J A. RP?LQTCPQC 9. 5FC NJCSP?J A?TGRW GQ ? N?PR MD UFGAF A?TGRW >. ?. @. A. BMPQ?J A?TGRW RFMP?AGA A?TGRW ?@BMKGL?J A?TGRW NCPGA?PBG?J A?TGRW 12. 8FGAF RWNC MD CNGRFCJG?J ACJJ GQ DMSLB GL RFC SPGL?PW @J?BBCP >. QOS?KMSQ ?. AS@MGB?J @. AMJSKL?P A. RP?LQGRGML?J 13. 8FGAF RWNC MD AMLLCARGTC RGQQSC F?Q RFC KMQR DG@CPQ >. JMMQC AMLLCARGTC RGQQSC ?. DG@PMSQ AMLLCARGTC RGQQSC @. A?PRGJ?EC A. @MLC 14. 8FGAF RWNC MD AMLLCARGTC RGQQSC F?Q ? KGLCP?JGXCB BGDDCPCLR K?RPGV >. JMMQC AMLLCARGTC RGQQSC ?. DG@PMSQ AMLLCARGTC RGQQSC @. A?PRGJ?EC A. @MLC 15. 5FC ACJJ DMSLB GL @MLC RF?R @PC?IQ GR BMUL GQ A?JJCB ?L >>>>>>>>. >. MQRCM@J?QR ?. MQRCMAWRC @. MQRCMAJ?QR A. MQRCML 16. 5FC ACJJ DMSLB GL @MLC RF?R K?ICQ RFC @MLC GQ A?JJCB ?L >>>>>>>>. >. MQRCM@J?QR ?. MQRCMAWRC @. MQRCMAJ?QR A. MQRCML 17. 1J?QK? GQ RFC >>>>>>>>. >. DG@CPQ GL @JMMB ?. K?RPGV MD @JMMB @. ACJJ RF?R NF?EMAWRGXCQ @?ARCPG? A. ACJJ DP?EKCLR DMSLB GL RFC RGQQSC 18. 5FC RWNC MD KSQAJC ACJJ SLBCP TMJSLR?PW AMLRPMJ GQ RFC >>>>>>>>. >. QKMMRF KSQAJC ?. QICJCR?J KSQAJC @. A?PBG?A KSQAJC A. TGQACP?J KSQAJC 19. 5FC N?PR MD ? LCSPML RF?R AMLR?GLQ RFC LSAJCSQ GQ RFC >. ACJJ @MBW ?. BCLBPGRC @. ?VML A. EJG?J 20. 8FCL D?ACB UGRF ? QSBBCL BPMN GL CLTGPMLKCLR?J RCKNCP?RSPC, ?L CLBMRFCPKGA ?LGK?J UGJJ:
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22. 8FGAF KCRFMB MD FC?R CVAF?LEC MAASPQ BSPGLE BGPCAR AMLR?AR @CRUCCL RFC QMSPAC ?LB ?LGK?J >. ?. @. A. P?BG?RGML CT?NMP?RGML AMLTCARGML AMLBSARGML
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962
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964
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965
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966
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GL RFC KMSRFQ MD K?LW ?LGK?JQ. 5FCPC ?PC RFPCC K?HMP EJ?LBQ RF?R QCAPCRC Q?JGT?qRFC N?PMRGB, RFC QS@K?LBG@SJ?P, ?LB RFC QS@JGLES?J. 4?JGT? AMLR?GLQ KSASQ RF?R KMGQRCLQ DMMB ?LB @SDDCPQ RFC N) MD RFC DMMB. 4?JGT? ?JQM AMLR?GLQ GKKSLMEJM@SJGLQ ?LB JWQMXWKCQ, UFGAF F?TC ?LRG@?ARCPG?J ?ARGML RM PCBSAC RMMRF BCA?W @W GLFG@GRGLE EPMURF MD QMKC @?ARCPG?. 4?JGT? ?JQM AMLR?GLQ ?L CLXWKC A?JJCB salivary amylase RF?R @CEGLQ RFC NPMACQQ MD AMLTCPRGLE QR?PAFCQ GL RFC DMMB GLRM ? BGQ?AAF?PGBC A?JJCB K?JRMQC. "LMRFCP CLXWKC A?JJCB lipase GQ NPMBSACB @W RFC ACJJQ GL RFC RMLESC. -GN?QCQ ?PC ? AJ?QQ MD CLXWKCQ RF?R A?L @PC?I BMUL RPGEJWACPGBCQ. 5FC JGLES?J JGN?QC @CEGLQ RFC @PC?IBMUL MD D?R AMKNMLCLRQ GL RFC DMMB. 5FC AFCUGLE ?LB UCRRGLE ?ARGML NPMTGBCB @W RFC RCCRF ?LB Q?JGT? NPCN?PC RFC DMMB GLRM ? K?QQ A?JJCB RFC bolus DMP QU?JJMUGLE. 5FC RMLESC FCJNQ GL QU?JJMUGLEqKMTGLE RFC @MJSQ DPMK RFC KMSRF GLRM RFC NF?PWLV. 5FC NF?PWLV MNCLQ RM RUM N?QQ?ECU?WQ: RFC RP?AFC?, UFGAF JC?BQ RM RFC JSLEQ, ?LB RFC CQMNF?ESQ, UFGAF JC?BQ RM RFC QRMK?AF. 5FC RP?AFC? F?Q ?L MNCLGLE A?JJCB RFC EJMRRGQ, UFGAF GQ AMTCPCB @W ? A?PRGJ?EGLMSQ DJ?N A?JJCB RFC CNGEJMRRGQ. 8FCL QU?JJMUGLE, RFC CNGEJMRRGQ AJMQCQ RFC EJMRRGQ ?LB DMMB N?QQCQ GLRM RFC CQMNF?ESQ ?LB LMR RFC RP?AFC?. 5FGQ ?PP?LECKCLR ?JJMUQ DMMB RM @C ICNR MSR MD RFC RP?AFC?.
!@>LI< 34.9 $FDBPQFLK LC CLLA ?BDFKP FK QEB (>) LO>I @>SFQV. &LLA FP J>PQF@>QBA ?V QBBQE >KA JLFPQBKBA ?V P>IFS> PB@OBQBA COLJ QEB (?) P>IFS>OV DI>KAP. %KWVJBP FK QEB P>IFS> ?BDFK QL AFDBPQ PQ>O@EBP >KA C>QP. 7FQE QEB EBIM LC QEB QLKDRB, QEB OBPRIQFKD ?LIRP FP JLSBA FKQL QEB BPLME>DRP ?V PT>IILTFKD. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V QEB .>QFLK>I #>K@BO )KPQFQRQB)
E845.'-:8 5FC esophagus GQ ? RS@SJ?P MPE?L RF?R AMLLCARQ RFC KMSRF RM RFC QRMK?AF. 5FC AFCUCB ?LB QMDRCLCB DMMB N?QQCQ RFPMSEF RFC CQMNF?ESQ ?DRCP @CGLE QU?JJMUCB. 5FC QKMMRF KSQAJCQ MD RFC CQMNF?ESQ SLBCPEM ? QCPGCQ MD U?TC JGIC KMTCKCLRQ A?JJCB peristalsis RF?R NSQF RFC DMMB RMU?PB RFC QRMK?AF, ?Q GJJSQRP?RCB GL Figure 34.10. 5FC NCPGQR?JQGQ U?TC GQ SLGBGPCARGML?JqGR KMTCQ DMMB DPMK RFC KMSRF RM RFC QRMK?AF, ?LB PCTCPQC KMTCKCLR GQ LMR NMQQG@JC. 5FC NCPGQR?JRGA KMTCKCLR MD RFC CQMNF?ESQ GQ ?L GLTMJSLR?PW PCDJCV; GR R?ICQ NJ?AC GL PCQNMLQC RM RFC ?AR MD QU?JJMUGLE.
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967
" PGLE-JGIC KSQAJC A?JJCB ? sphincter DMPKQ T?JTCQ GL RFC BGECQRGTC QWQRCK. 5FC E?QRPM-CQMNF?EC?J QNFGLARCP GQ JMA?RCB ?R RFC QRMK?AF CLB MD RFC CQMNF?ESQ. *L PCQNMLQC RM QU?JJMUGLE ?LB RFC NPCQQSPC CVCPRCB @W RFC @MJSQ MD DMMB, RFGQ QNFGLARCP MNCLQ, ?LB RFC @MJSQ CLRCPQ RFC QRMK?AF. 8FCL RFCPC GQ LM QU?JJMUGLE ?ARGML, RFGQ QNFGLARCP GQ QFSR ?LB NPCTCLRQ RFC AMLRCLRQ MD RFC QRMK?AF DPMK RP?TCJGLE SN RFC CQMNF?ESQ. .?LW ?LGK?JQ F?TC ? RPSC QNFGLARCP; FMUCTCP, GL FSK?LQ, RFCPC GQ LM RPSC QNFGLARCP, @SR RFC CQMNF?ESQ PCK?GLQ AJMQCB UFCL RFCPC GQ LM QU?JJMUGLE ?ARGML. "AGB PCDJSV MP tFC?PR@SPLu MAASPQ UFCL RFC ?AGBGA BGECQRGTC HSGACQ CQA?NC GLRM RFC CQMNF?ESQ. !942'). " J?PEC N?PR MD BGECQRGML MAASPQ GL RFC QRMK?AF, QFMUL GL Figure 34.11. 5FC stomach GQ ? Q?AJGIC MPE?L RF?R QCAPCRCQ E?QRPGA BGECQRGTC HSGACQ. 5FC N) GL RFC QRMK?AF GQ @CRUCCL 1.5 ?LB 2.5. 5FGQ FGEFJW ?AGBGA CLTGPMLKCLR GQ PCOSGPCB DMP RFC AFCKGA?J @PC?IBMUL MD DMMB ?LB RFC CVRP?ARGML MD LSRPGCLRQ. 8FCL CKNRW, RFC QRMK?AF GQ ? P?RFCP QK?JJ MPE?L; FMUCTCP, GR A?L CVN?LB RM SN RM 20 RGKCQ GRQ PCQRGLE QGXC UFCL DGJJCB UGRF DMMB. 5FGQ AF?P?ARCPGQRGA GQ N?PRGASJ?PJW SQCDSJ DMP ?LGK?JQ RF?R LCCB RM C?R UFCL DMMB GQ ?T?GJ?@JC.
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7EF@E LC QEB CLIILTFKD PQ>QBJBKQP >?LRQ QEB AFDBPQFSB PVPQBJ FP C>IPB #EVJB FP > JFUQROB LC CLLA >KA AFDBPQFSB GRF@BP QE>Q FP MOLAR@BA FK QEB PQLJ>@E. &LLA BKQBOP QEB I>ODB FKQBPQFKB ?BCLOB QEB PJ>II FKQBPQFKB. )K QEB PJ>II FKQBPQFKB, @EVJB JFUBP TFQE ?FIB, TEF@E BJRIPFCFBP C>QP. 4EB PQLJ>@E FP PBM>O>QBA COLJ QEB PJ>II FKQBPQFKB ?V QEB MVILOF@ PMEFK@QBO.
5FC QRMK?AF GQ ?JQM RFC K?HMP QGRC DMP NPMRCGL BGECQRGML GL ?LGK?JQ MRFCP RF?L PSKGL?LRQ. 1PMRCGL BGECQRGML GQ KCBG?RCB @W ?L CLXWKC A?JJCB NCNQGL GL RFC QRMK?AF AF?K@CP. Pepsin GQ QCAPCRCB @W RFC AFGCD ACJJQ GL RFC QRMK?AF GL ?L GL?ARGTC DMPK A?JJCB pepsinogen. 1CNQGL @PC?IQ NCNRGBC @MLBQ ?LB AJC?TCQ NPMRCGLQ GLRM QK?JJCP NMJWNCNRGBCQ; GR ?JQM FCJNQ ?ARGT?RC KMPC NCNQGLMECL, QR?PRGLE ? NMQGRGTC DCCB@?AI KCAF?LGQK RF?R ECLCP?RCQ KMPC NCNQGL. "LMRFCP ACJJ RWNCqN?PGCR?J ACJJQqQCAPCRC FWBPMECL ?LB AFJMPGBC GMLQ, UFGAF AMK@GLC GL RFC JSKCL RM DMPK FWBPMAFJMPGA ?AGB, RFC NPGK?PW ?AGBGA AMKNMLCLR MD RFC QRMK?AF HSGACQ. )WBPMAFJMPGA ?AGB FCJNQ RM AMLTCPR RFC GL?ARGTC NCNQGLMECL RM NCNQGL. 5FC FGEFJW ?AGBGA CLTGPMLKCLR ?JQM IGJJQ K?LW KGAPMMPE?LGQKQ GL RFC DMMB ?LB, AMK@GLCB UGRF RFC ?ARGML MD RFC CLXWKC NCNQGL, PCQSJRQ GL RFC FWBPMJWQGQ MD NPMRCGL GL RFC DMMB. $FCKGA?J BGECQRGML GQ D?AGJGR?RCB @W RFC AFSPLGLE ?ARGML MD RFC QRMK?AF. $MLRP?ARGML ?LB PCJ?V?RGML MD QKMMRF KSQAJCQ KGVCQ RFC QRMK?AF AMLRCLRQ ?@MSR CTCPW 20 KGLSRCQ. 5FC N?PRG?JJW BGECQRCB DMMB ?LB E?QRPGA HSGAC KGVRSPC GQ A?JJCB chyme. $FWKC N?QQCQ DPMK RFC QRMK?AF RM RFC QK?JJ GLRCQRGLC. 'SPRFCP NPMRCGL BGECQRGML R?ICQ NJ?AC GL RFC QK?JJ GLRCQRGLC. (?QRPGA CKNRWGLE MAASPQ UGRFGL RUM RM QGV FMSPQ ?DRCP ? KC?J. 0LJW ? QK?JJ ?KMSLR MD AFWKC GQ PCJC?QCB GLRM RFC QK?JJ GLRCQRGLC ?R ? RGKC. 5FC KMTCKCLR MD AFWKC DPMK RFC QRMK?AF GLRM RFC QK?JJ GLRCQRGLC GQ PCESJ?RCB @W RFC NWJMPGA QNFGLARCP.
968
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8FCL BGECQRGLE NPMRCGL ?LB QMKC D?RQ, RFC QRMK?AF JGLGLE KSQR @C NPMRCARCB DPMK ECRRGLE BGECQRCB @W NCNQGL. 5FCPC ?PC RUM NMGLRQ RM AMLQGBCP UFCL BCQAPG@GLE FMU RFC QRMK?AF JGLGLE GQ NPMRCARCB. 'GPQR, ?Q NPCTGMSQJW KCLRGMLCB, RFC CLXWKC NCNQGL GQ QWLRFCQGXCB GL RFC GL?ARGTC DMPK. 5FGQ NPMRCARQ RFC AFGCD ACJJQ, @CA?SQC NCNQGLMECL BMCQ LMR F?TC RFC Q?KC CLXWKC DSLARGML?JGRW MD NCNQGL. 4CAMLB, RFC QRMK?AF F?Q ? RFGAI KSASQ JGLGLE RF?R NPMRCARQ RFC SLBCPJWGLE RGQQSC DPMK RFC ?ARGML MD RFC BGECQRGTC HSGACQ. 8FCL RFGQ KSASQ JGLGLE GQ PSNRSPCB, SJACPQ A?L DMPK GL RFC QRMK?AF. 6JACPQ ?PC MNCL UMSLBQ GL MP ML ?L MPE?L A?SQCB @W @?ARCPG? (H,30*6)(*;,9 7@3690) UFCL RFC KSASQ JGLGLE GQ PSNRSPCB ?LB D?GJQ RM PCDMPK. !2'11 I39+89/3+ $FWKC KMTCQ DPMK RFC QRMK?AF RM RFC QK?JJ GLRCQRGLC. 5FC small intestine GQ RFC MPE?L UFCPC RFC BGECQRGML MD NPMRCGL, D?RQ, ?LB A?P@MFWBP?RCQ GQ AMKNJCRCB. 5FC QK?JJ GLRCQRGLC GQ ? JMLE RS@C-JGIC MPE?L UGRF ? FGEFJW DMJBCB QSPD?AC AMLR?GLGLE DGLECP-JGIC NPMHCARGMLQ A?JJCB RFC villi. 5FC ?NGA?J QSPD?AC MD C?AF TGJJSQ F?Q K?LW KGAPMQAMNGA NPMHCARGMLQ A?JJCB KGAPMTGJJG. 5FCQC QRPSARSPCQ, GJJSQRP?RCB GL Figure 34.12, ?PC JGLCB UGRF CNGRFCJG?J ACJJQ ML RFC JSKGL?J QGBC ?LB ?JJMU DMP RFC LSRPGCLRQ RM @C ?@QMP@CB DPMK RFC BGECQRCB DMMB ?LB ?@QMP@CB GLRM RFC @JMMB QRPC?K ML RFC MRFCP QGBC. 5FC TGJJG ?LB KGAPMTGJJG, UGRF RFCGP K?LW DMJBQ, GLAPC?QC RFC QSPD?AC ?PC? MD RFC GLRCQRGLC ?LB GLAPC?QC ?@QMPNRGML CDDGAGCLAW MD RFC LSRPGCLRQ. "@QMP@CB LSRPGCLRQ GL RFC @JMMB ?PC A?PPGCB GLRM RFC FCN?RGA NMPR?J TCGL, UFGAF JC?BQ RM RFC JGTCP. 5FCPC, RFC JGTCP PCESJ?RCQ RFC BGQRPG@SRGML MD LSRPGCLRQ RM RFC PCQR MD RFC @MBW ?LB PCKMTCQ RMVGA QS@QR?LACQ, GLAJSBGLE BPSEQ, ?JAMFMJ, ?LB QMKC N?RFMECLQ.
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t$-QF?NCB,u DGVCB N?PR MD RFC QK?JJ GLRCQRGLC GQ A?JJCB RFC duodenum ?LB GQ QFMUL GL Figure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ile GQ NPMBSACB GL RFC JGTCP ?LB QRMPCB ?LB AMLACLRP?RCB GL RFC E?JJ@J?BBCP. #GJC AMLR?GLQ @GJC Q?JRQ UFGAF CKSJQGDW JGNGBQ UFGJC RFC N?LAPC?Q NPMBSACQ CLXWKCQ RF?R A?R?@MJGXC QR?PAFCQ, BGQ?AAF?PGBCQ, NPMRCGLQ, ?LB D?RQ. 5FCQC BGECQRGTC HSGACQ @PC?I BMUL RFC DMMB N?PRGAJCQ GL RFC AFWKC GLRM EJSAMQC, RPGEJWACPGBCQ, ?LB ?KGLM ?AGBQ. 4MKC AFCKGA?J BGECQRGML MD DMMB R?ICQ NJ?AC GL RFC BSMBCLSK. "@QMPNRGML MD D?RRW ?AGBQ ?JQM R?ICQ NJ?AC GL RFC BSMBCLSK.
#(!04%2 34 X !.)-!, .542)4)/. !.$ 4(% $)'%34)6% 3934%-
969
5FC QCAMLB N?PR MD RFC QK?JJ GLRCQRGLC GQ A?JJCB RFC jejunum, QFMUL GL Figure 34.11. )CPC, FWBPMJWQGQ MD LSRPGCLRQ GQ AMLRGLSCB UFGJC KMQR MD RFC A?P@MFWBP?RCQ ?LB ?KGLM ?AGBQ ?PC ?@QMP@CB RFPMSEF RFC GLRCQRGL?J JGLGLE. 5FC @SJI MD AFCKGA?J BGECQRGML ?LB LSRPGCLR ?@QMPNRGML MAASPQ GL RFC HCHSLSK. 5FC ileum, ?JQM GJJSQRP?RCB GL Figure 34.11 GQ RFC J?QR N?PR MD RFC QK?JJ GLRCQRGLC ?LB FCPC RFC @GJC Q?JRQ ?LB TGR?KGLQ ?PC ?@QMP@CB GLRM @JMMB QRPC?K. 5FC SLBGECQRCB DMMB GQ QCLR RM RFC AMJML DPMK RFC GJCSK TG? NCPGQR?JRGA KMTCKCLRQ MD RFC KSQAJC. 5FC GJCSK CLBQ ?LB RFC J?PEC GLRCQRGLC @CEGLQ ?R RFC GJCMACA?J T?JTC. 5FC TCPKGDMPK, tUMPK-JGIC,u ?NNCLBGV GQ JMA?RCB ?R RFC GJCMACA?J T?JTC. 5FC ?NNCLBGV MD FSK?LQ QCAPCRCQ LM CLXWKCQ ?LB F?Q ?L GLQGELGDGA?LR PMJC GL GKKSLGRW. L'7-+ I39+89/3+ 5FC large intestine, GJJSQRP?RCB GL Figure 34.13, PC?@QMP@Q RFC U?RCP DPMK RFC SLBGECQRCB DMMB K?RCPG?J ?LB NPMACQQCQ RFC U?QRC K?RCPG?J. 5FC FSK?L J?PEC GLRCQRGLC GQ KSAF QK?JJCP GL JCLERF AMKN?PCB RM RFC QK?JJ GLRCQRGLC @SR J?PECP GL BG?KCRCP. *R F?Q RFPCC N?PRQ: RFC ACASK, RFC AMJML, ?LB RFC PCARSK. 5FC ACASK HMGLQ RFC GJCSK RM RFC AMJML ?LB GQ RFC PCACGTGLE NMSAF DMP RFC U?QRC K?RRCP. 5FC AMJML GQ FMKC RM K?LW @?ARCPG? MP tGLRCQRGL?J DJMP?u RF?R ?GB GL RFC BGECQRGTC NPMACQQCQ. 5FC AMJML A?L @C BGTGBCB GLRM DMSP PCEGMLQ, RFC ?QACLBGLE AMJML, RFC RP?LQTCPQC AMJML, RFC BCQACLBGLE AMJML ?LB RFC QGEKMGB AMJML. 5FC K?GL DSLARGMLQ MD RFC AMJML ?PC RM CVRP?AR RFC U?RCP ?LB KGLCP?J Q?JRQ DPMK SLBGECQRCB DMMB, ?LB RM QRMPC U?QRC K?RCPG?J. $?PLGTMPMSQ K?KK?JQ F?TC ? QFMPRCP J?PEC GLRCQRGLC AMKN?PCB RM FCP@GTMPMSQ K?KK?JQ BSC RM RFCGP BGCR.
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+)9:2 '3* A3:8 5FC rectum GQ RFC RCPKGL?J CLB MD RFC J?PEC GLRCQRGLC, ?Q QFMUL GL Figure 34.13. 5FC NPGK?PW PMJC MD RFC PCARSK GQ RM QRMPC RFC DCACQ SLRGJ BCDCA?RGML. 5FC DCACQ ?PC NPMNCJJCB SQGLE NCPGQR?JRGA KMTCKCLRQ BSPGLE CJGKGL?RGML. 5FC anus GQ ?L MNCLGLE ?R RFC D?P-CLB MD RFC BGECQRGTC RP?AR ?LB GQ RFC CVGR NMGLR DMP RFC U?QRC K?RCPG?J. 5UM QNFGLARCPQ @CRUCCL RFC PCARSK ?LB ?LSQ AMLRPMJ CJGKGL?RGML: RFC GLLCP QNFGLARCP GQ GLTMJSLR?PW ?LB RFC MSRCP QNFGLARCP GQ TMJSLR?PW. A))+8847> O7-'38 5FC MPE?LQ BGQASQQCB ?@MTC ?PC RFC MPE?LQ MD RFC BGECQRGTC RP?AR RFPMSEF UFGAF DMMB N?QQCQ. "AACQQMPW MPE?LQ ?PC MPE?LQ RF?R ?BB QCAPCRGMLQ (CLXWKCQ) RF?R A?R?@MJGXC DMMB GLRM LSRPGCLRQ. "AACQQMPW MPE?LQ GLAJSBC Q?JGT?PW EJ?LBQ, RFC JGTCP, RFC N?LAPC?Q, ?LB RFC E?JJ@J?BBCP. 5FC JGTCP, N?LAPC?Q, ?LB E?JJ@J?BBCP ?PC PCESJ?RCB @W FMPKMLCQ GL PCQNMLQC RM RFC DMMB AMLQSKCB. 5FC liver GQ RFC J?PECQR GLRCPL?J MPE?L GL FSK?LQ ?LB GR NJ?WQ ? TCPW GKNMPR?LR PMJC GL BGECQRGML MD D?RQ ?LB BCRMVGDWGLE @JMMB. 5FC JGTCP NPMBSACQ @GJC, ? BGECQRGTC HSGAC RF?R GQ PCOSGPCB DMP RFC @PC?IBMUL MD D?RRW AMKNMLCLRQ MD RFC DMMB GL RFC BSMBCLSK. 5FC JGTCP ?JQM NPMACQQCQ RFC TGR?KGLQ ?LB D?RQ ?LB QWLRFCQGXCQ K?LW NJ?QK? NPMRCGLQ. 5FC pancreas GQ ?LMRFCP GKNMPR?LR EJ?LB RF?R QCAPCRCQ BGECQRGTC HSGACQ. 5FC AFWKC NPMBSACB DPMK RFC QRMK?AF GQ FGEFJW ?AGBGA GL L?RSPC; RFC N?LAPC?RGA HSGACQ AMLR?GL FGEF JCTCJQ MD @GA?P@ML?RC, ?L ?JI?JG RF?R LCSRP?JGXCQ RFC ?AGBGA AFWKC. "BBGRGML?JJW, RFC N?LAPC?RGA HSGACQ AMLR?GL ? J?PEC T?PGCRW MD CLXWKCQ RF?R ?PC PCOSGPCB DMP RFC BGECQRGML MD NPMRCGL ?LB A?P@MFWBP?RCQ. 5FC gallbladder GQ ? QK?JJ MPE?L RF?R ?GBQ RFC JGTCP @W QRMPGLE @GJC ?LB AMLACLRP?RGLE @GJC Q?JRQ. 8FCL AFWKC AMLR?GLGLE D?RRW ?AGBQ CLRCPQ RFC BSMBCLSK, RFC @GJC GQ QCAPCRCB DPMK RFC E?JJ@J?BBCP GLRM RFC BSMBCLSK.
970
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34.2 R )LKI@K@FE 8E; EP +IF;L:K@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL UFW ?L ?LGK?JsQ BGCR QFMSJB @C @?J?LACB ?LB KCCR RFC LCCBQ MD RFC @MBW z %CDGLC RFC NPGK?PW AMKNMLCLRQ MD DMMB z %CQAPG@C RFC CQQCLRG?J LSRPGCLRQ PCOSGPCB DMP ACJJSJ?P DSLARGML RF?R A?LLMR @C QWLRFCQGXCB @W RFC ?LGK?J @MBW z &VNJ?GL FMU CLCPEW GQ NPMBSACB RFPMSEF BGCR ?LB BGECQRGML z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8F?R ?PC RFC DSLB?KCLR?J PCOSGPCKCLRQ MD RFC ?LGK?J BGCR 5FC ?LGK?J BGCR QFMSJB @C UCJJ @?J?LACB ?LB NPMTGBC LSRPGCLRQ PCOSGPCB DMP @MBGJW DSLARGML ?LB RFC KGLCP?JQ ?LB TGR?KGLQ PCOSGPCB DMP K?GLR?GLGLE QRPSARSPC ?LB PCESJ?RGML LCACQQ?PW DMP EMMB FC?JRF ?LB PCNPMBSARGTC A?N?@GJGRW. 5FCQC PCOSGPCKCLRQ DMP ? FSK?L ?PC GJJSQRP?RCB EP?NFGA?JJW GL Figure 34.14
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D/-+89/;+ P.'8+8 5FC PCQNMLQC RM DMMB @CEGLQ CTCL @CDMPC DMMB CLRCPQ RFC KMSRF. 5FC DGPQR NF?QC MD GLECQRGML, A?JJCB RFC cephalic phase, GQ AMLRPMJJCB @W RFC LCSP?J PCQNMLQC RM RFC QRGKSJSQ NPMTGBCB @W DMMB. "JJ ?QNCARQqQSAF ?Q QGEFR, QCLQC, ?LB QKCJJqRPGEECP RFC LCSP?J PCQNMLQCQ PCQSJRGLE GL Q?JGT?RGML ?LB QCAPCRGML MD E?QRPGA HSGACQ. 5FC E?QRPGA ?LB Q?JGT?PW QCAPCRGML GL RFC ACNF?JGA NF?QC A?L ?JQM R?IC NJ?AC BSC RM RFC RFMSEFR MD DMMB. 3GEFR LMU, GD WMS RFGLI ?@MSR ? NGCAC MD AFMAMJ?RC MP ? APGQNW NMR?RM AFGN, RFC GLAPC?QC GL Q?JGT?RGML GQ ? ACNF?JGA NF?QC PCQNMLQC RM RFC RFMSEFR. 5FC ACLRP?J LCPTMSQ QWQRCK NPCN?PCQ RFC QRMK?AF RM PCACGTC DMMB. 5FC gastric phase @CEGLQ MLAC RFC DMMB ?PPGTCQ GL RFC QRMK?AF. *R @SGJBQ ML RFC QRGKSJ?RGML NPMTGBCB BSPGLE RFC ACNF?JGA NF?QC. (?QRPGA ?AGBQ ?LB CLXWKCQ NPMACQQ RFC GLECQRCB K?RCPG?JQ. 5FC E?QRPGA NF?QC GQ QRGKSJ?RCB @W (1) BGQRCLQGML MD RFC QRMK?AF, (2) ? BCAPC?QC GL RFC N) MD RFC E?QRPGA AMLRCLRQ, ?LB (3) RFC NPCQCLAC MD SLBGECQRCB K?RCPG?J. 5FGQ NF?QC AMLQGQRQ MD JMA?J, FMPKML?J, ?LB LCSP?J PCQNMLQCQ. 5FCQC PCQNMLQCQ QRGKSJ?RC QCAPCRGMLQ ?LB NMUCPDSJ AMLRP?ARGMLQ.
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5FC intestinal phase @CEGLQ UFCL AFWKC CLRCPQ RFC QK?JJ GLRCQRGLC RPGEECPGLE BGECQRGTC QCAPCRGMLQ. 5FGQ NF?QC AMLRPMJQ RFC P?RC MD E?QRPGA CKNRWGLE. *L ?BBGRGML RM E?QRPGL CKNRWGLE, UFCL AFWKC CLRCPQ RFC QK?JJ GLRCQRGLC, GR RPGEECPQ MRFCP FMPKML?J ?LB LCSP?J CTCLRQ RF?R AMMPBGL?RC RFC ?ARGTGRGCQ MD RFC GLRCQRGL?J RP?AR, N?LAPC?Q, JGTCP, ?LB E?JJ@J?BBCP.
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5FC endocrine system AMLRPMJQ RFC PCQNMLQC MD RFC T?PGMSQ EJ?LBQ GL RFC @MBW ?LB RFC PCJC?QC MD FMPKMLCQ ?R RFC ?NNPMNPG?RC RGKCQ. 0LC MD RFC GKNMPR?LR D?ARMPQ SLBCP FMPKML?J AMLRPMJ GQ RFC QRMK?AF ?AGB CLTGPMLKCLR. %SPGLE RFC E?QRPGA NF?QC, RFC FMPKMLC gastrin GQ QCAPCRCB @W ( ACJJQ GL RFC QRMK?AF GL PCQNMLQC RM RFC NPCQCLAC MD NPMRCGLQ. (?QRPGL QRGKSJ?RCQ RFC PCJC?QC MD QRMK?AF ?AGB, MP FWBPMAFJMPGA ?AGB ()$J) UFGAF ?GBQ GL RFC BGECQRGML MD RFC NPMRCGLQ. )MUCTCP, UFCL RFC QRMK?AF GQ CKNRGCB, RFC ?AGBGA CLTGPMLKCLR LCCB LMR @C K?GLR?GLCB ?LB ? FMPKMLC A?JJCB somatostatin QRMNQ RFC PCJC?QC MD FWBPMAFJMPGA ?AGB. 5FGQ GQ AMLRPMJJCB @W ? LCE?RGTC DCCB@?AI KCAF?LGQK. *L RFC BSMBCLSK, BGECQRGTC QCAPCRGMLQ DPMK RFC JGTCP, N?LAPC?Q, ?LB E?JJ@J?BBCP NJ?W ?L GKNMPR?LR PMJC GL BGECQRGLE AFWKC BSPGLE RFC GLRCQRGL?J NF?QC. *L MPBCP RM LCSRP?JGXC RFC ?AGBGA AFWKC, ? FMPKMLC A?JJCB secretin QRGKSJ?RCQ RFC N?LAPC?Q RM NPMBSAC ?JI?JGLC @GA?P@ML?RC QMJSRGML ?LB BCJGTCP GR RM RFC BSMBCLSK. 4CAPCRGL ?ARQ GL R?LBCK UGRF ?LMRFCP FMPKMLC A?JJCB cholecystokinin ($$,). /MR MLJW BMCQ $$, QRGKSJ?RC RFC N?LAPC?Q RM NPMBSAC RFC PCOSGQGRC N?LAPC?RGA HSGACQ, GR ?JQM QRGKSJ?RCQ RFC E?JJ@J?BBCP RM PCJC?QC @GJC GLRM RFC BSMBCLSK.
7GQGR this website (http://openstaxcollege.org/l/enteric_endo) RM JC?PL KMPC ?@MSR RFC CLBMAPGLC QWQRCK. 3CTGCU RFC RCVR ?LB U?RAF RFC ?LGK?RGML MD FMU AMLRPMJ GQ GKNJCKCLRCB GL RFC CLBMAPGLC QWQRCK. "LMRFCP JCTCJ MD FMPKML?J AMLRPMJ MAASPQ GL PCQNMLQC RM RFC AMKNMQGRGML MD DMMB. 'MMBQ FGEF GL JGNGBQ R?IC ? JMLE RGKC RM BGECQR. " FMPKMLC A?JJCB gastric inhibitory peptide GQ QCAPCRCB @W RFC QK?JJ GLRCQRGLC RM QJMU BMUL RFC NCPGQR?JRGA KMTCKCLRQ MD RFC GLRCQRGLC RM ?JJMU D?RRW DMMBQ KMPC RGKC RM @C BGECQRCB ?LB ?@QMP@CB. 6LBCPQR?LBGLE RFC FMPKML?J AMLRPMJ MD RFC BGECQRGTC QWQRCK GQ ?L GKNMPR?LR ?PC? MD MLEMGLE PCQC?PAF. 4AGCLRGQRQ ?PC CVNJMPGLE RFC PMJC MD C?AF FMPKMLC GL RFC BGECQRGTC NPMACQQ ?LB BCTCJMNGLE U?WQ RM R?PECR RFCQC FMPKMLCQ. "BT?LACQ AMSJB JC?B RM ILMUJCBEC RF?R K?W FCJN RM @?RRJC RFC M@CQGRW CNGBCKGA.
984
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34.4 D@><JK@M< .PJK<D -<>LC8K@FE

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16. )MU BMCQ RFC NMJWE?QRPGA BGECQRGTC QWQRCK ?GB GL BGECQRGLE PMSEF?EC 17. )MU BM @GPBQ BGECQR RFCGP DMMB GL RFC ?@QCLAC MD RCCRF 18. 8F?R GQ RFC PMJC MD RFC ?AACQQMPW MPE?LQ GL BGECQRGML 19. &VNJ?GL FMU RFC TGJJG ?LB KGAPMTGJJG ?GB GL ?@QMPNRGML. 20. 8F?R ?PC CQQCLRG?J LSRPGCLRQ 21. 8F?R GQ RFC PMJC MD KGLCP?JQ GL K?GLR?GLGLE EMMB FC?JRF 22. %GQASQQ UFW M@CQGRW GQ ? EPMUGLE CNGBCKGA. 23. 5FCPC ?PC QCTCP?J L?RGMLQ UFCPC K?JLMSPGQFKCLR GQ ? AMKKML MAASPPCLAC. 8F?R K?W @C QMKC MD RFC FC?JRF AF?JJCLECQ NMQCB @W K?JLSRPGRGML 24. &VNJ?GL UFW QMKC BGCR?PW JGNGB GQ ? LCACQQ?PW N?PR MD ? @?J?LACB BGCR. 25. %CQAPG@C FMU FMPKMLCQ PCESJ?RC BGECQRGML. 26. %CQAPG@C MLC MP KMPC QACL?PGMQ UFCPC JMQQ MD FMPKML?J PCESJ?RGML MD BGECQRGML A?L JC?B RM BGQC?QCQ.
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8FCL WMSsPC PC?BGLE RFGQ @MMI, WMSP LCPTMSQ QWQRCK GQ NCPDMPKGLE QCTCP?J DSLARGMLQ QGKSJR?LCMSQJW. 5FC TGQS?J QWQRCK GQ NPMACQQGLE UF?R GQ QCCL ML RFC N?EC; RFC KMRMP QWQRCK AMLRPMJQ RFC RSPL MD RFC N?ECQ (MP AJGAI MD RFC KMSQC); RFC NPCDPMLR?J AMPRCV K?GLR?GLQ ?RRCLRGML. &TCL DSLB?KCLR?J DSLARGMLQ, JGIC @PC?RFGLE ?LB PCESJ?RGML MD @MBW RCKNCP?RSPC, ?PC AMLRPMJJCB @W RFC LCPTMSQ QWQRCK. " LCPTMSQ QWQRCK GQ ?L MPE?LGQKsQ AMLRPMJ ACLRCP: GR NPMACQQCQ QCLQMPW GLDMPK?RGML DPMK MSRQGBC (?LB GLQGBC) RFC @MBW ?LB AMLRPMJQ ?JJ @CF?TGMPQqDPMK C?RGLE RM QJCCNGLE RM DGLBGLE ? K?RC.
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z -GQR ?LB BCQAPG@C RFC DSLARGMLQ MD RFC QRPSARSP?J AMKNMLCLRQ MD ? LCSPML z -GQR ?LB BCQAPG@C RFC DMSP K?GL RWNCQ MD LCSPMLQ z $MKN?PC RFC DSLARGMLQ MD BGDDCPCLR RWNCQ MD EJG?J ACJJQ
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/CPTMSQ QWQRCKQ RFPMSEFMSR RFC ?LGK?J IGLEBMK T?PW GL QRPSARSPC ?LB AMKNJCVGRW, ?Q GJJSQRP?RCB @W RFC T?PGCRW MD ?LGK?JQ QFMUL GL Figure 35.2. 4MKC MPE?LGQKQ, JGIC QC? QNMLECQ, J?AI ? RPSC LCPTMSQ QWQRCK. 0RFCPQ, JGIC HCJJWDGQF, J?AI ? RPSC @P?GL ?LB GLQRC?B F?TC ? QWQRCK MD QCN?P?RC @SR AMLLCARCB LCPTC ACJJQ (LCSPMLQ) A?JJCB ? tLCPTC LCR.u &AFGLMBCPKQ QSAF ?Q QC? QR?PQ F?TC LCPTC ACJJQ RF?R ?PC @SLBJCB GLRM DG@CPQ A?JJCB LCPTCQ. 'J?RUMPKQ MD RFC NFWJSK 1J?RWFCJKGLRFCQ F?TC @MRF ? ACLRP?J LCPTMSQ QWQRCK ($/4), K?BC SN MD ? QK?JJ t@P?GLu ?LB RUM LCPTC AMPBQ, ?LB ? NCPGNFCP?J LCPTMSQ QWQRCK (1/4) AMLR?GLGLE ? QWQRCK MD LCPTCQ RF?R CVRCLB RFPMSEFMSR RFC @MBW. 5FC GLQCAR LCPTMSQ QWQRCK GQ KMPC AMKNJCV @SR ?JQM D?GPJW BCACLRP?JGXCB. *R AMLR?GLQ ? @P?GL, TCLRP?J LCPTC AMPB, ?LB E?LEJG? (AJSQRCPQ MD AMLLCARCB LCSPMLQ). 5FCQC E?LEJG? A?L AMLRPMJ KMTCKCLRQ ?LB @CF?TGMPQ UGRFMSR GLNSR DPMK RFC @P?GL. 0ARMNG K?W F?TC RFC KMQR AMKNJGA?RCB MD GLTCPRC@P?RC LCPTMSQ QWQRCKQqRFCW F?TC LCSPMLQ RF?R ?PC MPE?LGXCB GL QNCAG?JGXCB JM@CQ ?LB CWCQ RF?R ?PC QRPSARSP?JJW QGKGJ?P RM TCPRC@P?RC QNCAGCQ.
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5FC LCPTMSQ QWQRCK MD RFC AMKKML J?@MP?RMPW DJW, D96:67/03( 4,3(56.(:;,9, AMLR?GLQ ?PMSLB 100,000 LCSPMLQ, RFC Q?KC LSK@CP ?Q ? JM@QRCP. 5FGQ LSK@CP AMKN?PCQ RM 75 KGJJGML GL RFC KMSQC ?LB 300 KGJJGML GL RFC MARMNSQ. " FSK?L @P?GL AMLR?GLQ ?PMSLB 86 @GJJGML LCSPMLQ. %CQNGRC RFCQC TCPW BGDDCPCLR LSK@CPQ, RFC LCPTMSQ QWQRCKQ MD RFCQC ?LGK?JQ AMLRPMJ K?LW MD RFC Q?KC @CF?TGMPQqDPMK @?QGA PCDJCVCQ RM KMPC AMKNJGA?RCB @CF?TGMPQ JGIC DGLBGLE DMMB ?LB AMSPRGLE K?RCQ. 5FC ?@GJGRW MD LCSPMLQ RM AMKKSLGA?RC UGRF C?AF MRFCP ?Q UCJJ ?Q UGRF MRFCP RWNCQ MD ACJJQ SLBCPJGCQ ?JJ MD RFCQC @CF?TGMPQ. .MQR LCSPMLQ QF?PC RFC Q?KC ACJJSJ?P AMKNMLCLRQ. #SR LCSPMLQ ?PC ?JQM FGEFJW QNCAG?JGXCBqBGDDCPCLR RWNCQ MD LCSPMLQ F?TC BGDDCPCLR QGXCQ ?LB QF?NCQ RF?R PCJ?RC RM RFCGP DSLARGML?J PMJCQ. P'798 4, ' N+:743 -GIC MRFCP ACJJQ, C?AF LCSPML F?Q ? ACJJ @MBW (MP QMK?) RF?R AMLR?GLQ ? LSAJCSQ, QKMMRF ?LB PMSEF CLBMNJ?QKGA PCRGASJSK, (MJEG ?NN?P?RSQ, KGRMAFMLBPG?, ?LB MRFCP ACJJSJ?P AMKNMLCLRQ. /CSPMLQ ?JQM AMLR?GL SLGOSC QRPSARSPCQ, GJJSQRP?RCB GL Figure 35.3 DMP PCACGTGLE ?LB QCLBGLE RFC CJCARPGA?J QGEL?JQ RF?R K?IC LCSPML?J AMKKSLGA?RGML NMQQG@JC. Dendrites ?PC RPCC-JGIC QRPSARSPCQ RF?R CVRCLB ?U?W DPMK RFC ACJJ @MBW RM PCACGTC KCQQ?ECQ DPMK MRFCP LCSPMLQ ?R QNCAG?JGXCB HSLARGMLQ A?JJCB synapses. "JRFMSEF QMKC LCSPMLQ BM LMR F?TC ?LW BCLBPGRCQ, QMKC RWNCQ MD LCSPMLQ F?TC KSJRGNJC BCLBPGRCQ. %CLBPGRCQ A?L F?TC QK?JJ NPMRPSQGMLQ A?JJCB BCLBPGRGA QNGLCQ, UFGAF DSPRFCP GLAPC?QC QSPD?AC ?PC? DMP NMQQG@JC QWL?NRGA AMLLCARGMLQ. 0LAC ? QGEL?J GQ PCACGTCB @W RFC BCLBPGRC, GR RFCL RP?TCJQ N?QQGTCJW RM RFC ACJJ @MBW. 5FC ACJJ @MBW AMLR?GLQ ? QNCAG?JGXCB QRPSARSPC, RFC axon hillock RF?R GLRCEP?RCQ QGEL?JQ DPMK KSJRGNJC QWL?NQCQ ?LB QCPTCQ ?Q ? HSLARGML @CRUCCL RFC ACJJ @MBW ?LB ?L axon. "L ?VML GQ ? RS@C-JGIC QRPSARSPC RF?R NPMN?E?RCQ RFC GLRCEP?RCB QGEL?J RM QNCAG?JGXCB CLBGLEQ A?JJCB axon terminals. 5FCQC RCPKGL?JQ GL RSPL QWL?NQC ML MRFCP LCSPMLQ, KSQAJC, MP R?PECR MPE?LQ. $FCKGA?JQ PCJC?QCB ?R ?VML RCPKGL?JQ ?JJMU QGEL?JQ RM @C AMKKSLGA?RCB RM RFCQC MRFCP ACJJQ. /CSPMLQ SQS?JJW F?TC MLC MP RUM ?VMLQ, @SR QMKC LCSPMLQ, JGIC ?K?APGLC ACJJQ GL RFC PCRGL?, BM LMR AMLR?GL ?LW ?VMLQ. 4MKC ?VMLQ ?PC AMTCPCB UGRF myelin, UFGAF ?ARQ ?Q ?L GLQSJ?RMP RM KGLGKGXC BGQQGN?RGML MD RFC CJCARPGA?J QGEL?J ?Q GR RP?TCJQ BMUL RFC ?VML, EPC?RJW GLAPC?QGLE RFC QNCCB ML AMLBSARGML. 5FGQ GLQSJ?RGML GQ GKNMPR?LR ?Q RFC ?VML DPMK ? FSK?L KMRMP LCSPML A?L @C ?Q JMLE ?Q ? KCRCPqDPMK RFC @?QC MD RFC QNGLC RM RFC RMCQ. 5FC KWCJGL QFC?RF GQ LMR ?ARS?JJW N?PR MD RFC LCSPML. .WCJGL GQ NPMBSACB @W EJG?J ACJJQ. "JMLE RFC ?VML RFCPC ?PC NCPGMBGA E?NQ GL RFC KWCJGL QFC?RF. 5FCQC E?NQ ?PC A?JJCB nodes of Ranvier ?LB ?PC QGRCQ UFCPC RFC QGEL?J GQ tPCAF?PECBu ?Q GR RP?TCJQ ?JMLE RFC ?VML. *R GQ GKNMPR?LR RM LMRC RF?R ? QGLEJC LCSPML BMCQ LMR ?AR ?JMLCqLCSPML?J AMKKSLGA?RGML BCNCLBQ ML RFC AMLLCARGMLQ RF?R LCSPMLQ K?IC UGRF MLC ?LMRFCP (?Q UCJJ ?Q UGRF MRFCP ACJJQ, JGIC KSQAJC ACJJQ). %CLBPGRCQ DPMK ? QGLEJC LCSPML K?W PCACGTC QWL?NRGA AMLR?AR DPMK K?LW MRFCP LCSPMLQ. 'MP CV?KNJC, BCLBPGRCQ DPMK ? 1SPIGLHC ACJJ GL RFC ACPC@CJJSK ?PC RFMSEFR RM PCACGTC AMLR?AR DPMK ?Q K?LW ?Q 200,000 MRFCP LCSPMLQ.
992
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qKMQR @P?GL RSKMPQ ?PC A?SQCB @W KSR?RGMLQ GL EJG?. ">5+8 4, G1/' 5FCPC ?PC QCTCP?J BGDDCPCLR RWNCQ MD EJG? UGRF BGDDCPCLR DSLARGMLQ, RUM MD UFGAF ?PC QFMUL GL Figure 35.7. Astrocytes, QFMUL GL Figure 35.8a K?IC AMLR?AR UGRF @MRF A?NGJJ?PGCQ ?LB LCSPMLQ GL RFC $/4. 5FCW NPMTGBC LSRPGCLRQ ?LB MRFCP QS@QR?LACQ RM LCSPMLQ, PCESJ?RC RFC AMLACLRP?RGMLQ MD GMLQ ?LB AFCKGA?JQ GL RFC CVRP?ACJJSJ?P DJSGB, ?LB NPMTGBC QRPSARSP?J QSNNMPR DMP QWL?NQCQ. "QRPMAWRCQ ?JQM DMPK RFC @JMMB-@P?GL @?PPGCPq? QRPSARSPC RF?R @JMAIQ CLRP?LAC MD RMVGA QS@QR?LACQ GLRM RFC @P?GL. "QRPMAWRCQ, GL N?PRGASJ?P, F?TC @CCL QFMUL RFPMSEF A?JAGSK GK?EGLE CVNCPGKCLRQ RM @CAMKC ?ARGTC GL PCQNMLQC RM LCPTC ?ARGTGRW, RP?LQKGR A?JAGSK U?TCQ @CRUCCL ?QRPMAWRCQ, ?LB KMBSJ?RC RFC ?ARGTGRW MD QSPPMSLBGLE QWL?NQCQ. Satellite glia NPMTGBC LSRPGCLRQ ?LB QRPSARSP?J QSNNMPR DMP LCSPMLQ GL RFC 1/4. Microglia QA?TCLEC ?LB BCEP?BC BC?B ACJJQ ?LB NPMRCAR RFC @P?GL DPMK GLT?BGLE KGAPMMPE?LGQKQ. Oligodendrocytes, QFMUL GL Figure 35.8b DMPK KWCJGL QFC?RFQ ?PMSLB ?VMLQ GL RFC $/4. 0LC ?VML A?L @C KWCJGL?RCB @W QCTCP?J MJGEMBCLBPMAWRCQ, ?LB MLC MJGEMBCLBPMAWRC A?L NPMTGBC KWCJGL DMP KSJRGNJC LCSPMLQ. 5FGQ GQ BGQRGLARGTC DPMK RFC 1/4 UFCPC ? QGLEJC Schwann cell NPMTGBCQ KWCJGL DMP MLJW MLC ?VML ?Q RFC CLRGPC 4AFU?LL ACJJ QSPPMSLBQ RFC ?VML. Radial glia QCPTC ?Q QA?DDMJBQ DMP BCTCJMNGLE LCSPMLQ ?Q RFCW KGEP?RC RM RFCGP CLB BCQRGL?RGMLQ. Ependymal ACJJQ JGLC DJSGB-DGJJCB TCLRPGAJCQ MD RFC @P?GL ?LB RFC ACLRP?J A?L?J MD RFC QNGL?J AMPB. 5FCW ?PC GLTMJTCB GL RFC NPMBSARGML MD ACPC@PMQNGL?J DJSGB, UFGAF QCPTCQ ?Q ? ASQFGML DMP RFC @P?GL, KMTCQ RFC DJSGB @CRUCCL RFC QNGL?J AMPB ?LB RFC @P?GL, ?LB GQ ? AMKNMLCLR DMP RFC AFMPMGB NJCVSQ.
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996
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BCAPC?QC GL RFC BGDDCPCLAC GL TMJR?EC @CRUCCL RFC GLQGBC ?LB MSRQGBC MD RFC LCSPML. " QRGKSJSQ DPMK ? QCLQMPW ACJJ MP ?LMRFCP LCSPML BCNMJ?PGXCQ RFC R?PECR LCSPML RM GRQ RFPCQFMJB NMRCLRG?J (-55 K7). /?+ AF?LLCJQ GL RFC ?VML FGJJMAI MNCL, ?JJMUGLE NMQGRGTC GMLQ RM CLRCP RFC ACJJ (Figure 35.10 ?LB Figure 35.11). 0LAC RFC QMBGSK AF?LLCJQ MNCL, RFC LCSPML AMKNJCRCJW BCNMJ?PGXCQ RM ? KCK@P?LC NMRCLRG?J MD ?@MSR +40 K7. "ARGML NMRCLRG?JQ ?PC AMLQGBCPCB ?L "?JJ-MP LMRFGLE" CTCLR, GL RF?R, MLAC RFC RFPCQFMJB NMRCLRG?J GQ PC?AFCB, RFC LCSPML ?JU?WQ AMKNJCRCJW BCNMJ?PGXCQ. 0LAC BCNMJ?PGX?RGML GQ AMKNJCRC, RFC ACJJ KSQR LMU "PCQCR" GRQ KCK@P?LC TMJR?EC @?AI RM RFC PCQRGLE NMRCLRG?J. 5M ?AAMKNJGQF RFGQ, RFC /?+ AF?LLCJQ AJMQC ?LB A?LLMR @C MNCLCB. 5FGQ @CEGLQ RFC LCSPML'Q refractory period, GL UFGAF GR A?LLMR NPMBSAC ?LMRFCP ?ARGML NMRCLRG?J @CA?SQC GRQ QMBGSK AF?LLCJQ UGJJ LMR MNCL. "R RFC Q?KC RGKC, TMJR?ECE?RCB ,+ AF?LLCJQ MNCL, ?JJMUGLE ,+ RM JC?TC RFC ACJJ. "Q ,+ GMLQ JC?TC RFC ACJJ, RFC KCK@P?LC NMRCLRG?J MLAC ?E?GL @CAMKCQ LCE?RGTC. 5FC BGDDSQGML MD ,+ MSR MD RFC ACJJ ?ARS?JJW hyperpolarizes RFC ACJJ, GL RF?R RFC KCK@P?LC NMRCLRG?J @CAMKCQ KMPC LCE?RGTC RF?L RFC ACJJ'Q LMPK?J PCQRGLE NMRCLRG?J. "R RFGQ NMGLR, RFC QMBGSK AF?LLCJQ UGJJ PCRSPL RM RFCGP PCQRGLE QR?RC, KC?LGLE RFCW ?PC PC?BW RM MNCL ?E?GL GD RFC KCK@P?LC NMRCLRG?J ?E?GL CVACCBQ RFC RFPCQFMJB NMRCLRG?J. &TCLRS?JJW RFC CVRP? ,+ GMLQ BGDDSQC MSR MD RFC ACJJ RFPMSEF RFC NMR?QQGSK JC?I?EC AF?LLCJQ, @PGLEGLE RFC ACJJ DPMK GRQ FWNCPNMJ?PGXCB QR?RC, @?AI RM GRQ PCQRGLE KCK@P?LC NMRCLRG?J.
!@>LI< 35.11 4EB CLOJ>QFLK LC >K >@QFLK MLQBKQF>I @>K ?B AFSFABA FKQL CFSB PQBMP: (1) ! PQFJRIRP COLJ > PBKPLOV @BII LO >KLQEBO KBROLK @>RPBP QEB Q>ODBQ @BII QL ABMLI>OFWB QLT>OA QEB QEOBPELIA MLQBKQF>I. (2) )C QEB QEOBPELIA LC BU@FQ>QFLK FP OB>@EBA, >II .>+ @E>KKBIP LMBK >KA QEB JBJ?O>KB ABMLI>OFWBP. (3) !Q QEB MB>H >@QFLK MLQBKQF>I, ++ @E>KKBIP LMBK >KA ++ ?BDFKP QL IB>SB QEB @BII. !Q QEB P>JB QFJB, .>+ @E>KKBIP @ILPB. (4) 4EB JBJ?O>KB ?B@LJBP EVMBOMLI>OFWBA >P ++ FLKP @LKQFKRB QL IB>SB QEB @BII. 4EB EVMBOMLI>OFWBA JBJ?O>KB FP FK > OBCO>@QLOV MBOFLA >KA @>KKLQ CFOB. (5) 4EB ++ @E>KKBIP @ILPB >KA QEB .>+/++ QO>KPMLOQBO OBPQLOBP QEB OBPQFKD MLQBKQF>I.
0LQ>PPFRJ @E>KKBI ?IL@HBOP, PR@E >P >JFLA>OLKB >KA MOL@>FK>JFAB, TEF@E >OB RPBA QL QOB>Q >?KLOJ>I BIB@QOF@>I >@QFSFQV FK QEB EB>OQ, @>IIBA @>OAF>@ AVPOEVQEJF>, FJMBAB QEB JLSBJBKQ LC ++ QEOLRDE SLIQ>DB-D>QBA ++ @E>KKBIP. 7EF@E M>OQ LC QEB >@QFLK MLQBKQF>I TLRIA VLR BUMB@Q MLQ>PPFRJ @E>KKBIP QL >CCB@Q
#(!04%2 35 X 4(% .%26/53 3934%-
1001
!@>LI< 35.12 4EB >@QFLK MLQBKQF>I FP @LKAR@QBA ALTK QEB >ULK >P QEB >ULK JBJ?O>KB ABMLI>OFWBP, QEBK OBMLI>OFWBP.
5FGQ video (http://openstaxcollege.org/l/actionpotential) NPCQCLRQ ?L MTCPTGCU MD ?ARGML NMRCLRG?J. M>+1/3 '3* 9.+ P745'-'9/43 4, 9.+ A)9/43 P49+39/'1 'MP ?L ?ARGML NMRCLRG?J RM AMKKSLGA?RC GLDMPK?RGML RM ?LMRFCP LCSPML, GR KSQR RP?TCJ ?JMLE RFC ?VML ?LB PC?AF RFC ?VML RCPKGL?JQ UFCPC GR A?L GLGRG?RC LCSPMRP?LQKGRRCP PCJC?QC. 5FC QNCCB MD AMLBSARGML MD ?L ?ARGML NMRCLRG?J ?JMLE ?L ?VML GQ GLDJSCLACB @W @MRF RFC BG?KCRCP MD RFC ?VML ?LB RFC ?VMLsQ PCQGQR?LAC RM ASPPCLR JC?I. .WCJGL ?ARQ ?Q ?L GLQSJ?RMP RF?R NPCTCLRQ ASPPCLR DPMK JC?TGLE RFC ?VML; RFGQ GLAPC?QCQ RFC QNCCB MD ?ARGML NMRCLRG?J AMLBSARGML. *L BCKWCJGL?RGLE BGQC?QCQ JGIC KSJRGNJC QAJCPMQGQ, ?ARGML NMRCLRG?J AMLBSARGML QJMUQ @CA?SQC ASPPCLR JC?IQ DPMK NPCTGMSQJW GLQSJ?RCB ?VML ?PC?Q. 5FC LMBCQ MD 3?LTGCP, GJJSQRP?RCB GL Figure 35.13 ?PC E?NQ GL RFC KWCJGL QFC?RF ?JMLE RFC ?VML. 5FCQC SLKWCJGL?RCB QN?ACQ ?PC ?@MSR MLC KGAPMKCRCP JMLE ?LB AMLR?GL TMJR?EC E?RCB /?+ ?LB ,+ AF?LLCJQ. 'JMU MD GMLQ RFPMSEF RFCQC AF?LLCJQ, N?PRGASJ?PJW RFC /?+ AF?LLCJQ, PCECLCP?RCQ RFC ?ARGML NMRCLRG?J MTCP ?LB MTCP ?E?GL ?JMLE RFC ?VML. 5FGQ rHSKNGLEs MD RFC ?ARGML NMRCLRG?J DPMK MLC LMBC RM RFC LCVR GQ A?JJCB saltatory conduction. *D LMBCQ MD 3?LTGCP UCPC LMR NPCQCLR ?JMLE ?L ?VML, RFC ?ARGML NMRCLRG?J UMSJB NPMN?E?RC TCPW QJMUJW QGLAC /?+ ?LB ,+ AF?LLCJQ UMSJB F?TC RM AMLRGLSMSQJW PCECLCP?RC ?ARGML NMRCLRG?JQ ?R CTCPW NMGLR ?JMLE RFC ?VML GLQRC?B MD ?R QNCAGDGA NMGLRQ. /MBCQ MD 3?LTGCP ?JQM Q?TC CLCPEW DMP RFC LCSPML QGLAC RFC AF?LLCJQ MLJW LCCB RM @C NPCQCLR ?R RFC LMBCQ ?LB LMR ?JMLE RFC CLRGPC ?VML.
1002
#(!04%2 35 X 4(% .%26/53 3934%-
!@>LI< 35.13 .LABP LC 2>KSFBO >OB D>MP FK JVBIFK @LSBO>DB >ILKD >ULKP. .LABP @LKQ>FK SLIQ>DBD>QBA ++ >KA .>+ @E>KKBIP. !@QFLK MLQBKQF>IP QO>SBI ALTK QEB >ULK ?V GRJMFKD COLJ LKB KLAB QL QEB KBUQ.
.PE8GK@: /I8EJD@JJ@FE
5FC QWL?NQC MP tE?Nu GQ RFC NJ?AC UFCPC GLDMPK?RGML GQ RP?LQKGRRCB DPMK MLC LCSPML RM ?LMRFCP. 4WL?NQCQ SQS?JJW DMPK @CRUCCL ?VML RCPKGL?JQ ?LB BCLBPGRGA QNGLCQ, @SR RFGQ GQ LMR SLGTCPQ?JJW RPSC. 5FCPC ?PC ?JQM ?VML-RM-?VML, BCLBPGRC-RM-BCLBPGRC, ?LB ?VML-RM-ACJJ @MBW QWL?NQCQ. 5FC LCSPML RP?LQKGRRGLE RFC QGEL?J GQ A?JJCB RFC NPCQWL?NRGA LCSPML, ?LB RFC LCSPML PCACGTGLE RFC QGEL?J GQ A?JJCB RFC NMQRQWL?NRGA LCSPML. /MRC RF?R RFCQC BCQGEL?RGMLQ ?PC PCJ?RGTC RM ? N?PRGASJ?P QWL?NQCqKMQR LCSPMLQ ?PC @MRF NPCQWL?NRGA ?LB NMQRQWL?NRGA. 5FCPC ?PC RUM RWNCQ MD QWL?NQCQ: AFCKGA?J ?LB CJCARPGA?J. C.+2/)'1 !>3'58+ 8FCL ?L ?ARGML NMRCLRG?J PC?AFCQ RFC ?VML RCPKGL?J GR BCNMJ?PGXCQ RFC KCK@P?LC ?LB MNCLQ TMJR?ECE?RCB /?+ AF?LLCJQ. /?+ GMLQ CLRCP RFC ACJJ, DSPRFCP BCNMJ?PGXGLE RFC NPCQWL?NRGA KCK@P?LC. 5FGQ BCNMJ?PGX?RGML A?SQCQ TMJR?EC-E?RCB $?2+ AF?LLCJQ RM MNCL. $?JAGSK GMLQ CLRCPGLE RFC ACJJ GLGRG?RC ? QGEL?JGLE A?QA?BC RF?R A?SQCQ QK?JJ KCK@P?LC-@MSLB TCQGAJCQ, A?JJCB synaptic vesicles, AMLR?GLGLE LCSPMRP?LQKGRRCP KMJCASJCQ RM DSQC UGRF RFC NPCQWL?NRGA KCK@P?LC. 4WL?NRGA TCQGAJCQ ?PC QFMUL GL Figure 35.14, UFGAF GQ ?L GK?EC DPMK ? QA?LLGLE CJCARPML KGAPMQAMNC.
!@>LI< 35.14 4EFP MPBRAL@LILOBA FJ>DB Q>HBK TFQE > P@>KKFKD BIB@QOLK JF@OLP@LMB PELTP >K >ULK QBOJFK>I QE>Q T>P ?OLHBK LMBK QL OBSB>I PVK>MQF@ SBPF@IBP (?IRB >KA LO>KDB) FKPFAB QEB KBROLK. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V 4FK> #>OS>IEL, .)(-.)'-3; P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
'SQGML MD ? TCQGAJC UGRF RFC NPCQWL?NRGA KCK@P?LC A?SQCQ LCSPMRP?LQKGRRCP RM @C PCJC?QCB GLRM RFC synaptic cleft, RFC CVRP?ACJJSJ?P QN?AC @CRUCCL RFC NPCQWL?NRGA ?LB NMQRQWL?NRGA KCK@P?LCQ, ?Q GJJSQRP?RCB GL Figure 35.15. 5FC LCSPMRP?LQKGRRCP BGDDSQCQ ?APMQQ RFC QWL?NRGA AJCDR ?LB @GLBQ RM PCACNRMP NPMRCGLQ ML RFC NMQRQWL?NRGA KCK@P?LC.
#(!04%2 35 X 4(% .%26/53 3934%-
1003
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
5FC @GLBGLE MD ? QNCAGDGA LCSPMRP?LQKGRRCP A?SQCQ N?PRGASJ?P GML AF?LLCJQ, GL RFGQ A?QC JGE?LBE?RCB AF?LLCJQ, ML RFC NMQRQWL?NRGA KCK@P?LC RM MNCL. /CSPMRP?LQKGRRCPQ A?L CGRFCP F?TC CVAGR?RMPW MP GLFG@GRMPW CDDCARQ ML RFC NMQRQWL?NRGA KCK@P?LC, ?Q BCR?GJCB GL Table 35.1. 'MP CV?KNJC, UFCL ?ACRWJAFMJGLC GQ PCJC?QCB ?R RFC QWL?NQC @CRUCCL ? LCPTC ?LB KSQAJC (A?JJCB RFC LCSPMKSQASJ?P HSLARGML) @W ? NPCQWL?NRGA LCSPML, GR A?SQCQ NMQRQWL?NRGA /?+ AF?LLCJQ RM MNCL. /?+ CLRCPQ RFC NMQRQWL?NRGA ACJJ ?LB A?SQCQ RFC NMQRQWL?NRGA KCK@P?LC RM BCNMJ?PGXC. 5FGQ BCNMJ?PGX?RGML GQ A?JJCB ?L excitatory postsynaptic potential (EPSP) ?LB K?ICQ RFC NMQRQWL?NRGA LCSPML KMPC JGICJW RM DGPC ?L ?ARGML NMRCLRG?J. 3CJC?QC MD LCSPMRP?LQKGRRCP ?R GLFG@GRMPW QWL?NQCQ A?SQCQ inhibitory postsynaptic potentials (IPSPs), ? FWNCPNMJ?PGX?RGML MD RFC NPCQWL?NRGA KCK@P?LC. 'MP CV?KNJC, UFCL RFC LCSPMRP?LQKGRRCP ("#" (E?KK?-?KGLM@SRWPGA ?AGB) GQ PCJC?QCB DPMK ? NPCQWL?NRGA LCSPML, GR @GLBQ RM ?LB MNCLQ $J-
1004
#(!04%2 35 X 4(% .%26/53 3934%-
AF?LLCJQ. $J- GMLQ CLRCP RFC ACJJ ?LB FWNCPNMJ?PGXCQ RFC KCK@P?LC, K?IGLE RFC LCSPML JCQQ JGICJW RM DGPC ?L ?ARGML NMRCLRG?J. 0LAC LCSPMRP?LQKGQQGML F?Q MAASPPCB, RFC LCSPMRP?LQKGRRCP KSQR @C PCKMTCB DPMK RFC QWL?NRGA AJCDR QM RFC NMQRQWL?NRGA KCK@P?LC A?L tPCQCRu ?LB @C PC?BW RM PCACGTC ?LMRFCP QGEL?J. 5FGQ A?L @C ?AAMKNJGQFCB GL RFPCC U?WQ: RFC LCSPMRP?LQKGRRCP A?L BGDDSQC ?U?W DPMK RFC QWL?NRGA AJCDR, GR A?L @C BCEP?BCB @W CLXWKCQ GL RFC QWL?NRGA AJCDR, MP GR A?L @C PCAWAJCB (QMKCRGKCQ A?JJCB PCSNR?IC) @W RFC NPCQWL?NRGA LCSPML. 4CTCP?J BPSEQ ?AR ?R RFGQ QRCN MD LCSPMRP?LQKGQQGML. 'MP CV?KNJC, QMKC BPSEQ RF?R ?PC EGTCL RM "JXFCGKCPsQ N?RGCLRQ UMPI @W GLFG@GRGLE ?ACRWJAFMJGLCQRCP?QC, RFC CLXWKC RF?R BCEP?BCQ ?ACRWJAFMJGLC. 5FGQ GLFG@GRGML MD RFC CLXWKC CQQCLRG?JJW GLAPC?QCQ LCSPMRP?LQKGQQGML ?R QWL?NQCQ RF?R PCJC?QC ?ACRWJAFMJGLC. 0LAC PCJC?QCB, RFC ?ACRWJAFMJGLC QR?WQ GL RFC AJCDR ?LB A?L AMLRGLS?JJW @GLB ?LB SL@GLB RM NMQRQWL?NRGA PCACNRMPQ.
)<LIFKI8EJD@KKJFKB !JFKL >@FA .BROLMBMQFAB /89C< 35.2 u $LM>JFKB, PBOLQLKFK, KLOBMFKBMEOFKB
O8DGC<
'F:8K@FE
#.3 >KA/LO 0.3 #.3 >KA/LO 0.3
'IV@FKB, DIRQ>J>QB, >PM>OQ>QB, D>JJ> >JFKL?RQVOF@ #.3 >@FA 3R?PQ>K@B 0, BKALOMEFKP #.3 >KA/LO 0.3
E1+)97/)'1 !>3'58+ 8FGJC CJCARPGA?J QWL?NQCQ ?PC DCUCP GL LSK@CP RF?L AFCKGA?J QWL?NQCQ, RFCW ?PC DMSLB GL ?JJ LCPTMSQ QWQRCKQ ?LB NJ?W GKNMPR?LR ?LB SLGOSC PMJCQ. 5FC KMBC MD LCSPMRP?LQKGQQGML GL CJCARPGA?J QWL?NQCQ GQ OSGRC BGDDCPCLR DPMK RF?R GL AFCKGA?J QWL?NQCQ. *L ?L CJCARPGA?J QWL?NQC, RFC NPCQWL?NRGA ?LB NMQRQWL?NRGA KCK@P?LCQ ?PC TCPW AJMQC RMECRFCP ?LB ?PC ?ARS?JJW NFWQGA?JJW AMLLCARCB @W AF?LLCJ NPMRCGLQ DMPKGLE E?N HSLARGMLQ. (?N HSLARGMLQ ?JJMU ASPPCLR RM N?QQ BGPCARJW DPMK MLC ACJJ RM RFC LCVR. *L ?BBGRGML RM RFC GMLQ RF?R A?PPW RFGQ ASPPCLR, MRFCP KMJCASJCQ, QSAF ?Q "51, A?L BGDDSQC RFPMSEF RFC J?PEC E?N HSLARGML NMPCQ. 5FCPC ?PC ICW BGDDCPCLACQ @CRUCCL AFCKGA?J ?LB CJCARPGA?J QWL?NQCQ. #CA?SQC AFCKGA?J QWL?NQCQ BCNCLB ML RFC PCJC?QC MD LCSPMRP?LQKGRRCP KMJCASJCQ DPMK QWL?NRGA TCQGAJCQ RM N?QQ ML RFCGP QGEL?J, RFCPC GQ ?L ?NNPMVGK?RCJW MLC KGJJGQCAMLB BCJ?W @CRUCCL UFCL RFC ?VML NMRCLRG?J PC?AFCQ RFC NPCQWL?NRGA RCPKGL?J ?LB UFCL RFC LCSPMRP?LQKGRRCP JC?BQ RM MNCLGLE MD NMQRQWL?NRGA GML AF?LLCJQ. "BBGRGML?JJW, RFGQ QGEL?JGLE GQ SLGBGPCARGML?J. 4GEL?JGLE GL CJCARPGA?J QWL?NQCQ, GL AMLRP?QR, GQ TGPRS?JJW GLQR?LR?LCMSQ (UFGAF GQ GKNMPR?LR DMP QWL?NQCQ GLTMJTCB GL ICW PCDJCVCQ), ?LB QMKC CJCARPGA?J QWL?NQCQ ?PC @GBGPCARGML?J. &JCARPGA?J QWL?NQCQ ?PC ?JQM KMPC PCJG?@JC ?Q RFCW ?PC JCQQ JGICJW RM @C @JMAICB, ?LB RFCW ?PC GKNMPR?LR DMP QWLAFPMLGXGLE RFC CJCARPGA?J ?ARGTGRW MD ? EPMSN MD LCSPMLQ. 'MP CV?KNJC, CJCARPGA?J QWL?NQCQ GL RFC RF?J?KSQ ?PC RFMSEFR RM PCESJ?RC QJMU-U?TC QJCCN, ?LB BGQPSNRGML MD RFCQC QWL?NQCQ A?L A?SQC QCGXSPCQ.
.@>E8C .LDD8K@FE
4MKCRGKCQ ? QGLEJC &141 GQ QRPMLE CLMSEF RM GLBSAC ?L ?ARGML NMRCLRG?J GL RFC NMQRQWL?NRGA LCSPML, @SR MDRCL KSJRGNJC NPCQWL?NRGA GLNSRQ KSQR APC?RC &141Q ?PMSLB RFC Q?KC RGKC DMP RFC NMQRQWL?NRGA LCSPML RM @C QSDDGAGCLRJW BCNMJ?PGXCB RM DGPC ?L ?ARGML NMRCLRG?J. 5FGQ NPMACQQ GQ A?JJCB summation ?LB MAASPQ ?R RFC ?VML FGJJMAI, ?Q GJJSQRP?RCB GL Figure 35.16. "BBGRGML?JJW, MLC LCSPML MDRCL F?Q GLNSRQ DPMK K?LW NPCQWL?NRGA LCSPMLQqQMKC CVAGR?RMPW ?LB QMKC GLFG@GRMPWqQM *141Q A?L A?LACJ MSR &141Q ?LB TGAC TCPQ?. *R GQ RFC LCR AF?LEC GL NMQRQWL?NRGA KCK@P?LC TMJR?EC RF?R BCRCPKGLCQ UFCRFCP RFC NMQRQWL?NRGA ACJJ F?Q PC?AFCB GRQ RFPCQFMJB MD CVAGR?RGML LCCBCB RM DGPC ?L ?ARGML NMRCLRG?J. 5MECRFCP, QWL?NRGA QSKK?RGML ?LB RFC RFPCQFMJB DMP CVAGR?RGML ?AR ?Q ? DGJRCP QM RF?R P?LBMK tLMGQCu GL RFC QWQRCK GQ LMR RP?LQKGRRCB ?Q GKNMPR?LR GLDMPK?RGML.
#(!04%2 35 X 4(% .%26/53 3934%-
1005
!@>LI< 35.16 ! PFKDIB KBROLK @>K OB@BFSB ?LQE BU@FQ>QLOV >KA FKEF?FQLOV FKMRQP COLJ JRIQFMIB KBROLKP, OBPRIQFKD FK IL@>I JBJ?O>KB ABMLI>OFW>QFLK (%030 FKMRQ) >KA EVMBOMLI>OFW>QFLK ()030 FKMRQ). !II QEBPB FKMRQP >OB >AABA QLDBQEBO >Q QEB >ULK EFIIL@H. )C QEB %030P >OB PQOLKD BKLRDE QL LSBO@LJB QEB )030P >KA OB>@E QEB QEOBPELIA LC BU@FQ>QFLK, QEB KBROLK TFII CFOB.
1006
#(!04%2 35 X 4(% .%26/53 3934%-
BI8@E-:FDGLKQBO>I P@IBOLPFP (!,3, >IPL @>IIBA ,LR 'BEOFDwP $FPB>PB) FP > KBROLILDF@>I AFPB>PB @E>O>@QBOFWBA ?V QEB ABDBKBO>QFLK LC QEB JLQLO KBROLKP QE>Q @LKQOLI SLIRKQ>OV JLSBJBKQP. 4EB AFPB>PB ?BDFKP TFQE JRP@IB TB>HBKFKD >KA I>@H LC @LLOAFK>QFLK >KA BSBKQR>IIV ABPQOLVP QEB KBROLKP QE>Q @LKQOLI PMBB@E, ?OB>QEFKD, >KA PT>IILTFKD; FK QEB BKA, QEB AFPB>PB @>K IB>A QL M>O>IVPFP. !Q QE>Q MLFKQ, M>QFBKQP OBNRFOB >PPFPQ>K@B COLJ J>@EFKBP QL ?B >?IB QL ?OB>QEB >KA QL @LJJRKF@>QB. 3BSBO>I PMB@F>I QB@EKLILDFBP E>SB ?BBK ABSBILMBA QL >IILT xIL@HBA-FKy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u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
!@>LI< 35.17 7FQE ?O>FK-@LJMRQBO FKQBOC>@B QB@EKLILDV, KBRO>I PFDK>IP COLJ > M>O>IVWBA M>QFBKQ >OB @LIIB@QBA, AB@LABA, >KA QEBK CBA QL > QLLI, PR@E >P > @LJMRQBO, > TEBBI@E>FO, LO > OL?LQF@ >OJ.
#(!04%2 35 X 4(% .%26/53 3934%-
1007
8?RAF this video (http://openstaxcollege.org/l/paralyzation) GL UFGAF ? N?P?JWXCB UMK?L SQC ? @P?GL-AMLRPMJJCB PM@MRGA ?PK RM @PGLE ? BPGLI RM FCP KMSRF, ?KMLE MRFCP GK?ECQ MD @P?GL-AMKNSRCP GLRCPD?AC RCAFLMJMEW GL ?ARGML.
.PE8GK@: +C8JK@:@KP
4WL?NQCQ ?PC LMR QR?RGA QRPSARSPCQ. 5FCW A?L @C UC?ICLCB MP QRPCLERFCLCB. 5FCW A?L @C @PMICL, ?LB LCU QWL?NQCQ A?L @C K?BC. 4WL?NRGA NJ?QRGAGRW ?JJMUQ DMP RFCQC AF?LECQ, UFGAF ?PC ?JJ LCCBCB DMP ? DSLARGMLGLE LCPTMSQ QWQRCK. *L D?AR, QWL?NRGA NJ?QRGAGRW GQ RFC @?QGQ MD JC?PLGLE ?LB KCKMPW. 5UM NPMACQQCQ GL N?PRGASJ?P, JMLE-RCPK NMRCLRG?RGML (-51) ?LB JMLE-RCPK BCNPCQQGML (-5%) ?PC GKNMPR?LR DMPKQ MD QWL?NRGA NJ?QRGAGRW RF?R MAASP GL QWL?NQCQ GL RFC FGNNMA?KNSQ, ? @P?GL PCEGML RF?R GQ GLTMJTCB GL QRMPGLE KCKMPGCQ. L43--9+72 P49+39/'9/43 (L"P) Long-term potentiation (LTP) GQ ? NCPQGQRCLR QRPCLERFCLGLE MD ? QWL?NRGA AMLLCARGML. -51 GQ @?QCB ML RFC )C@@G?L NPGLAGNJC: ACJJQ RF?R DGPC RMECRFCP UGPC RMECRFCP. 5FCPC ?PC T?PGMSQ KCAF?LGQKQ, LMLC DSJJW SLBCPQRMMB, @CFGLB RFC QWL?NRGA QRPCLERFCLGLE QCCL UGRF -51. 0LC ILMUL KCAF?LGQK GLTMJTCQ ? RWNC MD NMQRQWL?NRGA EJSR?K?RC PCACNRMP, A?JJCB /.%" (/-.CRFWJ-%-?QN?PR?RC) PCACNRMPQ, QFMUL GL Figure 35.18. 5FCQC PCACNRMPQ ?PC LMPK?JJW @JMAICB @W K?ELCQGSK GMLQ; FMUCTCP, UFCL RFC NMQRQWL?NRGA LCSPML GQ BCNMJ?PGXCB @W KSJRGNJC NPCQWL?NRGA GLNSRQ GL OSGAI QSAACQQGML (CGRFCP DPMK MLC LCSPML MP KSJRGNJC LCSPMLQ), RFC K?ELCQGSK GMLQ ?PC DMPACB MSR ?JJMUGLE $? GMLQ RM N?QQ GLRM RFC NMQRQWL?NRGA ACJJ. /CVR, $?2+ GMLQ CLRCPGLE RFC ACJJ GLGRG?RC ? QGEL?JGLE A?QA?BC RF?R A?SQCQ ? BGDDCPCLR RWNC MD EJSR?K?RC PCACNRMP, A?JJCB ".1" (i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ong-term depression (LTD) GQ CQQCLRG?JJW RFC PCTCPQC MD -51: GR GQ ? JMLE-RCPK UC?ICLGLE MD ? QWL?NRGA AMLLCARGML. 0LC KCAF?LGQK ILMUL RM A?SQC -5% ?JQM GLTMJTCQ ".1" PCACNRMPQ. *L RFGQ QGRS?RGML, A?JAGSK RF?R CLRCPQ RFPMSEF /.%" PCACNRMPQ GLGRG?RCQ ? BGDDCPCLR QGEL?JGLE A?QA?BC, UFGAF PCQSJRQ GL RFC PCKMT?J MD ".1" PCACNRMPQ DPMK RFC NMQRQWL?NRGA KCK@P?LC, ?Q GJJSQRP?RCB GL Figure 35.18. 5FC BCAPC?QC GL ".1" PCACNRMPQ GL RFC KCK@P?LC K?ICQ RFC NMQRQWL?NRGA LCSPML JCQQ PCQNMLQGTC RM EJSR?K?RC PCJC?QCB DPMK RFC NPCQWL?NRGA LCSPML. 8FGJC GR K?W QCCK AMSLRCPGLRSGRGTC, -5% K?W @C HSQR ?Q GKNMPR?LR DMP JC?PLGLE ?LB KCKMPW ?Q -51. 5FC UC?ICLGLE ?LB NPSLGLE MD SLSQCB QWL?NQCQ ?JJMUQ DMP SLGKNMPR?LR AMLLCARGMLQ RM @C JMQR ?LB K?ICQ RFC QWL?NQCQ RF?R F?TC SLBCPEMLC -51 RF?R KSAF QRPMLECP @W AMKN?PGQML.
1008
#(!04%2 35 X 4(% .%26/53 3934%-
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
35.3 R /?< C<EKI8C )<IMFLJ .PJK<D

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z *BCLRGDW RFC QNGL?J AMPB, ACPC@P?J JM@CQ, ?LB MRFCP @P?GL ?PC?Q ML ? BG?EP?K MD RFC @P?GL z %CQAPG@C RFC @?QGA DSLARGMLQ MD RFC QNGL?J AMPB, ACPC@P?J JM@CQ, ?LB MRFCP @P?GL ?PC?Q 5FC ACLRP?J LCPTMSQ QWQRCK ($/4) GQ K?BC SN MD RFC @P?GL, ? N?PR MD UFGAF GQ QFMUL GL Figure 35.19 ?LB QNGL?J AMPB ?LB GQ AMTCPCB UGRF RFPCC J?WCPQ MD NPMRCARGTC AMTCPGLEQ A?JJCB meninges (DPMK RFC (PCCI UMPB DMP KCK@P?LC). 5FC MSRCPKMQR J?WCP GQ RFC dura mater (-?RGL DMP tF?PB KMRFCPu). "Q RFC -?RGL QSEECQRQ, RFC NPGK?PW DSLARGML DMP RFGQ RFGAI J?WCP GQ RM NPMRCAR RFC @P?GL ?LB QNGL?J AMPB. 5FC BSP? K?RCP ?JQM AMLR?GLQ TCGL-JGIC QRPSARSPCQ RF?R A?PPW @JMMB DPMK RFC @P?GL @?AI RM RFC FC?PR. 5FC KGBBJC J?WCP GQ RFC UC@-JGIC arachnoid mater. 5FC J?QR J?WCP GQ RFC pia mater (-?RGL DMP tQMDR KMRFCPu), UFGAF BGPCARJW AMLR?ARQ ?LB AMTCPQ RFC @P?GL ?LB QNGL?J AMPB JGIC NJ?QRGA UP?N. 5FC QN?AC @CRUCCL RFC ?P?AFLMGB ?LB NG? K?RCPQ GQ DGJJCB UGRF cerebrospinal fluid (CSF). $4' GQ NPMBSACB @W ? RGQQSC A?JJCB choroid plexus GL DJSGB-DGJJCB AMKN?PRKCLRQ GL RFC $/4 A?JJCB ventricles. 5FC @P?GL DJM?RQ GL $4', UFGAF ?ARQ ?Q ? ASQFGML ?LB QFMAI ?@QMP@CP ?LB K?ICQ RFC @P?GL LCSRP?JJW @SMW?LR. $4' ?JQM DSLARGMLQ RM AGPASJ?RC AFCKGA?J QS@QR?LACQ RFPMSEFMSR RFC @P?GL ?LB GLRM RFC QNGL?J AMPB.
#(!04%2 35 X 4(% .%26/53 3934%-
1009
5FC CLRGPC @P?GL AMLR?GLQ MLJW ?@MSR 8.5 R?@JCQNMMLQ MD $4', @SR $4' GQ AMLQR?LRJW NPMBSACB GL RFC TCLRPGAJCQ. 5FGQ APC?RCQ ? NPM@JCK UFCL ? TCLRPGAJC GQ @JMAICBqRFC $4' @SGJBQ SN ?LB APC?RCQ QUCJJGLE ?LB RFC @P?GL GQ NSQFCB ?E?GLQR RFC QISJJ. 5FGQ QUCJJGLE AMLBGRGML GQ A?JJCB FWBPMACNF?JSQ (tU?RCP FC?Bu) ?LB A?L A?SQC QCGXSPCQ, AMELGRGTC NPM@JCKQ, ?LB CTCL BC?RF GD ? QFSLR GQ LMR GLQCPRCB RM PCKMTC RFC DJSGB ?LB NPCQQSPC.
!@>LI< 35.19 4EB @BOB?O>I @LOQBU FP @LSBOBA ?V QEOBB I>VBOP LC JBKFKDBP: QEB ARO>, >O>@EKLFA, >KA MF> J>QBOP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V 'O>VwP !K>QLJV)
BI8@E
5FC @P?GL GQ RFC N?PR MD RFC ACLRP?J LCPTMSQ QWQRCK RF?R GQ AMLR?GLCB GL RFC AP?LG?J A?TGRW MD RFC QISJJ. *R GLAJSBCQ RFC ACPC@P?J AMPRCV, JGK@GA QWQRCK, @?Q?J E?LEJG?, RF?J?KSQ, FWNMRF?J?KSQ, ?LB ACPC@CJJSK. 5FCPC ?PC RFPCC BGDDCPCLR U?WQ RF?R ? @P?GL A?L @C QCARGMLCB GL MPBCP RM TGCU GLRCPL?J QRPSARSPCQ: ? Q?EGRR?J QCARGML ASRQ RFC @P?GL JCDR RM PGEFR, ?Q QFMUL GL Figure 35.21b, ? AMPML?J QCARGML ASRQ RFC @P?GL DPMLR RM @?AI, ?Q QFMUL GL Figure 35.20a, ?LB ? FMPGXMLR?J QCARGML ASRQ RFC @P?GL RMN RM @MRRMK. C+7+(7'1 C479+= 5FC MSRCPKMQR N?PR MD RFC @P?GL GQ ? RFGAI NGCAC MD LCPTMSQ QWQRCK RGQQSC A?JJCB RFC cerebral cortex, UFGAF GQ DMJBCB GLRM FGJJQ A?JJCB gyri (QGLESJ?P: EWPSQ) ?LB T?JJCWQ A?JJCB sulci (QGLESJ?P: QSJASQ). 5FC AMPRCV GQ K?BC SN MD RUM FCKGQNFCPCQqPGEFR ?LB JCDRqUFGAF ?PC QCN?P?RCB @W ? J?PEC QSJASQ. " RFGAI DG@CP @SLBJC A?JJCB RFC corpus callosum (-?RGL: tRMSEF @MBWu) AMLLCARQ RFC RUM FCKGQNFCPCQ ?LB ?JJMUQ GLDMPK?RGML RM @C N?QQCB DPMK MLC QGBC RM RFC MRFCP. "JRFMSEF RFCPC ?PC QMKC @P?GL DSLARGMLQ RF?R ?PC JMA?JGXCB KMPC RM MLC FCKGQNFCPC RF?L RFC MRFCP, RFC DSLARGMLQ MD RFC RUM FCKGQNFCPCQ ?PC J?PECJW PCBSLB?LR. *L D?AR, QMKCRGKCQ (TCPW P?PCJW) ?L CLRGPC FCKGQNFCPC GQ PCKMTCB RM RPC?R QCTCPC CNGJCNQW. 8FGJC N?RGCLRQ BM QSDDCP QMKC BCDGAGRQ DMJJMUGLE RFC QSPECPW, RFCW A?L F?TC QSPNPGQGLEJW DCU NPM@JCKQ, CQNCAG?JJW UFCL RFC QSPECPW GQ NCPDMPKCB ML AFGJBPCL UFM F?TC TCPW GKK?RSPC LCPTMSQ QWQRCKQ.
(@) (?) !@>LI< 35.20 4EBPB FIIRPQO>QFLKP PELT QEB (>) @LOLK>I >KA (?) P>DFQQ>I PB@QFLKP LC QEB ERJ>K ?O>FK.
*L MRFCP QSPECPGCQ RM RPC?R QCTCPC CNGJCNQW, RFC AMPNSQ A?JJMQSK GQ ASR GLQRC?B MD PCKMTGLE ?L CLRGPC FCKGQNFCPC. 5FGQ A?SQCQ ? AMLBGRGML A?JJCB QNJGR-@P?GL, UFGAF EGTCQ GLQGEFRQ GLRM SLGOSC DSLARGMLQ MD RFC
1010
#(!04%2 35 X 4(% .%26/53 3934%-
RUM FCKGQNFCPCQ. 'MP CV?KNJC, UFCL ?L M@HCAR GQ NPCQCLRCB RM N?RGCLRQs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
4CC this website (http://openstaxcollege.org/l/split-brain) RM JC?PL KMPC ?@MSR QNJGR-@P?GL N?RGCLRQ ?LB RM NJ?W ? E?KC UFCPC WMS A?L KMBCJ RFC QNJGR-@P?GL CVNCPGKCLRQ WMSPQCJD. &?AF AMPRGA?J FCKGQNFCPC AMLR?GLQ PCEGMLQ A?JJCB JM@CQ RF?R ?PC GLTMJTCB GL BGDDCPCLR DSLARGMLQ. 4AGCLRGQRQ SQC T?PGMSQ RCAFLGOSCQ RM BCRCPKGLC UF?R @P?GL ?PC?Q ?PC GLTMJTCB GL BGDDCPCLR DSLARGMLQ: RFCW CV?KGLC N?RGCLRQ UFM F?TC F?B GLHSPGCQ MP BGQC?QCQ RF?R ?DDCAR QNCAGDGA ?PC?Q ?LB QCC FMU RFMQC ?PC?Q ?PC PCJ?RCB RM DSLARGML?J BCDGAGRQ. 5FCW ?JQM AMLBSAR ?LGK?J QRSBGCQ UFCPC RFCW QRGKSJ?RC @P?GL ?PC?Q ?LB QCC GD RFCPC ?PC ?LW @CF?TGMP?J AF?LECQ. 5FCW SQC ? RCAFLGOSC A?JJCB RP?LQK?ELCRGA QRGKSJ?RGML (5.4) RM RCKNMP?PGJW BC?ARGT?RC QNCAGDGA N?PRQ MD RFC AMPRCV SQGLE QRPMLE K?ELCRQ NJ?ACB MSRQGBC RFC FC?B; ?LB RFCW SQC DSLARGML?J K?ELCRGA PCQML?LAC GK?EGLE (D.3*) RM JMMI ?R AF?LECQ GL MVWECL?RCB @JMMB DJMU GL N?PRGASJ?P @P?GL PCEGMLQ RF?R AMPPCJ?RC UGRF QNCAGDGA @CF?TGMP?J R?QIQ. 5FCQC RCAFLGOSCQ, ?LB MRFCPQ, F?TC EGTCL EPC?R GLQGEFR GLRM RFC DSLARGMLQ MD BGDDCPCLR @P?GL PCEGMLQ @SR F?TC ?JQM QFMUCB RF?R ?LW EGTCL @P?GL ?PC? A?L @C GLTMJTCB GL KMPC RF?L MLC @CF?TGMP MP NPMACQQ, ?LB ?LW EGTCL @CF?TGMP MP NPMACQQ ECLCP?JJW GLTMJTCQ LCSPMLQ GL KSJRGNJC @P?GL ?PC?Q. 5F?R @CGLE Q?GB, C?AF FCKGQNFCPC MD RFC K?KK?JG?L ACPC@P?J AMPRCV A?L @C @PMICL BMUL GLRM DMSP DSLARGML?JJW ?LB QN?RG?JJW BCDGLCB JM@CQ: DPMLR?J, N?PGCR?J, RCKNMP?J, ?LB MAAGNGR?J. Figure 35.21 GJJSQRP?RCQ RFCQC DMSP JM@CQ MD RFC FSK?L ACPC@P?J AMPRCV.
!@>LI< 35.21 4EB ERJ>K @BOB?O>I @LOQBU FK@IRABP QEB COLKQ>I, M>OFBQ>I, QBJMLO>I, >KA L@@FMFQ>I IL?BP.
5FC frontal lobe GQ JMA?RCB ?R RFC DPMLR MD RFC @P?GL, MTCP RFC CWCQ. 5FGQ JM@C AMLR?GLQ RFC MJD?ARMPW @SJ@, UFGAF NPMACQQCQ QKCJJQ. 5FC DPMLR?J JM@C ?JQM AMLR?GLQ RFC KMRMP AMPRCV, UFGAF GQ GKNMPR?LR DMP NJ?LLGLE ?LB GKNJCKCLRGLE KMTCKCLR. "PC?Q UGRFGL RFC KMRMP AMPRCV K?N RM BGDDCPCLR KSQAJC EPMSNQ, ?LB RFCPC GQ QMKC MPE?LGX?RGML RM RFGQ K?N, ?Q QFMUL GL Figure 35.22. 'MP CV?KNJC, RFC LCSPMLQ RF?R AMLRPMJ KMTCKCLR MD RFC DGLECPQ ?PC LCVR RM RFC LCSPMLQ RF?R AMLRPMJ KMTCKCLR MD RFC F?LB. /CSPMLQ GL RFC DPMLR?J JM@C ?JQM AMLRPMJ AMELGRGTC DSLARGMLQ JGIC K?GLR?GLGLE ?RRCLRGML, QNCCAF, ?LB BCAGQGML-K?IGLE. 4RSBGCQ MD FSK?LQ UFM F?TC B?K?ECB RFCGP DPMLR?J JM@CQ QFMU RF?R N?PRQ MD RFGQ ?PC? ?PC GLTMJTCB GL NCPQML?JGRW, QMAG?JGX?RGML, ?LB ?QQCQQGLE PGQI.
#(!04%2 35 X 4(% .%26/53 3934%-
1011
!@>LI< 35.22 $FCCBOBKQ M>OQP LC QEB JLQLO @LOQBU @LKQOLI AFCCBOBKQ JRP@IB DOLRMP. -RP@IB DOLRMP QE>Q >OB KBFDE?LOP FK QEB ?LAV >OB DBKBO>IIV @LKQOLIIBA ?V KBFDE?LOFKD OBDFLKP LC QEB JLQLO @LOQBU >P TBII. &LO BU>JMIB, QEB KBROLKP QE>Q @LKQOLI CFKDBO JLSBJBKQ >OB KB>O QEB KBROLKP QE>Q @LKQOLI E>KA JLSBJBKQ.
5FC parietal lobe GQ JMA?RCB ?R RFC RMN MD RFC @P?GL. /CSPMLQ GL RFC N?PGCR?J JM@C ?PC GLTMJTCB GL QNCCAF ?LB ?JQM PC?BGLE. 5UM MD RFC N?PGCR?J JM@CsQ K?GL DSLARGMLQ ?PC NPMACQQGLE somatosensationqRMSAF QCLQ?RGMLQ JGIC NPCQQSPC, N?GL, FC?R, AMJBq?LB NPMACQQGLE proprioceptionqRFC QCLQC MD FMU N?PRQ MD RFC @MBW ?PC MPGCLRCB GL QN?AC. 5FC N?PGCR?J JM@C AMLR?GLQ ? QMK?RMQCLQMPW K?N MD RFC @MBW QGKGJ?P RM RFC KMRMP AMPRCV. 5FC occipital lobe GQ JMA?RCB ?R RFC @?AI MD RFC @P?GL. *R GQ NPGK?PGJW GLTMJTCB GL TGQGMLqQCCGLE, PCAMELGXGLE, ?LB GBCLRGDWGLE RFC TGQS?J UMPJB. 5FC temporal lobe GQ JMA?RCB ?R RFC @?QC MD RFC @P?GL @W WMSP C?PQ ?LB GQ NPGK?PGJW GLTMJTCB GL NPMACQQGLE ?LB GLRCPNPCRGLE QMSLBQ. *R ?JQM AMLR?GLQ RFC hippocampus ((PCCI DMP tQC?FMPQCu)q? QRPSARSPC RF?R NPMACQQCQ KCKMPW DMPK?RGML. 5FC FGNNMA?KNSQ GQ GJJSQRP?RCB GL Figure 35.24. 5FC PMJC MD RFC FGNNMA?KNSQ GL KCKMPW U?Q N?PRG?JJW BCRCPKGLCB @W QRSBWGLE MLC D?KMSQ CNGJCNRGA N?RGCLR, )., UFM F?B @MRF QGBCQ MD FGQ FGNNMA?KNSQ PCKMTCB GL ?L ?RRCKNR RM ASPC FGQ CNGJCNQW. )GQ QCGXSPCQ UCLR ?U?W, @SR FC AMSJB LM JMLECP DMPK LCU KCKMPGCQ (?JRFMSEF FC AMSJB PCKCK@CP QMKC D?ARQ DPMK @CDMPC FGQ QSPECPW ?LB AMSJB JC?PL LCU KMRMP R?QIQ).
1012
#(!04%2 35 X 4(% .%26/53 3934%-
C<I<9I8C CFIK<O
#LJM>OBA QL LQEBO SBOQB?O>QBP, J>JJ>IP E>SB BU@BMQFLK>IIV I>ODB ?O>FKP CLO QEBFO ?LAV PFWB. !K BKQFOB >IIFD>QLOw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
!@>LI< 35.23 ->JJ>IP E>SB I>ODBO ?O>FK-QL-?LAV O>QFLP QE>K LQEBO SBOQB?O>QBP. 7FQEFK J>JJ>IP, FK@OB>PBA @LOQF@>I CLIAFKD >KA PROC>@B >OB> FP @LOOBI>QBA TFQE @LJMIBU ?BE>SFLO.
B'8'1 G'3-1/' *LRCPAMLLCARCB @P?GL ?PC?Q A?JJCB RFC basal ganglia (MP basal nuclei), QFMUL GL Figure 35.20b, NJ?W GKNMPR?LR PMJCQ GL KMTCKCLR AMLRPMJ ?LB NMQRSPC. %?K?EC RM RFC @?Q?J E?LEJG?, ?Q GL 1?PIGLQMLsQ BGQC?QC, JC?BQ RM KMRMP GKN?GPKCLRQ JGIC ? QFSDDJGLE E?GR UFCL U?JIGLE. 5FC @?Q?J E?LEJG? ?JQM PCESJ?RC KMRGT?RGML. 'MP CV?KNJC, UFCL ? U?QN QRGLE JCB RM @GJ?RCP?J @?Q?J E?LEJG? B?K?EC GL ? 25-WC?P-MJB @SQGLCQQK?L, FC @CE?L RM QNCLB ?JJ FGQ B?WQ GL @CB ?LB QFMUCB LM GLRCPCQR GL ?LWRFGLE MP ?LW@MBW. #SR UFCL FC U?Q CVRCPL?JJW QRGKSJ?RCBq?Q UFCL QMKCMLC ?QICB RM NJ?W ? A?PB E?KC UGRF FGKqFC U?Q ?@JC RM DSLARGML LMPK?JJW. *LRCPCQRGLEJW, FC ?LB MRFCP QGKGJ?P N?RGCLRQ BM LMR PCNMPR DCCJGLE @MPCB MP DPSQRP?RCB @W RFCGP QR?RC. ".'1'2:8 5FC thalamus ((PCCI DMP tGLLCP AF?K@CPu), GJJSQRP?RCB GL Figure 35.24, ?ARQ ?Q ? E?RCU?W RM ?LB DPMK RFC AMPRCV. *R PCACGTCQ QCLQMPW ?LB KMRMP GLNSRQ DPMK RFC @MBW ?LB ?JQM PCACGTCQ DCCB@?AI DPMK RFC AMPRCV. 5FGQ DCCB@?AI KCAF?LGQK A?L KMBSJ?RC AMLQAGMSQ ?U?PCLCQQ MD QCLQMPW ?LB KMRMP GLNSRQ BCNCLBGLE ML RFC ?RRCLRGML ?LB ?PMSQ?J QR?RC MD RFC ?LGK?J. 5FC RF?J?KSQ FCJNQ PCESJ?RC AMLQAGMSQLCQQ, ?PMSQ?J, ?LB QJCCN QR?RCQ. " P?PC ECLCRGA BGQMPBCP A?JJCB D?R?J D?KGJG?J GLQMKLG? A?SQCQ RFC BCECLCP?RGML MD RF?J?KGA LCSPMLQ ?LB EJG?. 5FGQ BGQMPBCP NPCTCLRQ ?DDCARCB N?RGCLRQ DPMK @CGLE ?@JC RM QJCCN, ?KMLE MRFCP QWKNRMKQ, ?LB GQ CTCLRS?JJW D?R?J.
#(!04%2 35 X 4(% .%26/53 3934%-
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!@>LI< 35.24 4EB IFJ?F@ PVPQBJ OBDRI>QBP BJLQFLK >KA LQEBO ?BE>SFLOP. )Q FK@IRABP M>OQP LC QEB @BOB?O>I @LOQBU IL@>QBA KB>O QEB @BKQBO LC QEB ?O>FK, FK@IRAFKD QEB @FKDRI>QB DVORP >KA QEB EFMML@>JMRP >P TBII >P QEB QE>I>JRP, EVMLQE>I>JRP >KA >JVDA>I>.
H>549.'1'2:8 #CJMU RFC RF?J?KSQ GQ RFC hypothalamus, QFMUL GL Figure 35.24. 5FC FWNMRF?J?KSQ AMLRPMJQ RFC CLBMAPGLC QWQRCK @W QCLBGLE QGEL?JQ RM RFC NGRSGR?PW EJ?LB, ? NC?-QGXCB CLBMAPGLC EJ?LB RF?R PCJC?QCQ QCTCP?J BGDDCPCLR FMPKMLCQ RF?R ?DDCAR MRFCP EJ?LBQ ?Q UCJJ ?Q MRFCP ACJJQ. 5FGQ PCJ?RGMLQFGN KC?LQ RF?R RFC FWNMRF?J?KSQ PCESJ?RCQ GKNMPR?LR @CF?TGMPQ RF?R ?PC AMLRPMJJCB @W RFCQC FMPKMLCQ. 5FC FWNMRF?J?KSQ GQ RFC @MBWsQ RFCPKMQR?RqGR K?ICQ QSPC ICW DSLARGMLQ JGIC DMMB ?LB U?RCP GLR?IC, CLCPEW CVNCLBGRSPC, ?LB @MBW RCKNCP?RSPC ?PC ICNR ?R ?NNPMNPG?RC JCTCJQ. /CSPMLQ UGRFGL RFC FWNMRF?J?KSQ ?JQM PCESJ?RC AGPA?BG?L PFWRFKQ, QMKCRGKCQ A?JJCB QJCCN AWAJCQ. L/2(/) !>89+2 5FC limbic system GQ ? AMLLCARCB QCR MD QRPSARSPCQ RF?R PCESJ?RCQ CKMRGML, ?Q UCJJ ?Q @CF?TGMPQ PCJ?RCB RM DC?P ?LB KMRGT?RGML. *R NJ?WQ ? PMJC GL KCKMPW DMPK?RGML ?LB GLAJSBCQ N?PRQ MD RFC RF?J?KSQ ?LB FWNMRF?J?KSQ ?Q UCJJ ?Q RFC FGNNMA?KNSQ. 0LC GKNMPR?LR QRPSARSPC UGRFGL RFC JGK@GA QWQRCK GQ ? RCKNMP?J JM@C QRPSARSPC A?JJCB RFC amygdala ((PCCI DMP t?JKMLBu), GJJSQRP?RCB GL Figure 35.24. 5FC RUM ?KWEB?J? ?PC GKNMPR?LR @MRF DMP RFC QCLQ?RGML MD DC?P ?LB DMP PCAMELGXGLE DC?PDSJ D?ACQ. 5FC cingulate gyrus FCJNQ PCESJ?RC CKMRGMLQ ?LB N?GL. C+7+(+11:2 5FC cerebellum (-?RGL DMP tJGRRJC @P?GLu), QFMUL GL Figure 35.21, QGRQ ?R RFC @?QC MD RFC @P?GL ML RMN MD RFC @P?GLQRCK. 5FC ACPC@CJJSK AMLRPMJQ @?J?LAC ?LB ?GBQ GL AMMPBGL?RGLE KMTCKCLR ?LB JC?PLGLE LCU KMRMP R?QIQ. B7'/389+2 5FC brainstem, GJJSQRP?RCB GL Figure 35.21, AMLLCARQ RFC PCQR MD RFC @P?GL UGRF RFC QNGL?J AMPB. *R AMLQGQRQ MD RFC KGB@P?GL, KCBSJJ? M@JMLE?R?, ?LB RFC NMLQ. .MRMP ?LB QCLQMPW LCSPMLQ CVRCLB RFPMSEF RFC @P?GLQRCK ?JJMUGLE DMP RFC PCJ?W MD QGEL?JQ @CRUCCL RFC @P?GL ?LB QNGL?J AMPB. "QACLBGLE LCSP?J N?RFU?WQ APMQQ GL RFGQ QCARGML MD RFC @P?GL ?JJMUGLE RFC JCDR FCKGQNFCPC MD RFC ACPC@PSK RM AMLRPMJ RFC PGEFR QGBC MD RFC @MBW ?LB TGAC TCPQ?. 5FC @P?GLQRCK AMMPBGL?RCQ KMRMP AMLRPMJ QGEL?JQ QCLR DPMK RFC @P?GL RM RFC @MBW. 5FC @P?GLQRCK AMLRPMJQ QCTCP?J GKNMPR?LR DSLARGMLQ MD RFC @MBW GLAJSBGLE ?JCPRLCQQ, ?PMSQ?J, @PC?RFGLE, @JMMB NPCQQSPC, BGECQRGML, FC?PR P?RC, QU?JJMUGLE, U?JIGLE, ?LB QCLQMPW ?LB KMRMP GLDMPK?RGML GLRCEP?RGML.
.G@E8C CFI;
$MLLCARGLE RM RFC @P?GLQRCK ?LB CVRCLBGLE BMUL RFC @MBW RFPMSEF RFC QNGL?J AMJSKL GQ RFC spinal cord, QFMUL GL Figure 35.21. 5FC QNGL?J AMPB GQ ? RFGAI @SLBJC MD LCPTC RGQQSC RF?R A?PPGCQ GLDMPK?RGML ?@MSR
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RFC @MBW RM RFC @P?GL ?LB DPMK RFC @P?GL RM RFC @MBW. 5FC QNGL?J AMPB GQ AMLR?GLCB UGRFGL RFC @MLCQ MD RFC TCPRC@P?RC AMJSKL @SR GQ ?@JC RM AMKKSLGA?RC QGEL?JQ RM ?LB DPMK RFC @MBW RFPMSEF GRQ AMLLCARGMLQ UGRF QNGL?J LCPTCQ (N?PR MD RFC NCPGNFCP?J LCPTMSQ QWQRCK). " APMQQ-QCARGML MD RFC QNGL?J AMPB JMMIQ JGIC ? UFGRC MT?J AMLR?GLGLE ? EP?W @SRRCPDJW-QF?NC, ?Q GJJSQRP?RCB GL Figure 35.25. .WCJGL?RCB ?VMLQ K?IC SN RFC tUFGRC K?RRCPu ?LB LCSPML ?LB EJG?J ACJJ @MBGCQ K?IC SN RFC tEP?W K?RRCP.u (P?W K?RRCP GQ ?JQM AMKNMQCB MD GLRCPLCSPMLQ, UFGAF AMLLCAR RUM LCSPMLQ C?AF JMA?RCB GL BGDDCPCLR N?PRQ MD RFC @MBW. "VMLQ ?LB ACJJ @MBGCQ GL RFC BMPQ?J (D?AGLE RFC @?AI MD RFC ?LGK?J) QNGL?J AMPB AMLTCW KMQRJW QCLQMPW GLDMPK?RGML DPMK RFC @MBW RM RFC @P?GL. "VMLQ ?LB ACJJ @MBGCQ GL RFC TCLRP?J (D?AGLE RFC DPMLR MD RFC ?LGK?J) QNGL?J AMPB NPGK?PGJW RP?LQKGR QGEL?JQ AMLRPMJJGLE KMTCKCLR DPMK RFC @P?GL RM RFC @MBW. 5FC QNGL?J AMPB ?JQM AMLRPMJQ KMRMP PCDJCVCQ. 5FCQC PCDJCVCQ ?PC OSGAI, SLAMLQAGMSQ KMTCKCLRQqJGIC ?SRMK?RGA?JJW PCKMTGLE ? F?LB DPMK ? FMR M@HCAR. 3CDJCVCQ ?PC QM D?QR @CA?SQC RFCW GLTMJTC JMA?J QWL?NRGA AMLLCARGMLQ. 'MP CV?KNJC, RFC ILCC PCDJCV RF?R ? BMARMP RCQRQ BSPGLE ? PMSRGLC NFWQGA?J GQ AMLRPMJJCB @W ? QGLEJC QWL?NQC @CRUCCL ? QCLQMPW LCSPML ?LB ? KMRMP LCSPML. 8FGJC ? PCDJCV K?W MLJW PCOSGPC RFC GLTMJTCKCLR MD MLC MP RUM QWL?NQCQ, QWL?NQCQ UGRF GLRCPLCSPMLQ GL RFC QNGL?J AMJSKL RP?LQKGR GLDMPK?RGML RM RFC @P?GL RM AMLTCW UF?R F?NNCLCB (RFC ILCC HCPICB, MP RFC F?LB U?Q FMR). *L RFC 6LGRCB 4R?RCQ, RFCPC ?PMSLB 10,000 QNGL?J AMPB GLHSPGCQ C?AF WC?P. #CA?SQC RFC QNGL?J AMPB GQ RFC GLDMPK?RGML QSNCPFGEFU?W AMLLCARGLE RFC @P?GL UGRF RFC @MBW, B?K?EC RM RFC QNGL?J AMPB A?L JC?B RM N?P?JWQGQ. 5FC CVRCLR MD RFC N?P?JWQGQ BCNCLBQ ML RFC JMA?RGML MD RFC GLHSPW ?JMLE RFC QNGL?J AMPB ?LB UFCRFCP RFC QNGL?J AMPB U?Q AMKNJCRCJW QCTCPCB. 'MP CV?KNJC, GD RFC QNGL?J AMPB GQ B?K?ECB ?R RFC JCTCJ MD RFC LCAI, GR A?L A?SQC N?P?JWQGQ DPMK RFC LCAI BMUL, UFCPC?Q B?K?EC RM RFC QNGL?J AMJSKL DSPRFCP BMUL K?W JGKGR N?P?JWQGQ RM RFC JCEQ. 4NGL?J AMPB GLHSPGCQ ?PC LMRMPGMSQJW BGDDGASJR RM RPC?R @CA?SQC QNGL?J LCPTCQ BM LMR PCECLCP?RC, ?JRFMSEF MLEMGLE PCQC?PAF QSEECQRQ RF?R QRCK ACJJ RP?LQNJ?LRQ K?W @C ?@JC RM ?AR ?Q ? @PGBEC RM PCAMLLCAR QCTCPCB LCPTCQ. 3CQC?PAFCPQ ?PC ?JQM JMMIGLE ?R U?WQ RM NPCTCLR RFC GLDJ?KK?RGML RF?R UMPQCLQ LCPTC B?K?EC ?DRCP GLHSPW. 0LC QSAF RPC?RKCLR GQ RM NSKN RFC @MBW UGRF AMJB Q?JGLC RM GLBSAC FWNMRFCPKG?. 5FGQ AMMJGLE A?L NPCTCLR QUCJJGLE ?LB MRFCP NPMACQQCQ RF?R ?PC RFMSEFR RM UMPQCL QNGL?J AMPB GLHSPGCQ.
!@>LI< 35.25 ! @OLPP-PB@QFLK LC QEB PMFK>I @LOA PELTP DO>V J>QQBO (@LKQ>FKFKD @BII ?LAFBP >KA FKQBOKBROLKP) >KA TEFQB J>QQBO (@LKQ>FKFKD >ULKP).
35.4 R /?< +<I@G?<I8C )<IMFLJ .PJK<D

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC MPE?LGX?RGML ?LB DSLARGMLQ MD RFC QWKN?RFCRGA ?LB N?P?QWKN?RFCRGA LCPTMSQ QWQRCKQ z %CQAPG@C RFC MPE?LGX?RGML ?LB DSLARGML MD RFC QCLQMPW-QMK?RGA LCPTMSQ QWQRCK 5FC NCPGNFCP?J LCPTMSQ QWQRCK (1/4) GQ RFC AMLLCARGML @CRUCCL RFC ACLRP?J LCPTMSQ QWQRCK ?LB RFC PCQR MD RFC @MBW. 5FC $/4 GQ JGIC RFC NMUCP NJ?LR MD RFC LCPTMSQ QWQRCK. *R APC?RCQ RFC QGEL?JQ RF?R AMLRPMJ RFC DSLARGMLQ MD RFC @MBW. 5FC 1/4 GQ JGIC RFC UGPCQ RF?R EM RM GLBGTGBS?J FMSQCQ. 8GRFMSR RFMQC tUGPCQ,u RFC QGEL?JQ NPMBSACB @W RFC $/4 AMSJB LMR AMLRPMJ RFC @MBW (?LB RFC $/4 UMSJB LMR @C ?@JC RM PCACGTC QCLQMPW GLDMPK?RGML DPMK RFC @MBW CGRFCP). 5FC 1/4 A?L @C @PMICL BMUL GLRM RFC autonomic nervous system, UFGAF AMLRPMJQ @MBGJW DSLARGMLQ UGRFMSR AMLQAGMSQ AMLRPMJ, ?LB RFC sensory-somatic nervous system, UFGAF RP?LQKGRQ QCLQMPW GLDMPK?RGML DPMK RFC QIGL, KSQAJCQ, ?LB QCLQMPW MPE?LQ RM RFC $/4 ?LB QCLBQ KMRMP AMKK?LBQ DPMK RFC $/4 RM RFC KSQAJCQ.
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ALKFEFD@: )<IMFLJ .PJK<D
!@>LI< 35.26 )K QEB >RQLKLJF@ KBOSLRP PVPQBJ, > MOBD>KDIFLKF@ KBROLK LC QEB #.3 PVK>MPBP TFQE > MLPQD>KDIFLKF@ KBROLK LC QEB 0.3. 4EB MLPQD>KDIFLKF@ KBROLK, FK QROK, >@QP LK > Q>ODBQ LOD>K. !RQLKLJF@ OBPMLKPBP >OB JBAF>QBA ?V QEB PVJM>QEBQF@ >KA QEB M>O>PVJM>QEBQF@ PVPQBJP, TEF@E >OB >KQ>DLKFPQF@ QL LKB >KLQEBO. 4EB PVJM>QEBQF@ PVPQBJ >@QFS>QBP QEB xCFDEQ LO CIFDEQy OBPMLKPB, TEFIB QEB M>O>PVJM>QEBQF@ PVPQBJ >@QFS>QBP QEB xOBPQ >KA AFDBPQy OBPMLKPB.
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GL RSPL, QWL?NQCQ ML RFC R?PECR MPE?L, ?Q GJJSQRP?RCB GL Figure 35.26. 5FCPC ?PC RUM BGTGQGMLQ MD RFC ?SRMLMKGA LCPTMSQ QWQRCK RF?R MDRCL F?TC MNNMQGLE CDDCARQ: RFC QWKN?RFCRGA LCPTMSQ QWQRCK ?LB RFC N?P?QWKN?RFCRGA LCPTMSQ QWQRCK. !>25'9.+9/) N+7;4:8 !>89+2 5FC sympathetic nervous system GQ PCQNMLQG@JC DMP RFC tDGEFR MP DJGEFRu PCQNMLQC RF?R MAASPQ UFCL ?L ?LGK?J CLAMSLRCPQ ? B?LECPMSQ QGRS?RGML. 0LC U?W RM PCKCK@CP RFGQ GQ RM RFGLI MD RFC QSPNPGQC ? NCPQML DCCJQ UFCL CLAMSLRCPGLE ? QL?IC (tQL?ICu ?LB tQWKN?RFCRGAu @MRF @CEGL UGRF tQu). &V?KNJCQ MD DSLARGMLQ AMLRPMJJCB @W RFC QWKN?RFCRGA LCPTMSQ QWQRCK GLAJSBC ?L ?AACJCP?RCB FC?PR P?RC ?LB GLFG@GRCB BGECQRGML. 5FCQC DSLARGMLQ FCJN NPCN?PC ?L MPE?LGQKsQ @MBW DMP RFC NFWQGA?J QRP?GL PCOSGPCB RM CQA?NC ? NMRCLRG?JJW B?LECPMSQ QGRS?RGML MP RM DCLB MDD ? NPCB?RMP.
!@>LI< 35.27 4EB PVJM>QEBQF@ >KA M>O>PVJM>QEBQF@ KBOSLRP PVPQBJP LCQBK E>SB LMMLPFKD BCCB@QP LK Q>ODBQ LOD>KP.
.MQR NPCE?LEJGMLGA LCSPMLQ GL RFC QWKN?RFCRGA LCPTMSQ QWQRCK MPGEGL?RC GL RFC QNGL?J AMPB, ?Q GJJSQRP?RCB GL Figure 35.27. 5FC ?VMLQ MD RFCQC LCSPMLQ PCJC?QC acetylcholine ML NMQRE?LEJGMLGA LCSPMLQ UGRFGL QWKN?RFCRGA E?LEJG? (RFC QWKN?RFCRGA E?LEJG? DMPK ? AF?GL RF?R CVRCLBQ ?JMLEQGBC RFC QNGL?J AMPB). 5FC ?ACRWJAFMJGLC ?ARGT?RCQ RFC NMQRE?LEJGMLGA LCSPMLQ. 1MQRE?LEJGMLGA LCSPMLQ RFCL PCJC?QC norepinephrine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q?JJMUGLE RFC ?LGK?J RM PCKCK@CP RFC B?LECPMSQ QGRS?RGML ?LB ?TMGB GR GL RFC DSRSPC.
#(!04%2 35 X 4(% .%26/53 3934%-
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P'7'8>25'9.+9/) N+7;4:8 !>89+2 8FGJC RFC QWKN?RFCRGA LCPTMSQ QWQRCK GQ ?ARGT?RCB GL QRPCQQDSJ QGRS?RGMLQ, RFC parasympathetic nervous system ?JJMUQ ?L ?LGK?J RM tPCQR ?LB BGECQR.u 0LC U?W RM PCKCK@CP RFGQ GQ RM RFGLI RF?R BSPGLE ? PCQRDSJ QGRS?RGML JGIC ? NGALGA, RFC N?P?QWKN?RFCRGA LCPTMSQ QWQRCK GQ GL AMLRPMJ (tNGALGAu ?LB tN?P?QWKN?RFCRGAu @MRF QR?PR UGRF tNu). 1?P?QWKN?RFCRGA NPCE?LEJGMLGA LCSPMLQ F?TC ACJJ @MBGCQ JMA?RCB GL RFC @P?GLQRCK ?LB GL RFC Q?AP?J (RMU?PB RFC @MRRMK) QNGL?J AMPB, ?Q QFMUL GL Figure 35.27. 5FC ?VMLQ MD RFC NPCE?LEJGMLGA LCSPMLQ PCJC?QC ?ACRWJAFMJGLC ML RFC NMQRE?LEJGMLGA LCSPMLQ, UFGAF ?PC ECLCP?JJW JMA?RCB TCPW LC?P RFC R?PECR MPE?LQ. .MQR NMQRE?LEJGMLGA LCSPMLQ PCJC?QC ?ACRWJAFMJGLC MLRM R?PECR MPE?LQ, ?JRFMSEF QMKC PCJC?QC LGRPGA MVGBC. 5FC N?P?QWKN?RFCRGA LCPTMSQ QWQRCK PCQCRQ MPE?L DSLARGML ?DRCP RFC QWKN?RFCRGA LCPTMSQ QWQRCK GQ ?ARGT?RCB (RFC AMKKML ?BPCL?JGLC BSKN WMS DCCJ ?DRCP ? rDGEFR-MP-DJGEFRs CTCLR). &DDCARQ MD ?ACRWJAFMJGLC PCJC?QC ML R?PECR MPE?LQ GLAJSBC QJMUGLE MD FC?PR P?RC, JMUCPCB @JMMB NPCQQSPC, ?LB QRGKSJ?RGML MD BGECQRGML.
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qMLC CLBGLE MD RFC LCSPML GQ ?R RFC MPE?L ?LB RFC MRFCP BGPCARJW AMLR?ARQ ? $/4 LCSPML. "ACRWJAFMJGLC GQ RFC K?GL LCSPMRP?LQKGRRCP PCJC?QCB ?R RFCQC QWL?NQCQ. )SK?LQ F?TC 12 cranial nerves, LCPTCQ RF?R CKCPEC DPMK MP CLRCP RFC QISJJ (AP?LGSK), ?Q MNNMQCB RM RFC QNGL?J LCPTCQ, UFGAF CKCPEC DPMK RFC TCPRC@P?J AMJSKL. &?AF AP?LG?J LCPTC GQ ?AAMPBCB ? L?KC, UFGAF ?PC BCR?GJCB GL Figure 35.28. 4MKC AP?LG?J LCPTCQ RP?LQKGR MLJW QCLQMPW GLDMPK?RGML. 'MP CV?KNJC, RFC MJD?ARMPW LCPTC RP?LQKGRQ GLDMPK?RGML ?@MSR QKCJJQ DPMK RFC LMQC RM RFC @P?GLQRCK. 0RFCP AP?LG?J LCPTCQ RP?LQKGR ?JKMQR QMJCJW KMRMP GLDMPK?RGML. 'MP CV?KNJC, RFC MASJMKMRMP LCPTC AMLRPMJQ RFC MNCLGLE ?LB AJMQGLE MD RFC CWCJGB ?LB QMKC CWC KMTCKCLRQ. 0RFCP AP?LG?J LCPTCQ AMLR?GL ? KGV MD QCLQMPW ?LB KMRMP DG@CPQ. 'MP CV?KNJC, RFC EJMQQMNF?PWLEC?J LCPTC F?Q ? PMJC GL @MRF R?QRC (QCLQMPW) ?LB QU?JJMUGLE (KMRMP).
!@>LI< 35.28 4EB ERJ>K ?O>FK @LKQ>FKP 12 @O>KF>I KBOSBP QE>Q OB@BFSB PBKPLOV FKMRQ >KA @LKQOLI JLQLO LRQMRQ CLO QEB EB>A >KA KB@H.
Spinal nerves RP?LQKGR QCLQMPW ?LB KMRMP GLDMPK?RGML @CRUCCL RFC QNGL?J AMPB ?LB RFC PCQR MD RFC @MBW. &?AF MD RFC 31 QNGL?J LCPTCQ (GL FSK?LQ) AMLR?GLQ @MRF QCLQMPW ?LB KMRMP ?VMLQ. 5FC QCLQMPW LCSPML ACJJ @MBGCQ ?PC EPMSNCB GL QRPSARSPCQ A?JJCB BMPQ?J PMMR E?LEJG? ?LB ?PC QFMUL GL Figure 35.29.
1018
#(!04%2 35 X 4(% .%26/53 3934%-
&?AF QCLQMPW LCSPML F?Q MLC NPMHCARGMLqUGRF ? QCLQMPW PCACNRMP CLBGLE GL QIGL, KSQAJC, MP QCLQMPW MPE?LQq?LB ?LMRFCP RF?R QWL?NQCQ UGRF ? LCSPML GL RFC BMPQ?J QNGL?J AMPB. .MRMP LCSPMLQ F?TC ACJJ @MBGCQ GL RFC TCLRP?J EP?W K?RRCP MD RFC QNGL?J AMPB RF?R NPMHCAR RM KSQAJC RFPMSEF RFC TCLRP?J PMMR. 5FCQC LCSPMLQ ?PC SQS?JJW QRGKSJ?RCB @W GLRCPLCSPMLQ UGRFGL RFC QNGL?J AMPB @SR ?PC QMKCRGKCQ BGPCARJW QRGKSJ?RCB @W QCLQMPW LCSPMLQ.
!@>LI< 35.29 3MFK>I KBOSBP @LKQ>FK ?LQE PBKPLOV >KA JLQLO >ULKP. 4EB PLJ>P LC PBKPLOV KBROLKP >OB IL@>QBA FK ALOP>I OLLQ D>KDIF>. 4EB PLJ>P LC JLQLO KBROLKP >OB CLRKA FK QEB SBKQO>I MLOQFLK LC QEB DO>V J>QQBO LC QEB PMFK>I @LOA.
35.5 R )<IMFLJ .PJK<D D@JFI;<IJ

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC QWKNRMKQ, NMRCLRG?J A?SQCQ, ?LB RPC?RKCLR MD QCTCP?J CV?KNJCQ MD LCPTMSQ QWQRCK BGQMPBCPQ " LCPTMSQ QWQRCK RF?R DSLARGMLQ AMPPCARJW GQ ? D?LR?QRGA?JJW AMKNJCV, UCJJ-MGJCB K?AFGLCqQWL?NQCQ DGPC ?NNPMNPG?RCJW, KSQAJCQ KMTC UFCL LCCBCB, KCKMPGCQ ?PC DMPKCB ?LB QRMPCB, ?LB CKMRGMLQ ?PC UCJJ PCESJ?RCB. 6LDMPRSL?RCJW, C?AF WC?P KGJJGMLQ MD NCMNJC GL RFC 6LGRCB 4R?RCQ BC?J UGRF QMKC QMPR MD LCPTMSQ QWQRCK BGQMPBCP. 8FGJC QAGCLRGQRQ F?TC BGQAMTCPCB NMRCLRG?J A?SQCQ MD K?LW MD RFCQC BGQC?QCQ, ?LB TG?@JC RPC?RKCLRQ DMP QMKC, MLEMGLE PCQC?PAF QCCIQ RM DGLB U?WQ RM @CRRCP NPCTCLR ?LB RPC?R ?JJ MD RFCQC BGQMPBCPQ.
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Neurodegenerative disorders ?PC GJJLCQQCQ AF?P?ARCPGXCB @W ? JMQQ MD LCPTMSQ QWQRCK DSLARGMLGLE RF?R ?PC SQS?JJW A?SQCB @W LCSPML?J BC?RF. 5FCQC BGQC?QCQ ECLCP?JJW UMPQCL MTCP RGKC ?Q KMPC ?LB KMPC LCSPMLQ BGC. 5FC QWKNRMKQ MD ? N?PRGASJ?P LCSPMBCECLCP?RGTC BGQC?QC ?PC PCJ?RCB RM UFCPC GL RFC LCPTMSQ QWQRCK RFC BC?RF MD LCSPMLQ MAASPQ. 4NGLMACPC@CJJ?P ?R?VG?, DMP CV?KNJC, JC?BQ RM LCSPML?J BC?RF GL RFC ACPC@CJJSK. 5FC BC?RF MD RFCQC LCSPMLQ A?SQCQ NPM@JCKQ GL @?J?LAC ?LB U?JIGLE. /CSPMBCECLCP?RGTC BGQMPBCPQ GLAJSBC )SLRGLERMLsQ BGQC?QC, ?KWMRPMNFGA J?RCP?J QAJCPMQGQ, "JXFCGKCPsQ BGQC?QC ?LB MRFCP RWNCQ MD BCKCLRG? BGQMPBCPQ, ?LB 1?PIGLQMLsQ BGQC?QC. )CPC, "JXFCGKCPsQ ?LB 1?PIGLQMLsQ BGQC?QC UGJJ @C BGQASQQCB GL KMPC BCNRF. A1?.+/2+7B8 D/8+'8+ Alzheimers disease GQ RFC KMQR AMKKML A?SQC MD BCKCLRG? GL RFC CJBCPJW. *L 2012, ?L CQRGK?RCB 5.4 KGJJGML "KCPGA?LQ QSDDCPCB DPMK "JXFCGKCPsQ BGQC?QC, ?LB N?WKCLRQ DMP RFCGP A?PC ?PC CQRGK?RCB ?R $200 @GJJGML. 3MSEFJW MLC GL CTCPW CGEFR NCMNJC ?EC 65 MP MJBCP F?Q RFC BGQC?QC. %SC RM RFC ?EGLE MD RFC @?@W@MMKCP ECLCP?RGML, RFCPC ?PC NPMHCARCB RM @C ?Q K?LW ?Q 13 KGJJGML "JXFCGKCPsQ N?RGCLRQ GL RFC 6LGRCB 4R?RCQ GL RFC WC?P 2050. 4WKNRMKQ MD "JXFCGKCPsQ BGQC?QC GLAJSBC BGQPSNRGTC KCKMPW JMQQ, AMLDSQGML ?@MSR RGKC MP NJ?AC, BGDDGASJRW NJ?LLGLE MP CVCASRGLE R?QIQ, NMMP HSBEKCLR, ?LB NCPQML?JGRW AF?LECQ. 1PM@JCKQ QKCJJGLE ACPR?GL QACLRQ A?L ?JQM @C GLBGA?RGTC MD "JXFCGKCPsQ BGQC?QC ?LB K?W QCPTC ?Q ?L C?PJW U?PLGLE QGEL. .?LW MD RFCQC QWKNRMKQ ?PC ?JQM AMKKML GL NCMNJC UFM ?PC ?EGLE LMPK?JJW, QM GR GQ RFC QCTCPGRW ?LB JMLECTGRW MD RFC QWKNRMKQ RF?R BCRCPKGLC UFCRFCP ? NCPQML GQ QSDDCPGLE DPMK "JXFCGKCPsQ.
#(!04%2 35 X 4(% .%26/53 3934%-
1019
"JXFCGKCPsQ BGQC?QC U?Q L?KCB DMP "JMGQ "JXFCGKCP, ? (CPK?L NQWAFG?RPGQR UFM NS@JGQFCB ? PCNMPR GL 1911 ?@MSR ? UMK?L UFM QFMUCB QCTCPC BCKCLRG? QWKNRMKQ. "JMLE UGRF FGQ AMJJC?ESCQ, FC CV?KGLCB RFC UMK?LsQ @P?GL DMJJMUGLE FCP BC?RF ?LB PCNMPRCB RFC NPCQCLAC MD ?@LMPK?J AJSKNQ, UFGAF ?PC LMU A?JJCB ?KWJMGB NJ?OSCQ, ?JMLE UGRF R?LEJCB @P?GL DG@CPQ A?JJCB LCSPMDG@PGJJ?PW R?LEJCQ. "KWJMGB NJ?OSCQ, LCSPMDG@PGJJ?PW R?LEJCQ, ?LB ?L MTCP?JJ QFPGLIGLE MD @P?GL TMJSKC ?PC AMKKMLJW QCCL GL RFC @P?GLQ MD "JXFCGKCPsQ N?RGCLRQ. -MQQ MD LCSPMLQ GL RFC FGNNMA?KNSQ GQ CQNCAG?JJW QCTCPC GL ?BT?LACB "JXFCGKCPsQ N?RGCLRQ. Figure 35.30 AMKN?PCQ ? LMPK?J @P?GL RM RFC @P?GL MD ?L "JXFCGKCPsQ N?RGCLR. .?LW PCQC?PAF EPMSNQ ?PC CV?KGLGLE RFC A?SQCQ MD RFCQC F?JJK?PIQ MD RFC BGQC?QC. 0LC DMPK MD RFC BGQC?QC GQ SQS?JJW A?SQCB @W KSR?RGMLQ GL MLC MD RFPCC ILMUL ECLCQ. 5FGQ P?PC DMPK MD C?PJW MLQCR "JXFCGKCPsQ BGQC?QC ?DDCARQ DCUCP RF?L DGTC NCPACLR MD N?RGCLRQ UGRF RFC BGQC?QC ?LB A?SQCQ BCKCLRG? @CEGLLGLE @CRUCCL RFC ?ECQ MD 30 ?LB 60. 5FC KMPC NPCT?JCLR, J?RC-MLQCR DMPK MD RFC BGQC?QC JGICJW ?JQM F?Q ? ECLCRGA AMKNMLCLR. 0LC N?PRGASJ?P ECLC, ?NMJGNMNPMRCGL & ("10&) F?Q ? T?PG?LR (&4) RF?R GLAPC?QCQ ? A?PPGCPsQ JGICJGFMMB MD ECRRGLE RFC BGQC?QC. .?LW MRFCP ECLCQ F?TC @CCL GBCLRGDGCB RF?R KGEFR @C GLTMJTCB GL RFC N?RFMJMEW.
7GQGR this website (http://openstaxcollege.org/l/alzheimers) DMP TGBCM JGLIQ BGQASQQGLE ECLCRGAQ ?LB "JXFCGKCPsQ BGQC?QC. 6LDMPRSL?RCJW, RFCPC GQ LM ASPC DMP "JXFCGKCPsQ BGQC?QC. $SPPCLR RPC?RKCLRQ DMASQ ML K?L?EGLE RFC QWKNRMKQ MD RFC BGQC?QC. #CA?SQC BCAPC?QC GL RFC ?ARGTGRW MD AFMJGLCPEGA LCSPMLQ (LCSPMLQ RF?R SQC RFC LCSPMRP?LQKGRRCP ?ACRWJAFMJGLC) GQ AMKKML GL "JXFCGKCPsQ BGQC?QC, QCTCP?J BPSEQ SQCB RM RPC?R RFC BGQC?QC UMPI @W GLAPC?QGLE ?ACRWJAFMJGLC LCSPMRP?LQKGQQGML, MDRCL @W GLFG@GRGLE RFC CLXWKC RF?R @PC?IQ BMUL ?ACRWJAFMJGLC GL RFC QWL?NRGA AJCDR. 0RFCP AJGLGA?J GLRCPTCLRGMLQ DMASQ ML @CF?TGMP?J RFCP?NGCQ JGIC NQWAFMRFCP?NW, QCLQMPW RFCP?NW, ?LB AMELGRGTC CVCPAGQCQ. 4GLAC "JXFCGKCPsQ BGQC?QC ?NNC?PQ RM FGH?AI RFC LMPK?J ?EGLE NPMACQQ, PCQC?PAF GLRM NPCTCLRGML GQ NPCT?JCLR. 4KMIGLE, M@CQGRW, ?LB A?PBGMT?QASJ?P NPM@JCKQ K?W @C PGQI D?ARMPQ DMP RFC BGQC?QC, QM RPC?RKCLRQ DMP RFMQC K?W ?JQM FCJN RM NPCTCLR "JXFCGKCPsQ BGQC?QC. 4MKC QRSBGCQ F?TC QFMUL RF?R NCMNJC UFM PCK?GL GLRCJJCARS?JJW ?ARGTC @W NJ?WGLE E?KCQ, PC?BGLE, NJ?WGLE KSQGA?J GLQRPSKCLRQ, ?LB @CGLE QMAG?JJW ?ARGTC GL J?RCP JGDC F?TC ? PCBSACB PGQI MD BCTCJMNGLE RFC BGQC?QC.
!@>LI< 35.30 #LJM>OBA QL > KLOJ>I ?O>FK (IBCQ), QEB ?O>FK COLJ > M>QFBKQ TFQE !IWEBFJBOwP AFPB>PB (OFDEQ) PELTP > AO>J>QF@ KBROLABDBKBO>QFLK, M>OQF@RI>OIV TFQEFK QEB SBKQOF@IBP >KA EFMML@>JMRP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V x'>OO>KALy/7FHFJBAF> #LJJLKP ?>PBA LK LOFDFK>I FJ>DBP ?V !$%!2: "!IWEBFJBO'P $FPB>PB %AR@>QFLK >KA 2BCBOO>I #BKQBO, > PBOSF@B LC QEB .>QFLK>I )KPQFQRQB LK !DFKDy)
1020
#(!04%2 35 X 4(% .%26/53 3934%-
P'70/3843B8 D/8+'8+ -GIC "JXFCGKCPsQ BGQC?QC, Parkinsons disease GQ ? LCSPMBCECLCP?RGTC BGQC?QC. *R U?Q DGPQR AF?P?ARCPGXCB @W +?KCQ 1?PIGLQML GL 1817. &?AF WC?P, 50,000-60,000 NCMNJC GL RFC 6LGRCB 4R?RCQ ?PC BG?ELMQCB UGRF RFC BGQC?QC. 1?PIGLQMLsQ BGQC?QC A?SQCQ RFC JMQQ MD BMN?KGLC LCSPMLQ GL RFC QS@QR?LRG? LGEP?, ? KGB@P?GL QRPSARSPC RF?R PCESJ?RCQ KMTCKCLR. -MQQ MD RFCQC LCSPMLQ A?SQCQ K?LW QWKNRMKQ GLAJSBGLE RPCKMP (QF?IGLE MD DGLECPQ MP ? JGK@), QJMUCB KMTCKCLR, QNCCAF AF?LECQ, @?J?LAC ?LB NMQRSPC NPM@JCKQ, ?LB PGEGB KSQAJCQ. 5FC AMK@GL?RGML MD RFCQC QWKNRMKQ MDRCL A?SQCQ ? AF?P?ARCPGQRGA QJMU FSLAFCB QFSDDJGLE U?JI, GJJSQRP?RCB GL Figure 35.31. 1?RGCLRQ UGRF 1?PIGLQMLsQ BGQC?QC A?L ?JQM CVFG@GR NQWAFMJMEGA?J QWKNRMKQ, QSAF ?Q BCKCLRG? MP CKMRGML?J NPM@JCKQ. "JRFMSEF QMKC N?RGCLRQ F?TC ? DMPK MD RFC BGQC?QC ILMUL RM @C A?SQCB @W ? QGLEJC KSR?RGML, DMP KMQR N?RGCLRQ RFC CV?AR A?SQCQ MD 1?PIGLQMLsQ BGQC?QC PCK?GL SLILMUL: RFC BGQC?QC JGICJW PCQSJRQ DPMK ? AMK@GL?RGML MD ECLCRGA ?LB CLTGPMLKCLR?J D?ARMPQ (QGKGJ?P RM "JXFCGKCPsQ BGQC?QC). 1MQR-KMPRCK ?L?JWQGQ MD @P?GLQ DPMK 1?PIGLQMLsQ N?RGCLRQ QFMUQ RFC NPCQCLAC MD -CUW @MBGCQq?@LMPK?J NPMRCGL AJSKNQqGL BMN?KGLCPEGA LCSPMLQ. 5FC NPCT?JCLAC MD RFCQC -CUW @MBGCQ MDRCL AMPPCJ?RCQ UGRF RFC QCTCPGRW MD RFC BGQC?QC. 5FCPC GQ LM ASPC DMP 1?PIGLQMLsQ BGQC?QC, ?LB RPC?RKCLR GQ DMASQCB ML C?QGLE QWKNRMKQ. 0LC MD RFC KMQR AMKKMLJW NPCQAPG@CB BPSEQ DMP 1?PIGLQMLsQ GQ --%01", UFGAF GQ ? AFCKGA?J RF?R GQ AMLTCPRCB GLRM BMN?KGLC @W LCSPMLQ GL RFC @P?GL. 5FGQ AMLTCPQGML GLAPC?QCQ RFC MTCP?JJ JCTCJ MD BMN?KGLC LCSPMRP?LQKGQQGML ?LB A?L FCJN AMKNCLQ?RC DMP RFC JMQQ MD BMN?KGLCPEGA LCSPMLQ GL RFC QS@QR?LRG? LGEP?. 0RFCP BPSEQ UMPI @W GLFG@GRGLE RFC CLXWKC RF?R @PC?IQ BMUL BMN?KGLC.
!@>LI< 35.31 0>OHFKPLKwP M>QFBKQP LCQBK E>SB > @E>O>@QBOFPQF@ ERK@EBA T>IH.
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/CSPMBCTCJMNKCLR?J BGQMPBCPQ MAASP UFCL RFC BCTCJMNKCLR MD RFC LCPTMSQ QWQRCK GQ BGQRSP@CB. 5FCPC ?PC QCTCP?J BGDDCPCLR AJ?QQCQ MD LCSPMBCTCJMNKCLR?J BGQMPBCPQ. 4MKC, JGIC %MUL 4WLBPMKC, A?SQC GLRCJJCARS?J BCDGAGRQ. 0RFCPQ QNCAGDGA?JJW ?DDCAR AMKKSLGA?RGML, JC?PLGLE, MP RFC KMRMP QWQRCK. 4MKC BGQMPBCPQ JGIC ?SRGQK QNCARPSK BGQMPBCP ?LB ?RRCLRGML BCDGAGR/FWNCP?ARGTGRW BGQMPBCP F?TC AMKNJCV QWKNRMKQ. A:9/82 Autism spectrum disorder (ASD) GQ ? LCSPMBCTCJMNKCLR?J BGQMPBCP. *RQ QCTCPGRW BGDDCPQ DPMK NCPQML RM NCPQML. &QRGK?RCQ DMP RFC NPCT?JCLAC MD RFC BGQMPBCP F?TC AF?LECB P?NGBJW GL RFC N?QR DCU BCA?BCQ. $SPPCLR CQRGK?RCQ QSEECQR RF?R MLC GL 88 AFGJBPCL UGJJ BCTCJMN RFC BGQMPBCP. "4% GQ DMSP RGKCQ KMPC NPCT?JCLR GL K?JCQ RF?L DCK?JCQ.
#(!04%2 35 X 4(% .%26/53 3934%-
1021
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#(!04%2 35 X 4(% .%26/53 3934%-
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s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sQ AMLQAGMSQ PCQNMLQC RM SLNJC?Q?LR CVNCPGCLACQ :FIGLJ :8CCFJLD RFGAI DG@CP @SLBJC RF?R AMLLCARQ RFC ACPC@P?J FCKGQNFCPCQ :I8E@8C E<IM< QCLQMPW ?LB/MP KMRMP LCPTC RF?R CK?L?RCQ DPMK RFC @P?GL ;<E;I@K< QRPSARSPC RF?R CVRCLBQ ?U?W DPMK RFC ACJJ @MBW RM PCACGTC KCQQ?ECQ DPMK MRFCP LCSPMLQ ;<GFC8I@Q8K@FE AF?LEC GL RFC KCK@P?LC NMRCLRG?J RM ? JCQQ LCE?RGTC T?JSC ;LI8 D8K<I RMSEF MSRCPKMQR J?WCP RF?R AMTCPQ RFC ACLRP?J LCPTMSQ QWQRCK
1026
#(!04%2 35 X 4(% .%26/53 3934%-
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tHSKNGLEu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tT?JJCWQu GL RFC AMPRCV JLDD8K@FE NPMACQQ MD KSJRGNJC NPCQWL?NRGA GLNSRQ APC?RGLE &141Q ?PMSLB RFC Q?KC RGKC DMP RFC NMQRQWL?NRGA LCSPML RM @C QSDDGAGCLRJW BCNMJ?PGXCB RM DGPC ?L ?ARGML NMRCLRG?J JPDG8K?<K@: E<IMFLJ JPJK<D BGTGQGML MD ?SRMLMKGA LCPTMSQ QWQRCK ?ARGT?RCB BSPGLE QRPCQQDSJ tDGEFR MP DJGEFRu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
C#A+/ - .0((A-4
35.1 )<LIFEJ 8E; "C@8C C<CCJ
5FC LCPTMSQ QWQRCK GQ K?BC SN MD LCSPMLQ ?LB EJG?. /CSPMLQ ?PC QNCAG?JGXCB ACJJQ RF?R ?PC A?N?@JC MD QCLBGLE CJCARPGA?J ?Q UCJJ ?Q AFCKGA?J QGEL?JQ. .MQR LCSPMLQ AMLR?GL BCLBPGRCQ, UFGAF PCACGTC RFCQC QGEL?JQ, ?LB ?VMLQ RF?R QCLB QGEL?JQ RM MRFCP LCSPMLQ MP RGQQSCQ. 5FCPC ?PC DMSP K?GL RWNCQ MD LCSPMLQ: SLGNMJ?P, @GNMJ?P, KSJRGNMJ?P, ?LB NQCSBMSLGNMJ?P LCSPMLQ. (JG? ?PC LML-LCSPML?J ACJJQ GL RFC LCPTMSQ QWQRCK RF?R QSNNMPR LCSPML?J BCTCJMNKCLR ?LB QGEL?JGLE. 5FCPC ?PC QCTCP?J RWNCQ MD EJG? RF?R QCPTC BGDDCPCLR DSLARGMLQ.
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35.2 #FN )<LIFEJ CFDDLE@:8K<

/CSPMLQ F?TC AF?PECB KCK@P?LCQ @CA?SQC RFCPC ?PC BGDDCPCLR AMLACLRP?RGMLQ MD GMLQ GLQGBC ?LB MSRQGBC MD RFC ACJJ. 7MJR?EC-E?RCB GML AF?LLCJQ AMLRPMJ RFC KMTCKCLR MD GMLQ GLRM ?LB MSR MD ? LCSPML. 8FCL ? LCSPML?J KCK@P?LC GQ BCNMJ?PGXCB RM ?R JC?QR RFC RFPCQFMJB MD CVAGR?RGML, ?L ?ARGML NMRCLRG?J GQ DGPCB. 5FC ?ARGML NMRCLRG?J GQ RFCL NPMN?E?RCB ?JMLE ? KWCJGL?RCB ?VML RM RFC ?VML RCPKGL?JQ. *L ? AFCKGA?J QWL?NQC, RFC ?ARGML NMRCLRG?J A?SQCQ PCJC?QC MD LCSPMRP?LQKGRRCP KMJCASJCQ GLRM RFC QWL?NRGA AJCDR. 5FPMSEF @GLBGLE RM NMQRQWL?NRGA PCACNRMPQ, RFC LCSPMRP?LQKGRRCP A?L A?SQC CVAGR?RMPW MP GLFG@GRMPW NMQRQWL?NRGA NMRCLRG?JQ @W BCNMJ?PGXGLE MP FWNCPNMJ?PGXGLE, PCQNCARGTCJW, RFC NMQRQWL?NRGA KCK@P?LC. *L CJCARPGA?J QWL?NQCQ, RFC ?ARGML NMRCLRG?J GQ BGPCARJW AMKKSLGA?RCB RM RFC NMQRQWL?NRGA ACJJ RFPMSEF E?N HSLARGMLQqJ?PEC AF?LLCJ NPMRCGLQ RF?R AMLLCAR RFC NPC-?LB NMQRQWL?NRGA KCK@P?LCQ. 4WL?NQCQ ?PC LMR QR?RGA QRPSARSPCQ ?LB A?L @C QRPCLERFCLCB ?LB UC?ICLCB. 5UM KCAF?LGQKQ MD QWL?NRGA NJ?QRGAGRW ?PC JMLE-RCPK NMRCLRG?RGML ?LB JMLE-RCPK BCNPCQQGML.
35.3 /?< C<EKI8C )<IMFLJ .PJK<D

5FC TCPRC@P?RC ACLRP?J LCPTMSQ QWQRCK AMLR?GLQ RFC @P?GL ?LB RFC QNGL?J AMPB, UFGAF ?PC AMTCPCB ?LB NPMRCARCB @W RFPCC KCLGLECQ. 5FC @P?GL AMLR?GLQ QRPSARSP?JJW ?LB DSLARGML?JJW BCDGLCB PCEGMLQ. *L K?KK?JQ, RFCQC GLAJSBC RFC AMPRCV (UFGAF A?L @C @PMICL BMUL GLRM DMSP NPGK?PW DSLARGML?J JM@CQ: DPMLR?J, RCKNMP?J, MAAGNGR?J, ?LB N?PGCR?J), @?Q?J E?LEJG?, RF?J?KSQ, FWNMRF?J?KSQ, JGK@GA QWQRCK, ACPC@CJJSK, ?LB @P?GLQRCKq?JRFMSEF QRPSARSPCQ GL QMKC MD RFCQC BCQGEL?RGMLQ MTCPJ?N. 8FGJC DSLARGMLQ K?W @C NPGK?PGJW JMA?JGXCB RM MLC QRPSARSPC GL RFC @P?GL, KMQR AMKNJCV DSLARGMLQ, JGIC J?LES?EC ?LB QJCCN, GLTMJTC LCSPMLQ GL KSJRGNJC @P?GL PCEGMLQ. 5FC QNGL?J AMPB GQ RFC GLDMPK?RGML QSNCPFGEFU?W RF?R AMLLCARQ RFC @P?GL UGRF RFC PCQR MD RFC @MBW RFPMSEF GRQ AMLLCARGMLQ UGRF NCPGNFCP?J LCPTCQ. *R RP?LQKGRQ QCLQMPW ?LB KMRMP GLNSR ?LB ?JQM AMLRPMJQ KMRMP PCDJCVCQ.
35.4 /?< +<I@G?<I8C )<IMFLJ .PJK<D

5FC NCPGNFCP?J LCPTMSQ QWQRCK AMLR?GLQ @MRF RFC ?SRMLMKGA ?LB QCLQMPW-QMK?RGA LCPTMSQ QWQRCKQ. 5FC ?SRMLMKGA LCPTMSQ QWQRCK NPMTGBCQ SLAMLQAGMSQ AMLRPMJ MTCP TGQACP?J DSLARGMLQ ?LB F?Q RUM BGTGQGMLQ: RFC QWKN?RFCRGA ?LB N?P?QWKN?RFCRGA LCPTMSQ QWQRCKQ. 5FC QWKN?RFCRGA LCPTMSQ QWQRCK GQ ?ARGT?RCB GL QRPCQQDSJ QGRS?RGMLQ RM NPCN?PC RFC ?LGK?J DMP ? tDGEFR MP DJGEFRu PCQNMLQC. 5FC N?P?QWKN?RFCRGA LCPTMSQ QWQRCK GQ ?ARGTC BSPGLE PCQRDSJ NCPGMBQ. 5FC QCLQMPW-QMK?RGA LCPTMSQ QWQRCK GQ K?BC MD AP?LG?J ?LB QNGL?J LCPTCQ RF?R RP?LQKGR QCLQMPW GLDMPK?RGML DPMK QIGL ?LB KSQAJC RM RFC $/4 ?LB KMRMP AMKK?LBQ DPMK RFC $/4 RM RFC KSQAJCQ.
35.5 )<IMFLJ .PJK<D D@JFI;<IJ

4MKC ECLCP?J RFCKCQ CKCPEC DPMK RFC Q?KNJGLE MD LCPTMSQ QWQRCK BGQMPBCPQ NPCQCLRCB ?@MTC. 5FC A?SQCQ DMP KMQR BGQMPBCPQ ?PC LMR DSJJW SLBCPQRMMBq?R JC?QR LMR DMP ?JJ N?RGCLRQq?LB JGICJW GLTMJTC ? AMK@GL?RGML MD L?RSPC (ECLCRGA KSR?RGMLQ RF?R @CAMKC PGQI D?ARMPQ) ?LB LSPRSPC (CKMRGML?J RP?SK?, QRPCQQ, F?X?PBMSQ AFCKGA?J CVNMQSPC). #CA?SQC RFC A?SQCQ F?TC WCR RM @C DSJJW BCRCPKGLCB, RPC?RKCLR MNRGMLQ ?PC MDRCL J?AIGLE ?LB MLJW ?BBPCQQ QWKNRMKQ.
A-/ C*)) C/$*) ,0 ./$*).

1. Figure 35.3 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ GQ D?JQC >. 5FC QMK? GQ RFC ACJJ @MBW MD ? LCPTC ACJJ. ?. .WCJGL QFC?RF NPMTGBCQ ?L GLQSJ?RGLE J?WCP RM RFC BCLBPGRCQ. @. "VMLQ A?PPW RFC QGEL?J DPMK RFC QMK? RM RFC R?PECR. A. %CLBPGRCQ A?PPW RFC QGEL?J RM RFC QMK?. 2. Figure 35.11 1MR?QQGSK AF?LLCJ @JMAICPQ, QSAF ?Q ?KGMB?PMLC ?LB NPMA?GL?KGBC, UFGAF ?PC SQCB RM RPC?R ?@LMPK?J CJCARPGA?J ?ARGTGRW GL RFC FC?PR, A?JJCB A?PBG?A BWQPFWRFKG?, GKNCBC RFC KMTCKCLR MD ,+ RFPMSEF TMJR?EC-E?RCB ,+ AF?LLCJQ. 8FGAF N?PR MD RFC ?ARGML NMRCLRG?J UMSJB WMS CVNCAR NMR?QQGSK AF?LLCJQ RM ?DDCAR 3. Figure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
#(!04%2 35 X 4(% .%26/53 3934%-
1029
- 1$ 2 ,0 ./$*).
4. /CSPMLQ AMLR?GL >>>>>>>>, UFGAF A?L PCACGTC QGEL?JQ DPMK MRFCP LCSPMLQ. >. ?VMLQ ?. KGRMAFMLBPG? @. BCLBPGRCQ A. (MJEG @MBGCQ 5. "(L) >>>>>>>> LCSPML F?Q MLC ?VML ?LB MLC BCLBPGRC CVRCLBGLE BGPCARJW DPMK RFC ACJJ @MBW. >. ?. @. A. SLGNMJ?P @GNMJ?P KSJRGNMJ?P NQCSBMSLGNMJ?P >. ?. @. A. JGK@GA QWQRCK AMPNSQ A?JJMQSK ACPC@CJJSK NGRSGR?PW
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sQ BGQC?QC GQ ? A?SQCB @W RFC BCECLCP?RGML MD LCSPMLQ RF?R PCJC?QC >>>>>>>>. >. ?. @. A. QCPMRMLGL BMN?KGLC EJSR?K?RC LMPCNGLCNFPGLC
6. (JG? RF?R NPMTGBC KWCJGL DMP LCSPMLQ GL RFC @P?GL ?PC A?JJCB >>>>>>>>. >. 4AFU?LL ACJJQ ?. MJGEMBCLBPMAWRCQ @. KGAPMEJG? A. ?QRPMAWRCQ 7. 'MP ? LCSPML RM DGPC ?L ?ARGML NMRCLRG?J, GRQ KCK@P?LC KSQR PC?AF >>>>>>>>. >. FWNCPNMJ?PGX?RGML ?. RFC RFPCQFMJB MD CVAGR?RGML @. RFC PCDP?ARMPW NCPGMB A. GLFG@GRMPW NMQRQWL?NRGA NMRCLRG?J 8. "DRCP ?L ?ARGML NMRCLRG?J, RFC MNCLGLE MD ?BBGRGML?J TMJR?EC-E?RCB >>>>>>>> AF?LLCJQ ?LB RFC GL?ARGT?RGML MD QMBGSK AF?LLCJQ, A?SQC RFC KCK@P?LC RM PCRSPL RM GRQ PCQRGLE KCK@P?LC NMRCLRG?J. >. QMBGSK ?. NMR?QQGSK @. A?JAGSK A. AFJMPGBC 9. 8F?R GQ RFC RCPK DMP NPMRCGL AF?LLCJQ RF?R AMLLCAR RUM LCSPMLQ ?R ?L CJCARPGA?J QWL?NQC >. ?. @. A. QWL?NRGA TCQGAJCQ TMJR?EC-E?RCB GML AF?LLCJQ E?N HSLARGML NPMRCGL QMBGSK-NMR?QQGSK CVAF?LEC NSKNQ
17. >>>>>>>> KCBGA?RGMLQ ?PC MDRCL SQCB RM RPC?R N?RGCLRQ UGRF "%)%. >. 5P?LOSGJGXCP ?. "LRG@GMRGA @. 4RGKSJ?LR A. "LRG-QCGXSPC 18. 4RPMICQ ?PC MDRCL A?SQCB @W >>>>>>>>. >. ?. @. A. LCSPMBCECLCP?RGML @JMMB AJMRQ MP @SPQR @JMMB TCQQCJQ QCGXSPCQ TGPSQCQ
10. 5FC >>>>>>>> JM@C AMLR?GLQ RFC TGQS?J AMPRCV. >. ?. @. A. DPMLR?J N?PGCR?J RCKNMP?J MAAGNGR?J
11. 5FC >>>>>>>> AMLLCARQ RFC RUM ACPC@P?J FCKGQNFCPCQ.
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19. )MU ?PC LCSPMLQ QGKGJ?P RM MRFCP ACJJQ )MU ?PC RFCW SLGOSC 20. .SJRGNJC QAJCPMQGQ A?SQCQ BCKWCJGL?RGML MD ?VMLQ GL RFC @P?GL ?LB QNGL?J AMPB. 8FW GQ RFGQ NPM@JCK?RGA 21. )MU BMCQ KWCJGL ?GB NPMN?E?RGML MD ?L ?ARGML NMRCLRG?J ?JMLE ?L ?VML )MU BM RFC LMBCQ MD 3?LTGCP FCJN RFGQ NPMACQQ 22. 8F?R ?PC RFC K?GL QRCNQ GL AFCKGA?J LCSPMRP?LQKGQQGML
1030
#(!04%2 35 X 4(% .%26/53 3934%-
23. 8F?R KCRFMBQ A?L @C SQCB RM BCRCPKGLC RFC DSLARGML MD ? N?PRGASJ?P @P?GL PCEGML 24. 8F?R ?PC RFC K?GL DSLARGMLQ MD RFC QNGL?J AMPB 25. 8F?R ?PC RFC K?GL BGDDCPCLACQ @CRUCCL RFC QWKN?RFCRGA ?LB N?P?QWKN?RFCRGA @P?LAFCQ MD RFC ?SRMLMKGA LCPTMSQ QWQRCK
26. 8F?R ?PC RFC K?GL DSLARGMLQ MD RFC QCLQMPWQMK?RGA LCPTMSQ QWQRCK 27. 8F?R ?PC RFC K?GL QWKNRMKQ MD "JXFCGKCPsQ BGQC?QC 28. 8F?R ?PC NMQQG@JC RPC?RKCLRQ DMP N?RGCLRQ UGRF K?HMP BCNPCQQGML
#(!04%2 36 X 3%.3/29 3934%-3
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*L KMPC ?BT?LACB ?LGK?JQ, RFC QCLQCQ ?PC AMLQR?LRJW ?R UMPI, K?IGLE RFC ?LGK?J ?U?PC MD QRGKSJGqQSAF ?Q JGEFR, MP QMSLB, MP RFC NPCQCLAC MD ? AFCKGA?J QS@QR?LAC GL RFC CVRCPL?J CLTGPMLKCLRq?LB KMLGRMPGLE GLDMPK?RGML ?@MSR RFC MPE?LGQKsQ GLRCPL?J CLTGPMLKCLR. "JJ @GJ?RCP?JJW QWKKCRPGA ?LGK?JQ F?TC ? QCLQMPW QWQRCK, ?LB RFC BCTCJMNKCLR MD ?LW QNCAGCQs QCLQMPW QWQRCK F?Q @CCL BPGTCL @W L?RSP?J QCJCARGML; RFSQ, QCLQMPW QWQRCKQ BGDDCP ?KMLE QNCAGCQ ?AAMPBGLE RM RFC BCK?LBQ MD RFCGP CLTGPMLKCLRQ. 5FC QF?PI, SLJGIC KMQR DGQF NPCB?RMPQ, GQ CJCARPMQCLQGRGTCqRF?R GQ, QCLQGRGTC RM CJCARPGA?J DGCJBQ NPMBSACB @W MRFCP ?LGK?JQ GL GRQ CLTGPMLKCLR. 8FGJC GR GQ FCJNDSJ RM RFGQ SLBCPU?RCP NPCB?RMP, CJCARPMQCLQGRGTGRW GQ ? QCLQC LMR DMSLB GL KMQR J?LB ?LGK?JQ.
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z *BCLRGDW RFC ECLCP?J ?LB QNCAG?J QCLQCQ GL FSK?LQ z %CQAPG@C RFPCC GKNMPR?LR QRCNQ GL QCLQMPW NCPACNRGML z &VNJ?GL RFC AMLACNR MD HSQR-LMRGAC?@JC BGDDCPCLAC GL QCLQMPW NCPACNRGML
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#(!04%2 36 X 3%.3/29 3934%-3
4CLQCQ NPMTGBC GLDMPK?RGML ?@MSR RFC @MBW ?LB GRQ CLTGPMLKCLR. )SK?LQ F?TC DGTC QNCAG?J QCLQCQ: MJD?ARGML (QKCJJ), ESQR?RGML (R?QRC), COSGJG@PGSK (@?J?LAC ?LB @MBW NMQGRGML), TGQGML, ?LB FC?PGLE. "BBGRGML?JJW, UC NMQQCQQ ECLCP?J QCLQCQ, ?JQM A?JJCB QMK?RMQCLQ?RGML, UFGAF PCQNMLB RM QRGKSJG JGIC RCKNCP?RSPC, N?GL, NPCQQSPC, ?LB TG@P?RGML. Vestibular sensation, UFGAF GQ ?L MPE?LGQKsQ QCLQC MD QN?RG?J MPGCLR?RGML ?LB @?J?LAC, proprioception (NMQGRGML MD @MLCQ, HMGLRQ, ?LB KSQAJCQ), ?LB RFC QCLQC MD JGK@ NMQGRGML RF?R GQ SQCB RM RP?AI kinesthesia (JGK@ KMTCKCLR) ?PC N?PR MD QMK?RMQCLQ?RGML. "JRFMSEF RFC QCLQMPW QWQRCKQ ?QQMAG?RCB UGRF RFCQC QCLQCQ ?PC TCPW BGDDCPCLR, ?JJ QF?PC ? AMKKML DSLARGML: RM AMLTCPR ? QRGKSJSQ (QSAF ?Q JGEFR, MP QMSLB, MP RFC NMQGRGML MD RFC @MBW) GLRM ?L CJCARPGA?J QGEL?J GL RFC LCPTMSQ QWQRCK. 5FGQ NPMACQQ GQ A?JJCB sensory transduction. 5FCPC ?PC RUM @PM?B RWNCQ MD ACJJSJ?P QWQRCKQ RF?R NCPDMPK QCLQMPW RP?LQBSARGML. *L MLC, ? LCSPML UMPIQ UGRF ? sensory receptor, ? ACJJ, MP ACJJ NPMACQQ RF?R GQ QNCAG?JGXCB RM CLE?EC UGRF ?LB BCRCAR ? QNCAGDGA QRGKSJSQ. 4RGKSJ?RGML MD RFC QCLQMPW PCACNRMP ?ARGT?RCQ RFC ?QQMAG?RCB ?DDCPCLR LCSPML, UFGAF A?PPGCQ GLDMPK?RGML ?@MSR RFC QRGKSJSQ RM RFC ACLRP?J LCPTMSQ QWQRCK. *L RFC QCAMLB RWNC MD QCLQMPW RP?LQBSARGML, ? QCLQMPW LCPTC CLBGLE PCQNMLBQ RM ? QRGKSJSQ GL RFC GLRCPL?J MP CVRCPL?J CLTGPMLKCLR: RFGQ LCSPML AMLQRGRSRCQ RFC QCLQMPW PCACNRMP. 'PCC LCPTC CLBGLEQ A?L @C QRGKSJ?RCB @W QCTCP?J BGDDCPCLR QRGKSJG, RFSQ QFMUGLE JGRRJC PCACNRMP QNCAGDGAGRW. 'MP CV?KNJC, N?GL PCACNRMPQ GL WMSP ESKQ ?LB RCCRF K?W @C QRGKSJ?RCB @W RCKNCP?RSPC AF?LECQ, AFCKGA?J QRGKSJ?RGML, MP NPCQQSPC.
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5FC DGPQR QRCN GL QCLQ?RGML GQ reception, UFGAF GQ RFC ?ARGT?RGML MD QCLQMPW PCACNRMPQ @W QRGKSJG QSAF ?Q KCAF?LGA?J QRGKSJG (@CGLE @CLR MP QOSGQFCB, DMP CV?KNJC), AFCKGA?JQ, MP RCKNCP?RSPC. 5FC PCACNRMP A?L RFCL PCQNMLB RM RFC QRGKSJG. 5FC PCEGML GL QN?AC GL UFGAF ? EGTCL QCLQMPW PCACNRMP A?L PCQNMLB RM ? QRGKSJSQ, @C GR D?P ?U?W MP GL AMLR?AR UGRF RFC @MBW, GQ RF?R PCACNRMPsQ receptive field. 5FGLI DMP ? KMKCLR ?@MSR RFC BGDDCPCLACQ GL PCACNRGTC DGCJBQ DMP RFC BGDDCPCLR QCLQCQ. 'MP RFC QCLQC MD RMSAF, ? QRGKSJSQ KSQR AMKC GLRM AMLR?AR UGRF @MBW. 'MP RFC QCLQC MD FC?PGLE, ? QRGKSJSQ A?L @C ? KMBCP?RC BGQR?LAC ?U?W (QMKC @?JCCL UF?JC QMSLBQ A?L NPMN?E?RC DMP K?LW IGJMKCRCPQ). 'MP TGQGML, ? QRGKSJSQ A?L @C TCPW D?P ?U?W; DMP CV?KNJC, RFC TGQS?J QWQRCK NCPACGTCQ JGEFR DPMK QR?PQ ?R CLMPKMSQ BGQR?LACQ.
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5FC KMQR DSLB?KCLR?J DSLARGML MD ? QCLQMPW QWQRCK GQ RFC RP?LQJ?RGML MD ? QCLQMPW QGEL?J RM ?L CJCARPGA?J QGEL?J GL RFC LCPTMSQ QWQRCK. 5FGQ R?ICQ NJ?AC ?R RFC QCLQMPW PCACNRMP, ?LB RFC AF?LEC GL CJCARPGA?J NMRCLRG?J RF?R GQ NPMBSACB GQ A?JJCB RFC receptor potential. )MU GQ QCLQMPW GLNSR, QSAF ?Q NPCQQSPC ML RFC QIGL, AF?LECB RM ? PCACNRMP NMRCLRG?J *L RFGQ CV?KNJC, ? RWNC MD PCACNRMP A?JJCB ? mechanoreceptor (?Q QFMUL GL Figure 36.2) NMQQCQQCQ QNCAG?JGXCB KCK@P?LCQ RF?R PCQNMLB RM NPCQQSPC. %GQRSP@?LAC MD RFCQC BCLBPGRCQ @W AMKNPCQQGLE RFCK MP @CLBGLE RFCK MNCLQ E?RCB GML AF?LLCJQ GL RFC NJ?QK? KCK@P?LC MD RFC QCLQMPW LCSPML, AF?LEGLE GRQ CJCARPGA?J NMRCLRG?J. 3CA?JJ RF?R GL RFC LCPTMSQ QWQRCK, ? NMQGRGTC AF?LEC MD ? LCSPMLsQ CJCARPGA?J NMRCLRG?J (?JQM A?JJCB RFC KCK@P?LC NMRCLRG?J), BCNMJ?PGXCQ RFC LCSPML. 3CACNRMP NMRCLRG?JQ ?PC EP?BCB NMRCLRG?JQ: RFC K?ELGRSBC MD RFCQC EP?BCB (PCACNRMP) NMRCLRG?JQ T?PGCQ UGRF RFC QRPCLERF MD RFC QRGKSJSQ. *D RFC K?ELGRSBC MD BCNMJ?PGX?RGML GQ QSDDGAGCLR (RF?R GQ, GD KCK@P?LC NMRCLRG?J PC?AFCQ ? RFPCQFMJB), RFC LCSPML UGJJ DGPC ?L ?ARGML NMRCLRG?J. *L KMQR A?QCQ, RFC AMPPCAR QRGKSJSQ GKNGLEGLE ML ? QCLQMPW PCACNRMP UGJJ BPGTC KCK@P?LC NMRCLRG?J GL ? NMQGRGTC BGPCARGML, ?JRFMSEF DMP QMKC PCACNRMPQ, QSAF ?Q RFMQC GL RFC TGQS?J QWQRCK, RFGQ GQ LMR ?JU?WQ RFC A?QC.
#(!04%2 36 X 3%.3/29 3934%-3
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4CLQMPW PCACNRMPQ DMP BGDDCPCLR QCLQCQ ?PC TCPW BGDDCPCLR DPMK C?AF MRFCP, ?LB RFCW ?PC QNCAG?JGXCB ?AAMPBGLE RM RFC RWNC MD QRGKSJSQ RFCW QCLQC: RFCW F?TC PCACNRMP QNCAGDGAGRW. 'MP CV?KNJC, RMSAF PCACNRMPQ, JGEFR PCACNRMPQ, ?LB QMSLB PCACNRMPQ ?PC C?AF ?ARGT?RCB @W BGDDCPCLR QRGKSJG. 5MSAF PCACNRMPQ ?PC LMR QCLQGRGTC RM JGEFR MP QMSLB; RFCW ?PC QCLQGRGTC MLJW RM RMSAF MP NPCQQSPC. )MUCTCP, QRGKSJG K?W @C AMK@GLCB ?R FGEFCP JCTCJQ GL RFC @P?GL, ?Q F?NNCLQ UGRF MJD?ARGML, AMLRPG@SRGLE RM MSP QCLQC MD R?QRC. E3)4*/3- '3* "7'382/88/43 4, !+3847> I3,472'9/43 'MSP ?QNCARQ MD QCLQMPW GLDMPK?RGML ?PC CLAMBCB @W QCLQMPW QWQRCKQ: RFC RWNC MD QRGKSJSQ, RFC JMA?RGML MD RFC QRGKSJSQ GL RFC PCACNRGTC DGCJB, RFC BSP?RGML MD RFC QRGKSJSQ, ?LB RFC PCJ?RGTC GLRCLQGRW MD RFC QRGKSJSQ. 5FSQ, ?ARGML NMRCLRG?JQ RP?LQKGRRCB MTCP ? QCLQMPW PCACNRMPsQ ?DDCPCLR ?VMLQ CLAMBC MLC RWNC MD QRGKSJSQ, ?LB RFGQ QCEPCE?RGML MD RFC QCLQCQ GQ NPCQCPTCB GL MRFCP QCLQMPW AGPASGRQ. 'MP CV?KNJC, ?SBGRMPW PCACNRMPQ RP?LQKGR QGEL?JQ MTCP RFCGP MUL BCBGA?RCB QWQRCK, ?LB CJCARPGA?J ?ARGTGRW GL RFC ?VMLQ MD RFC ?SBGRMPW PCACNRMPQ UGJJ @C GLRCPNPCRCB @W RFC @P?GL ?Q ?L ?SBGRMPW QRGKSJSQq? QMSLB. 5FC GLRCLQGRW MD ? QRGKSJSQ GQ MDRCL CLAMBCB GL RFC P?RC MD ?ARGML NMRCLRG?JQ NPMBSACB @W RFC QCLQMPW PCACNRMP. 5FSQ, ?L GLRCLQC QRGKSJSQ UGJJ NPMBSAC ? KMPC P?NGB RP?GL MD ?ARGML NMRCLRG?JQ, ?LB PCBSAGLE RFC QRGKSJSQ UGJJ JGICUGQC QJMU RFC P?RC MD NPMBSARGML MD ?ARGML NMRCLRG?JQ. " QCAMLB U?W GL UFGAF GLRCLQGRW GQ CLAMBCB GQ @W RFC LSK@CP MD PCACNRMPQ ?ARGT?RCB. "L GLRCLQC QRGKSJSQ KGEFR GLGRG?RC ?ARGML NMRCLRG?JQ GL ? J?PEC LSK@CP MD ?BH?ACLR PCACNRMPQ, UFGJC ? JCQQ GLRCLQC QRGKSJSQ KGEFR QRGKSJ?RC DCUCP PCACNRMPQ. *LRCEP?RGML MD QCLQMPW GLDMPK?RGML @CEGLQ ?Q QMML ?Q RFC GLDMPK?RGML GQ PCACGTCB GL RFC $/4, ?LB RFC @P?GL UGJJ DSPRFCP NPMACQQ GLAMKGLE QGEL?JQ.
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1034
#(!04%2 36 X 3%.3/29 3934%-3
?ARGML NMRCLRG?JQ DPMK QCLQMPW PCACNRMPQ RP?TCJ ?JMLE LCSPMLQ RF?R ?PC BCBGA?RCB RM ? N?PRGASJ?P QRGKSJSQ. 5FCQC LCSPMLQ ?PC BCBGA?RCB RM RF?R N?PRGASJ?P QRGKSJSQ ?LB QWL?NQC UGRF N?PRGASJ?P LCSPMLQ GL RFC @P?GL MP QNGL?J AMPB. "JJ QCLQMPW QGEL?JQ, CVACNR RFMQC DPMK RFC MJD?ARMPW QWQRCK, ?PC RP?LQKGRRCB RFMSEF RFC ACLRP?J LCPTMSQ QWQRCK ?LB ?PC PMSRCB RM RFC RF?J?KSQ ?LB RM RFC ?NNPMNPG?RC PCEGML MD RFC AMPRCV. 3CA?JJ RF?R RFC RF?J?KSQ GQ ? QRPSARSPC GL RFC DMPC@P?GL RF?R QCPTCQ ?Q ? AJC?PGLEFMSQC ?LB PCJ?W QR?RGML DMP QCLQMPW (?Q UCJJ ?Q KMRMP) QGEL?JQ. 8FCL RFC QCLQMPW QGEL?J CVGRQ RFC RF?J?KSQ, GR GQ AMLBSARCB RM RFC QNCAGDGA ?PC? MD RFC AMPRCV (Figure 36.3) BCBGA?RCB RM NPMACQQGLE RF?R N?PRGASJ?P QCLQC. )MU ?PC LCSP?J QGEL?JQ GLRCPNPCRCB *LRCPNPCR?RGML MD QCLQMPW QGEL?JQ @CRUCCL GLBGTGBS?JQ MD RFC Q?KC QNCAGCQ GQ J?PECJW QGKGJ?P, MUGLE RM RFC GLFCPGRCB QGKGJ?PGRW MD RFCGP LCPTMSQ QWQRCKQ; FMUCTCP, RFCPC ?PC QMKC GLBGTGBS?J BGDDCPCLACQ. " EMMB CV?KNJC MD RFGQ GQ GLBGTGBS?J RMJCP?LACQ RM ? N?GLDSJ QRGKSJSQ, QSAF ?Q BCLR?J N?GL, UFGAF ACPR?GLJW BGDDCP.
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%CCNCP GL RFC CNGBCPKGQ, LC?P RFC @?QC, ?PC Ruffini endings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acinian corpuscles (QCCL GL Figure 36.7) ?PC JMA?RCB BCCN GL RFC BCPKGQ MD @MRF EJ?@PMSQ ?LB F?GPW QIGL ?LB ?PC QRPSARSP?JJW QGKGJ?P RM .CGQQLCPsQ AMPNSQAJCQ; RFCW ?PC DMSLB GL RFC @MLC NCPGMQRCSK, HMGLR A?NQSJCQ, N?LAPC?Q ?LB MRFCP TGQACP?, @PC?QR, ?LB ECLGR?JQ. 5FCW ?PC P?NGBJW ?B?NRGLE KCAF?LMPCACNRMPQ RF?R QCLQC BCCN RP?LQGCLR (@SR LMR NPMJMLECB) NPCQQSPC ?LB FGEF-DPCOSCLAW TG@P?RGML. 1?AGLG?L PCACNRMPQ BCRCAR NPCQQSPC ?LB TG@P?RGML @W @CGLE AMKNPCQQCB, QRGKSJ?RGLE RFCGP GLRCPL?J BCLBPGRCQ. 5FCPC ?PC DCUCP 1?AGLG?L AMPNSQAJCQ ?LB 3SDDGLG CLBGLEQ GL QIGL RF?L RFCPC ?PC .CPICJsQ BGQIQ ?LB .CGQQLCPsQ AMPNSQAJCQ.
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#(!04%2 36 X 3%.3/29 3934%-3
1039
#?PMPCANRMPQ BCRCAR NPCQQSPC AF?LECQ GL ?L MPE?L. 5FCW ?PC DMSLB GL RFC U?JJQ MD RFC A?PMRGB ?PRCPW ?LB RFC ?MPR? UFCPC RFCW KMLGRMP @JMMB NPCQQSPC, ?LB GL RFC JSLEQ UFCPC RFCW BCRCAR RFC BCEPCC MD JSLE CVN?LQGML. 4RPCRAF PCACNRMPQ ?PC DMSLB ?R T?PGMSQ QGRCQ GL RFC BGECQRGTC ?LB SPGL?PW QWQRCKQ. *L ?BBGRGML RM RFCQC RUM RWNCQ MD BCCNCP PCACNRMPQ, RFCPC ?PC ?JQM P?NGBJW ?B?NRGLE F?GP PCACNRMPQ, UFGAF ?PC DMSLB ML LCPTC CLBGLEQ RF?R UP?N ?PMSLB RFC @?QC MD F?GP DMJJGAJCQ. 5FCPC ?PC ? DCU RWNCQ MD F?GP PCACNRMPQ RF?R BCRCAR QJMU ?LB P?NGB F?GP KMTCKCLR, ?LB RFCW BGDDCP GL RFCGP QCLQGRGTGRW RM KMTCKCLR. 4MKC F?GP PCACNRMPQ ?JQM BCRCAR QIGL BCDJCARGML, ?LB ACPR?GL P?NGBJW ?B?NRGLE F?GP PCACNRMPQ ?JJMU BCRCARGML MD QRGKSJG RF?R F?TC LMR WCR RMSAFCB RFC QIGL. I39+-7'9/43 4, !/-3'18 ,742 M+).'347+)+59478 5FC AMLDGESP?RGML MD RFC BGDDCPCLR RWNCQ MD PCACNRMPQ UMPIGLE GL AMLACPR GL FSK?L QIGL PCQSJRQ GL ? TCPW PCDGLCB QCLQC MD RMSAF. 5FC LMAGACNRGTC PCACNRMPQqRFMQC RF?R BCRCAR N?GLq?PC JMA?RCB LC?P RFC QSPD?AC. 4K?JJ, DGLCJW A?JG@P?RCB KCAF?LMPCACNRMPQq.CPICJsQ BGQIQ ?LB .CGQQLCPsQ AMPNSQAJCQq?PC JMA?RCB GL RFC SNNCP J?WCPQ ?LB A?L NPCAGQCJW JMA?JGXC CTCL ECLRJC RMSAF. 5FC J?PEC KCAF?LMPCACNRMPQq1?AGLG?L AMPNSQAJCQ ?LB 3SDDGLG CLBGLEQq?PC JMA?RCB GL RFC JMUCP J?WCPQ ?LB PCQNMLB RM BCCNCP RMSAF. ($MLQGBCP RF?R RFC BCCN NPCQQSPC RF?R PC?AFCQ RFMQC BCCNCP PCACNRMPQ UMSJB LMR LCCB RM @C DGLCJW JMA?JGXCB.) #MRF RFC SNNCP ?LB JMUCP J?WCPQ MD RFC QIGL FMJB P?NGBJW ?LB QJMUJW ?B?NRGLE PCACNRMPQ. #MRF NPGK?PW QMK?RMQCLQMPW AMPRCV ?LB QCAMLB?PW AMPRGA?J ?PC?Q ?PC PCQNMLQG@JC DMP NPMACQQGLE RFC AMKNJCV NGARSPC MD QRGKSJG RP?LQKGRRCB DPMK RFC GLRCPNJ?W MD KCAF?LMPCACNRMPQ. D+38/9> 4, M+).'347+)+59478 5FC BGQRPG@SRGML MD RMSAF PCACNRMPQ GL FSK?L QIGL GQ LMR AMLQGQRCLR MTCP RFC @MBW. *L FSK?LQ, RMSAF PCACNRMPQ ?PC JCQQ BCLQC GL QIGL AMTCPCB UGRF ?LW RWNC MD F?GP, QSAF ?Q RFC ?PKQ, JCEQ, RMPQM, ?LB D?AC. 5MSAF PCACNRMPQ ?PC BCLQCP GL EJ?@PMSQ QIGL (RFC RWNC DMSLB ML FSK?L DGLECPRGNQ ?LB JGNQ, DMP CV?KNJC), UFGAF GQ RWNGA?JJW KMPC QCLQGRGTC ?LB GQ RFGAICP RF?L F?GPW QIGL (4 RM 5 KK TCPQSQ 2 RM 3 KK). )MU GQ PCACNRMP BCLQGRW CQRGK?RCB GL ? FSK?L QS@HCAR 5FC PCJ?RGTC BCLQGRW MD NPCQQSPC PCACNRMPQ GL BGDDCPCLR JMA?RGMLQ ML RFC @MBW A?L @C BCKMLQRP?RCB CVNCPGKCLR?JJW SQGLE ? RUM-NMGLR BGQAPGKGL?RGML RCQR. *L RFGQ BCKMLQRP?RGML, RUM QF?PN NMGLRQ, QSAF ?Q RUM RFSK@R?AIQ, ?PC @PMSEFR GLRM AMLR?AR UGRF RFC QS@HCARsQ QIGL (RFMSEF LMR F?PB CLMSEF RM A?SQC N?GL MP @PC?I RFC QIGL). 5FC QS@HCAR PCNMPRQ GD FC MP QFC DCCJQ MLC NMGLR MP RUM NMGLRQ. *D RFC RUM NMGLRQ ?PC DCJR ?Q MLC NMGLR, GR A?L @C GLDCPPCB RF?R RFC RUM NMGLRQ ?PC @MRF GL RFC PCACNRGTC DGCJB MD ? QGLEJC QCLQMPW PCACNRMP. *D RUM NMGLRQ ?PC DCJR ?Q RUM QCN?P?RC NMGLRQ, C?AF GQ GL RFC PCACNRGTC DGCJB MD RUM QCN?P?RC QCLQMPW PCACNRMPQ. 5FC NMGLRQ AMSJB RFCL @C KMTCB AJMQCP ?LB PC-RCQRCB SLRGJ RFC QS@HCAR PCNMPRQ DCCJGLE MLJW MLC NMGLR, ?LB RFC QGXC MD RFC PCACNRGTC DGCJB MD ? QGLEJC PCACNRMP AMSJB @C CQRGK?RCB DPMK RF?R BGQR?LAC.
/?<IDFI<:<GK@FE
*L ?BBGRGML RM ,P?SQC CLB @SJ@Q RF?R BCRCAR AMJB ?LB 3SDDGLG CLBGLEQ RF?R BCRCAR U?PKRF, RFCPC ?PC BGDDCPCLR RWNCQ MD AMJB PCACNRMPQ ML QMKC DPCC LCPTC CLBGLEQ: RFCPKMPCACNRMPQ, JMA?RCB GL RFC BCPKGQ, QICJCR?J KSQAJCQ, JGTCP, ?LB FWNMRF?J?KSQ, RF?R ?PC ?ARGT?RCB @W BGDDCPCLR RCKNCP?RSPCQ. 5FCGP N?RFU?WQ GLRM RFC @P?GL PSL DPMK RFC QNGL?J AMPB RFPMSEF RFC RF?J?KSQ RM RFC NPGK?PW QMK?RMQCLQMPW AMPRCV. 8?PKRF ?LB AMJB GLDMPK?RGML DPMK RFC D?AC RP?TCJQ RFPMSEF MLC MD RFC AP?LG?J LCPTCQ RM RFC @P?GL. :MS ILMU DPMK CVNCPGCLAC RF?R ? RMJCP?@JW AMJB MP FMR QRGKSJSQ A?L OSGAIJW NPMEPCQQ RM ? KSAF KMPC GLRCLQC QRGKSJSQ RF?R GQ LM JMLECP RMJCP?@JC. "LW QRGKSJSQ RF?R GQ RMM GLRCLQC A?L @C NCPACGTCB ?Q N?GL @CA?SQC RCKNCP?RSPC QCLQ?RGMLQ ?PC AMLBSARCB ?JMLE RFC Q?KC N?RFU?WQ RF?R A?PPW N?GL QCLQ?RGMLQ
+8@E
1?GL GQ RFC L?KC EGTCL RM nociception, UFGAF GQ RFC LCSP?J NPMACQQGLE MD GLHSPGMSQ QRGKSJG GL PCQNMLQC RM RGQQSC B?K?EC. 1?GL GQ A?SQCB @W RPSC QMSPACQ MD GLHSPW, QSAF ?Q AMLR?AR UGRF ? FC?R QMSPAC RF?R A?SQCQ ? RFCPK?J @SPL MP AMLR?AR UGRF ? AMPPMQGTC AFCKGA?J. #SR N?GL ?JQM A?L @C A?SQCB @W F?PKJCQQ QRGKSJG RF?R KGKGA RFC ?ARGML MD B?K?EGLE QRGKSJG, QSAF ?Q AMLR?AR UGRF A?NQ?GAGLQ, RFC AMKNMSLBQ RF?R A?SQC NCNNCPQ RM R?QRC FMR ?LB UFGAF ?PC SQCB GL QCJD-BCDCLQC NCNNCP QNP?WQ ?LB ACPR?GL RMNGA?J KCBGA?RGMLQ. 1CNNCPQ R?QRC tFMRu @CA?SQC RFC NPMRCGL PCACNRMPQ RF?R @GLB A?NQ?GAGL MNCL RFC Q?KC A?JAGSK AF?LLCJQ RF?R ?PC ?ARGT?RCB @W U?PK PCACNRMPQ. /MAGACNRGML QR?PRQ ?R RFC QCLQMPW PCACNRMPQ, @SR N?GL, GL?QKSAF ?Q GR GQ RFC NCPACNRGML MD LMAGACNRGML, BMCQ LMR QR?PR SLRGJ GR GQ AMKKSLGA?RCB RM RFC @P?GL. 5FCPC ?PC QCTCP?J LMAGACNRGTC N?RFU?WQ RM ?LB RFPMSEF RFC @P?GL. .MQR ?VMLQ A?PPWGLE LMAGACNRGTC GLDMPK?RGML GLRM RFC @P?GL DPMK RFC QNGL?J AMPB NPMHCAR RM RFC RF?J?KSQ (?Q BM MRFCP QCLQMPW LCSPMLQ) ?LB RFC LCSP?J QGEL?J SLBCPEMCQ DGL?J NPMACQQGLE GL RFC NPGK?PW QMK?RMQCLQMPW AMPRCV. *LRCPCQRGLEJW, MLC LMAGACNRGTC N?RFU?W NPMHCARQ LMR RM RFC RF?J?KSQ @SR BGPCARJW RM RFC FWNMRF?J?KSQ GL RFC DMPC@P?GL, UFGAF KMBSJ?RCQ RFC A?PBGMT?QASJ?P ?LB LCSPMCLBMAPGLC DSLARGMLQ MD RFC ?SRMLMKGA LCPTMSQ QWQRCK. 3CA?JJ RF?R RFPC?RCLGLEqMP N?GLDSJqQRGKSJG QRGKSJ?RC RFC QWKN?RFCRGA @P?LAF MD RFC TGQACP?J QCLQMPW QWQRCK, PC?BWGLE ? DGEFR-MP-DJGEFR PCQNMLQC.
1040
#(!04%2 36 X 3%.3/29 3934%-3
7GCU RFGQ video (http://openstaxcollege.org/l/nociceptive) RF?R ?LGK?RCQ RFC DGTC NF?QCQ MD LMAGACNRGTC N?GL.
36.3 R /8JK< 8E; .D<CC

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL GL UF?R U?W QKCJJ ?LB R?QRC QRGKSJG BGDDCP DPMK MRFCP QCLQMPW QRGKSJG z *BCLRGDW RFC DGTC NPGK?PW R?QRCQ RF?R A?L @C BGQRGLESGQFCB @W FSK?LQ z &VNJ?GL GL ?L?RMKGA?J RCPKQ UFW ? BMEsQ QCLQC MD QKCJJ GQ KMPC ?ASRC RF?L ? FSK?LsQ 5?QRC, ?JQM A?JJCB gustation, ?LB QKCJJ, ?JQM A?JJCB olfaction, ?PC RFC KMQR GLRCPAMLLCARCB QCLQCQ GL RF?R @MRF GLTMJTC KMJCASJCQ MD RFC QRGKSJSQ CLRCPGLE RFC @MBW ?LB @MLBGLE RM PCACNRMPQ. 4KCJJ JCRQ ?L ?LGK?J QCLQC RFC NPCQCLAC MD DMMB MP MRFCP ?LGK?JQqUFCRFCP NMRCLRG?J K?RCQ, NPCB?RMPQ, MP NPCWq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
/8JK<J 8E; *;FIJ

#MRF R?QRC ?LB MBMP QRGKSJG ?PC KMJCASJCQ R?ICL GL DPMK RFC CLTGPMLKCLR. 5FC NPGK?PW R?QRCQ BCRCARCB @W FSK?LQ ?PC QUCCR, QMSP, @GRRCP, Q?JRW ?LB SK?KG. 5FC DGPQR DMSP R?QRCQ LCCB JGRRJC CVNJ?L?RGML. 5FC GBCLRGDGA?RGML MD umami ?Q ? DSLB?KCLR?J R?QRC MAASPPCB D?GPJW PCACLRJWqGR U?Q GBCLRGDGCB GL 1908 @W +?N?LCQC QAGCLRGQR ,GISL?C *ICB? UFGJC FC UMPICB UGRF QC?UCCB @PMRF, @SR GR U?Q LMR UGBCJW ?AACNRCB ?Q ? R?QRC RF?R AMSJB @C NFWQGMJMEGA?JJW BGQRGLESGQFCB SLRGJ K?LW WC?PQ J?RCP. 5FC R?QRC MD SK?KG, ?JQM ILMUL ?Q Q?TMPGLCQQ, GQ ?RRPG@SR?@JC RM RFC R?QRC MD RFC ?KGLM ?AGB --EJSR?K?RC. *L D?AR, KMLMQMBGSK EJSR?K?RC, MP .4(, GQ MDRCL SQCB GL AMMIGLE RM CLF?LAC RFC Q?TMPW R?QRC MD ACPR?GL DMMBQ. 8F?R GQ RFC ?B?NRGTC T?JSC MD @CGLE ?@JC RM BGQRGLESGQF SK?KG 4?TMPW QS@QR?LACQ RCLB RM @C FGEF GL NPMRCGL. "JJ MBMPQ RF?R UC NCPACGTC ?PC KMJCASJCQ GL RFC ?GP UC @PC?RFC. *D ? QS@QR?LAC BMCQ LMR PCJC?QC KMJCASJCQ GLRM RFC ?GP DPMK GRQ QSPD?AC, GR F?Q LM QKCJJ. "LB GD ? FSK?L MP MRFCP ?LGK?J BMCQ LMR F?TC ? PCACNRMP RF?R PCAMELGXCQ ? QNCAGDGA KMJCASJC, RFCL RF?R KMJCASJC F?Q LM QKCJJ. )SK?LQ F?TC ?@MSR 350 MJD?ARMPW PCACNRMP QS@RWNCQ RF?R UMPI GL T?PGMSQ AMK@GL?RGMLQ RM ?JJMU SQ RM QCLQC ?@MSR 10,000 BGDDCPCLR MBMPQ. $MKN?PC RF?R RM KGAC, DMP CV?KNJC, UFGAF F?TC ?@MSR 1,300 MJD?ARMPW PCACNRMP RWNCQ, ?LB RFCPCDMPC NPM@?@JW QCLQC KMPC MBMPQ. #MRF MBMPQ ?LB R?QRCQ GLTMJTC KMJCASJCQ RF?R QRGKSJ?RC QNCAGDGA AFCKMPCACNRMPQ. "JRFMSEF FSK?LQ AMKKMLJW BGQRGLESGQF R?QRC ?Q MLC QCLQC ?LB QKCJJ ?Q ?LMRFCP, RFCW UMPI RMECRFCP RM APC?RC RFC NCPACNRGML MD DJ?TMP. " NCPQMLsQ NCPACNRGML MD DJ?TMP GQ PCBSACB GD FC MP QFC F?Q AMLECQRCB L?Q?J N?QQ?ECQ.
-<:<GK@FE 8E; /I8EJ;L:K@FE

Odorants (MBMP KMJCASJCQ) CLRCP RFC LMQC ?LB BGQQMJTC GL RFC MJD?ARMPW CNGRFCJGSK, RFC KSAMQ? ?R RFC @?AI MD RFC L?Q?J A?TGRW (?Q GJJSQRP?RCB GL Figure 36.8). 5FC olfactory epithelium GQ ? AMJJCARGML MD QNCAG?JGXCB MJD?ARMPW PCACNRMPQ GL RFC @?AI MD RFC L?Q?J A?TGRW RF?R QN?LQ ?L ?PC? ?@MSR 5 AK2 GL FSK?LQ. 3CA?JJ RF?R QCLQMPW ACJJQ ?PC LCSPMLQ. "L olfactory receptor, UFGAF GQ ? BCLBPGRC MD ? QNCAG?JGXCB LCSPML, PCQNMLBQ UFCL GR @GLBQ ACPR?GL KMJCASJCQ GLF?JCB DPMK RFC CLTGPMLKCLR @W QCLBGLE GKNSJQCQ BGPCARJW RM RFC MJD?ARMPW @SJ@ MD RFC @P?GL. )SK?LQ F?TC ?@MSR 12 KGJJGML MJD?ARMPW PCACNRMPQ, BGQRPG@SRCB ?KMLE FSLBPCBQ MD BGDDCPCLR PCACNRMP RWNCQ RF?R PCQNMLB RM BGDDCPCLR MBMPQ. 5UCJTC KGJJGML QCCKQ JGIC ? J?PEC LSK@CP MD PCACNRMPQ, @SR AMKN?PC RF?R RM MRFCP ?LGK?JQ: P?@@GRQ F?TC ?@MSR 100 KGJJGML, KMQR BMEQ F?TC
#(!04%2 36 X 3%.3/29 3934%-3
1041
?@MSR 1 @GJJGML, ?LB @JMMBFMSLBQqBMEQ QCJCARGTCJW @PCB DMP RFCGP QCLQC MD QKCJJqF?TC ?@MSR 4 @GJJGML. 5FC MTCP?JJ QGXC MD RFC MJD?ARMPW CNGRFCJGSK ?JQM BGDDCPQ @CRUCCL QNCAGCQ, UGRF RF?R MD @JMMBFMSLBQ, DMP CV?KNJC, @CGLE K?LW RGKCQ J?PECP RF?L RF?R MD FSK?LQ. 0JD?ARMPW LCSPMLQ ?PC bipolar neurons (LCSPMLQ UGRF RUM NPMACQQCQ DPMK RFC ACJJ @MBW). &?AF LCSPML F?Q ? QGLEJC BCLBPGRC @SPGCB GL RFC MJD?ARMPW CNGRFCJGSK, ?LB CVRCLBGLE DPMK RFGQ BCLBPGRC ?PC 5 RM 20 PCACNRMP-J?BCL, F?GP-JGIC AGJG? RF?R RP?N MBMP?LR KMJCASJCQ. 5FC QCLQMPW PCACNRMPQ ML RFC AGJG? ?PC NPMRCGLQ, ?LB GR GQ RFC T?PG?RGMLQ GL RFCGP ?KGLM ?AGB AF?GLQ RF?R K?IC RFC PCACNRMPQ QCLQGRGTC RM BGDDCPCLR MBMP?LRQ. &?AF MJD?ARMPW QCLQMPW LCSPML F?Q MLJW MLC RWNC MD PCACNRMP ML GRQ AGJG?, ?LB RFC PCACNRMPQ ?PC QNCAG?JGXCB RM BCRCAR QNCAGDGA MBMP?LRQ, QM RFC @GNMJ?P LCSPMLQ RFCKQCJTCQ ?PC QNCAG?JGXCB. 8FCL ?L MBMP?LR @GLBQ UGRF ? PCACNRMP RF?R PCAMELGXCQ GR, RFC QCLQMPW LCSPML ?QQMAG?RCB UGRF RFC PCACNRMP GQ QRGKSJ?RCB. 0JD?ARMPW QRGKSJ?RGML GQ RFC MLJW QCLQMPW GLDMPK?RGML RF?R BGPCARJW PC?AFCQ RFC ACPC@P?J AMPRCV, UFCPC?Q MRFCP QCLQ?RGMLQ ?PC PCJ?WCB RFPMSEF RFC RF?J?KSQ.
!@>LI< 36.8 )K QEB ERJ>K LIC>@QLOV PVPQBJ, (>) ?FMLI>O LIC>@QLOV KBROLKP BUQBKA COLJ (?) QEB LIC>@QLOV BMFQEBIFRJ, TEBOB LIC>@QLOV OB@BMQLOP >OB IL@>QBA, QL QEB LIC>@QLOV ?RI?. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V 0>QOF@H *. ,VK@E, JBAF@>I FIIRPQO>QLO; #. #>OI *>CCB, -$, @>OAFLILDFPQ)
1042
#(!04%2 36 X 3%.3/29 3934%-3
+?<IFDFE<J
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w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
!@>LI< 36.9 4EB CIBEJBK OBPMLKPB FK QEFP QFDBO OBPRIQP FK QEB @ROIFKD LC QEB RMMBO IFM >KA EBIMP >FO?LOKB MEBOLJLKB JLIB@RIBP BKQBO QEB SLJBOLK>P>I LOD>K. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V "@E>AE"/&IF@HO)
"'89+ %CRCARGLE ? R?QRC (ESQR?RGML) GQ D?GPJW QGKGJ?P RM BCRCARGLE ?L MBMP (MJD?ARGML), EGTCL RF?R @MRF R?QRC ?LB QKCJJ PCJW ML AFCKGA?J PCACNRMPQ @CGLE QRGKSJ?RCB @W ACPR?GL KMJCASJCQ. 5FC NPGK?PW MPE?L MD R?QRC GQ RFC R?QRC @SB. " taste bud GQ ? AJSQRCP MD ESQR?RMPW PCACNRMPQ (R?QRC ACJJQ) RF?R ?PC JMA?RCB UGRFGL RFC @SKNQ ML RFC RMLESC A?JJCB papillae (QGLESJ?P: N?NGJJ?) (GJJSQRP?RCB GL Figure 36.10). 5FCPC ?PC QCTCP?J QRPSARSP?JJW BGQRGLAR N?NGJJ?C. 'GJGDMPK N?NGJJ?C, UFGAF ?PC JMA?RCB ?APMQQ RFC RMLESC, ?PC R?ARGJC, NPMTGBGLE DPGARGML RF?R FCJNQ RFC RMLESC KMTC QS@QR?LACQ, ?LB AMLR?GL LM R?QRC ACJJQ. *L AMLRP?QR, DSLEGDMPK N?NGJJ?C, UFGAF ?PC JMA?RCB K?GLJW ML RFC ?LRCPGMP RUM-RFGPBQ MD RFC RMLESC, C?AF AMLR?GL MLC RM CGEFR R?QRC @SBQ ?LB ?JQM F?TC PCACNRMPQ DMP NPCQQSPC ?LB RCKNCP?RSPC. 5FC J?PEC AGPASKT?JJ?RC N?NGJJ?C AMLR?GL SN RM 100 R?QRC @SBQ ?LB DMPK ? 7 LC?P RFC NMQRCPGMP K?PEGL MD RFC RMLESC.
#(!04%2 36 X 3%.3/29 3934%-3
1043
!@>LI< 36.10 (>) &LIF>QB, @FO@RJS>II>QB, >KA CRKDFCLOJ M>MFII>B >OB IL@>QBA LK AFCCBOBKQ OBDFLKP LC QEB QLKDRB. (?) &LIF>QB M>MFII>B >OB MOLJFKBKQ MOLQORPFLKP LK QEFP IFDEQ JF@OLDO>ME. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V .#); P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
*L ?BBGRGML RM RFMQC RUM RWNCQ MD AFCKGA?JJW ?LB KCAF?LGA?JJW QCLQGRGTC N?NGJJ?C ?PC DMJG?RC N?NGJJ?CqJC?D-JGIC N?NGJJ?C JMA?RCB GL N?P?JJCJ DMJBQ ?JMLE RFC CBECQ ?LB RMU?PB RFC @?AI MD RFC RMLESC, ?Q QCCL GL RFC Figure 36.10 KGAPMEP?NF. 'MJG?RC N?NGJJ?C AMLR?GL ?@MSR 1,300 R?QRC @SBQ UGRFGL RFCGP DMJBQ. 'GL?JJW, RFCPC ?PC AGPASKT?JJ?RC N?NGJJ?C, UFGAF ?PC U?JJ-JGIC N?NGJJ?C GL RFC QF?NC MD ?L GLTCPRCB t7u ?R RFC @?AI MD RFC RMLESC. &?AF MD RFCQC N?NGJJ?C GQ QSPPMSLBCB @W ? EPMMTC ?LB AMLR?GLQ ?@MSR 250 R?QRC @SBQ. &?AF R?QRC @SBsQ R?QRC ACJJQ ?PC PCNJ?ACB CTCPW 10 RM 14 B?WQ. 5FCQC ?PC CJMLE?RCB ACJJQ UGRF F?GPJGIC NPMACQQCQ A?JJCB KGAPMTGJJG ?R RFC RGNQ RF?R CVRCLB GLRM RFC R?QRC @SB NMPC (GJJSQRP?RC GL Figure 36.11). 'MMB KMJCASJCQ ( tastants) ?PC BGQQMJTCB GL Q?JGT?, ?LB RFCW @GLB UGRF ?LB QRGKSJ?RC RFC PCACNRMPQ ML RFC KGAPMTGJJG. 5FC PCACNRMPQ DMP R?QR?LRQ ?PC JMA?RCB ?APMQQ RFC MSRCP NMPRGML ?LB DPMLR MD RFC RMLESC, MSRQGBC MD RFC KGBBJC ?PC? UFCPC RFC DGJGDMPK N?NGJJ?C ?PC KMQR NPMKGLCLR.
!@>LI< 36.11 0LOBP FK QEB QLKDRB >IILT Q>PQ>KQP QL BKQBO Q>PQB MLOBP FK QEB QLKDRB. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V 6FK@BKWL 2FWWL)
*L FSK?LQ, RFCPC ?PC DGTC NPGK?PW R?QRCQ, ?LB C?AF R?QRC F?Q MLJW MLC AMPPCQNMLBGLE RWNC MD PCACNRMP. 5FSQ, JGIC MJD?ARGML, C?AF PCACNRMP GQ QNCAGDGA RM GRQ QRGKSJSQ (R?QR?LR). 5P?LQBSARGML MD RFC DGTC R?QRCQ F?NNCLQ RFPMSEF BGDDCPCLR KCAF?LGQKQ RF?R PCDJCAR RFC KMJCASJ?P AMKNMQGRGML MD RFC R?QR?LR. " Q?JRW R?QR?LR (AMLR?GLGLE /?$J) NPMTGBCQ RFC QMBGSK GMLQ (/?+) RF?R CLRCP RFC R?QRC LCSPMLQ ?LB CVAGRC RFCK BGPCARJW. 4MSP R?QR?LRQ ?PC ?AGBQ ?LB @CJMLE RM RFC RFCPKMPCACNRMP NPMRCGL D?KGJW. #GLBGLE MD ?L ?AGB MP MRFCP QMSP-R?QRGLE KMJCASJC RPGEECPQ ? AF?LEC GL RFC GML AF?LLCJ ?LB RFCQC GLAPC?QC FWBPMECL GML ()+) AMLACLRP?RGMLQ GL RFC R?QRC LCSPMLQ, RFSQ BCNMJ?PGXGLE RFCK. 4UCCR, @GRRCP, ?LB SK?KG R?QR?LRQ PCOSGPC ? (-NPMRCGL AMSNJCB PCACNRMP. 5FCQC R?QR?LRQ @GLB RM RFCGP PCQNCARGTC PCACNRMPQ, RFCPC@W CVAGRGLE RFC QNCAG?JGXCB LCSPMLQ ?QQMAG?RCB UGRF RFCK.
1044
#(!04%2 36 X 3%.3/29 3934%-3
#MRF R?QRGLE ?@GJGRGCQ ?LB QCLQC MD QKCJJ AF?LEC UGRF ?EC. *L FSK?LQ, RFC QCLQCQ BCAJGLC BP?K?RGA?JJW @W ?EC 50 ?LB AMLRGLSC RM BCAJGLC. " AFGJB K?W DGLB ? DMMB RM @C RMM QNGAW, UFCPC?Q ?L CJBCPJW NCPQML K?W DGLB RFC Q?KC DMMB RM @C @J?LB ?LB SL?NNCRGXGLE.
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0JD?ARMPW LCSPMLQ NPMHCAR DPMK RFC MJD?ARMPW CNGRFCJGSK RM RFC MJD?ARMPW @SJ@ ?Q RFGL, SLKWCJGL?RCB ?VMLQ. 5FC olfactory bulb GQ AMKNMQCB MD LCSP?J AJSQRCPQ A?JJCB glomeruli, ?LB C?AF EJMKCPSJSQ PCACGTCQ QGEL?JQ DPMK MLC RWNC MD MJD?ARMPW PCACNRMP, QM C?AF EJMKCPSJSQ GQ QNCAGDGA RM MLC MBMP?LR. 'PMK EJMKCPSJG, MJD?ARMPW QGEL?JQ RP?TCJ BGPCARJW RM RFC MJD?ARMPW AMPRCV ?LB RFCL RM RFC DPMLR?J AMPRCV ?LB RFC RF?J?KSQ. 3CA?JJ RF?R RFGQ GQ ? BGDDCPCLR N?RF DPMK KMQR MRFCP QCLQMPW GLDMPK?RGML, UFGAF GQ QCLR BGPCARJW RM RFC RF?J?KSQ @CDMPC CLBGLE SN GL RFC AMPRCV. 0JD?ARMPW QGEL?JQ ?JQM RP?TCJ BGPCARJW RM RFC ?KWEB?J?, RFCPC?DRCP PC?AFGLE RFC FWNMRF?J?KSQ, RF?J?KSQ, ?LB DPMLR?J AMPRCV. 5FC J?QR QRPSARSPC RF?R MJD?ARMPW QGEL?JQ BGPCARJW RP?TCJ RM GQ ? AMPRGA?J ACLRCP GL RFC RCKNMP?J JM@C QRPSARSPC GKNMPR?LR GL QN?RG?J, ?SRM@GMEP?NFGA?J, BCAJ?P?RGTC, ?LB CNGQMBGA KCKMPGCQ. 0JD?ARGML GQ DGL?JJW NPMACQQCB @W ?PC?Q MD RFC @P?GL RF?R BC?J UGRF KCKMPW, CKMRGMLQ, PCNPMBSARGML, ?LB RFMSEFR. 5?QRC LCSPMLQ NPMHCAR DPMK R?QRC ACJJQ GL RFC RMLESC, CQMNF?ESQ, ?LB N?J?RC RM RFC KCBSJJ?, GL RFC @P?GLQRCK. 'PMK RFC KCBSJJ?, R?QRC QGEL?JQ RP?TCJ RM RFC RF?J?KSQ ?LB RFCL RM RFC NPGK?PW ESQR?RMPW AMPRCV. *LDMPK?RGML DPMK BGDDCPCLR PCEGMLQ MD RFC RMLESC GQ QCEPCE?RCB GL RFC KCBSJJ?, RF?J?KSQ, ?LB AMPRCV.
36.4 R #<8I@E> 8E; 1<JK@9LC8I .<EJ8K@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC PCJ?RGMLQFGN MD ?KNJGRSBC ?LB DPCOSCLAW MD ? QMSLB U?TC RM ?RRPG@SRCQ MD QMSLB z 5P?AC RFC N?RF MD QMSLB RFPMSEF RFC ?SBGRMPW QWQRCK RM RFC QGRC MD RP?LQBSARGML MD QMSLB z *BCLRGDW RFC QRPSARSPCQ MD RFC TCQRG@SJ?P QWQRCK RF?R PCQNMLB RM EP?TGRW Audition, MP FC?PGLE, GQ GKNMPR?LR RM FSK?LQ ?LB RM MRFCP ?LGK?JQ DMP K?LW BGDDCPCLR GLRCP?ARGMLQ. *R CL?@JCQ ?L MPE?LGQK RM BCRCAR ?LB PCACGTC GLDMPK?RGML ?@MSR B?LECP, QSAF ?Q ?L ?NNPM?AFGLE NPCB?RMP, ?LB RM N?PRGAGN?RC GL AMKKSL?J CVAF?LECQ JGIC RFMQC AMLACPLGLE RCPPGRMPGCQ MP K?RGLE. 0L RFC MRFCP F?LB, ?JRFMSEF GR GQ NFWQGA?JJW JGLICB RM RFC ?SBGRMPW QWQRCK, RFC TCQRG@SJ?P QWQRCK GQ LMR GLTMJTCB GL FC?PGLE. *LQRC?B, ?L ?LGK?JsQ TCQRG@SJ?P QWQRCK BCRCARQ GRQ MUL KMTCKCLR, @MRF JGLC?P ?LB ?LESJ?P ?AACJCP?RGML ?LB BCACJCP?RGML, ?LB @?J?LAC.
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"SBGRMPW QRGKSJG ?PC QMSLB U?TCQ, UFGAF ?PC KCAF?LGA?J, NPCQQSPC U?TCQ RF?R KMTC RFPMSEF ? KCBGSK, QSAF ?Q ?GP MP U?RCP. 5FCPC ?PC LM QMSLB U?TCQ GL ? T?ASSK QGLAC RFCPC ?PC LM ?GP KMJCASJCQ RM KMTC GL U?TCQ. 5FC QNCCB MD QMSLB U?TCQ BGDDCPQ, @?QCB ML ?JRGRSBC, RCKNCP?RSPC, ?LB KCBGSK, @SR ?R QC? JCTCJ ?LB ? RCKNCP?RSPC MD 20^ $ (68^ '), QMSLB U?TCQ RP?TCJ GL RFC ?GP ?R ?@MSR 343 KCRCPQ NCP QCAMLB. "Q GQ RPSC DMP ?JJ U?TCQ, RFCPC ?PC DMSP K?GL AF?P?ARCPGQRGAQ MD ? QMSLB U?TC: DPCOSCLAW, U?TCJCLERF, NCPGMB, ?LB ?KNJGRSBC. 'PCOSCLAW GQ RFC LSK@CP MD U?TCQ NCP SLGR MD RGKC, ?LB GL QMSLB GQ FC?PB ?Q NGRAF. )GEF-DPCOSCLAW ({15.000)X) QMSLBQ ?PC FGEFCP-NGRAFCB (QFMPR U?TCJCLERF) RF?L JMU-DPCOSCLAW (JMLE U?TCJCLERFQ; z100)X) QMSLBQ. 'PCOSCLAW GQ KC?QSPCB GL AWAJCQ NCP QCAMLB, ?LB DMP QMSLB, RFC KMQR AMKKMLJW SQCB SLGR GQ FCPRX ()X), MP AWAJCQ NCP QCAMLB. .MQR FSK?LQ A?L NCPACGTC QMSLBQ UGRF DPCOSCLAGCQ @CRUCCL 30 ?LB 20,000 )X. 8MKCL ?PC RWNGA?JJW @CRRCP ?R FC?PGLE FGEF DPCOSCLAGCQ, @SR CTCPWMLCsQ ?@GJGRW RM FC?P FGEF DPCOSCLAGCQ BCAPC?QCQ UGRF ?EC. %MEQ BCRCAR SN RM ?@MSR 40,000 )X; A?RQ,
#(!04%2 36 X 3%.3/29 3934%-3
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60,000 )X; @?RQ, 100,000 )X; ?LB BMJNFGLQ 150,000 )X, ?LB "KCPGA?L QF?B (A36:( :(70+0::04(), ? DGQF, A?L FC?P 180,000 )X. 5FMQC DPCOSCLAGCQ ?@MTC RFC FSK?L P?LEC ?PC A?JJCB ultrasound. "KNJGRSBC, MP RFC BGKCLQGML MD ? U?TC DPMK NC?I RM RPMSEF, GL QMSLB GQ FC?PB ?Q TMJSKC ?LB GQ GJJSQRP?RCB GL Figure 36.12. 5FC QMSLB U?TCQ MD JMSBCP QMSLBQ F?TC EPC?RCP ?KNJGRSBC RF?L RFMQC MD QMDRCP QMSLBQ. 'MP QMSLB, TMJSKC GQ KC?QSPCB GL BCAG@CJQ (B#). 5FC QMDRCQR QMSLB RF?R ? FSK?L A?L FC?P GQ RFC XCPM NMGLR. )SK?LQ QNC?I LMPK?JJW ?R 60 BCAG@CJQ.
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!@>LI< 36.13 3LRKA QO>SBIP QEOLRDE QEB LRQBO B>O QL QEB JFAAIB B>O, TEF@E FP ?LRKABA LK FQP BUQBOFLO ?V QEB QVJM>KF@ JBJ?O>KB. 4EB JFAAIB B>O @LKQ>FKP QEOBB ?LKBP @>IIBA LPPF@IBP QE>Q QO>KPCBO QEB PLRKA T>SB QL QEB LS>I TFKALT, QEB BUQBOFLO ?LRKA>OV LC QEB FKKBO B>O. 4EB LOD>K LC #LOQF, TEF@E FP QEB LOD>K LC PLRKA QO>KPAR@QFLK, IFBP FKPFAB QEB @L@EIB>. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ,>OP #EFQQH>, !UBI "OL@HJ>KK)
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7G@P?RGLE M@HCARQ, QSAF ?Q TMA?J AMPBQ, APC?RC QMSLB U?TCQ MP NPCQQSPC U?TCQ GL RFC ?GP. 8FCL RFCQC NPCQQSPC U?TCQ PC?AF RFC C?P, RFC C?P RP?LQBSACQ RFGQ KCAF?LGA?J QRGKSJSQ (NPCQQSPC U?TC) GLRM ? LCPTC GKNSJQC (CJCARPGA?J QGEL?J) RF?R RFC @P?GL NCPACGTCQ ?Q QMSLB. 5FC NPCQQSPC U?TCQ QRPGIC RFC RWKN?LSK, A?SQGLE GR RM TG@P?RC. 5FC KCAF?LGA?J CLCPEW DPMK RFC KMTGLE RWKN?LSK RP?LQKGRQ RFC TG@P?RGMLQ RM RFC RFPCC @MLCQ MD RFC KGBBJC C?P. 5FC QR?NCQ RP?LQKGRQ RFC TG@P?RGMLQ RM ? RFGL BG?NFP?EK A?JJCB RFC oval window, UFGAF GQ RFC MSRCPKMQR QRPSARSPC MD RFC inner ear. 5FC QRPSARSPCQ MD RFC GLLCP C?P ?PC DMSLB GL RFC labyrinth, ? @MLW, FMJJMU QRPSARSPC RF?R GQ RFC KMQR GLRCPGMP NMPRGML MD RFC C?P. )CPC, RFC CLCPEW DPMK RFC QMSLB U?TC GQ RP?LQDCPPCB DPMK RFC QR?NCQ RFPMSEF RFC DJCVG@JC MT?J UGLBMU ?LB RM RFC DJSGB MD RFC AMAFJC?. 5FC TG@P?RGMLQ MD RFC MT?J UGLBMU APC?RC NPCQQSPC U?TCQ GL RFC DJSGB (NCPGJWKNF) GLQGBC RFC AMAFJC?. 5FC cochlea GQ ? UFMPJCB QRPSARSPC, JGIC RFC QFCJJ MD ? QL?GJ, ?LB GR AMLR?GLQ PCACNRMPQ DMP RP?LQBSARGML MD RFC KCAF?LGA?J U?TC GLRM ?L CJCARPGA?J QGEL?J (?Q GJJSQRP?RCB GL Figure 36.14). *LQGBC RFC AMAFJC?, RFC basilar membrane GQ ? KCAF?LGA?J ?L?JWXCP RF?R PSLQ RFC JCLERF MD RFC AMAFJC?, ASPJGLE RMU?PB RFC AMAFJC?sQ ACLRCP. 5FC KCAF?LGA?J NPMNCPRGCQ MD RFC @?QGJ?P KCK@P?LC AF?LEC ?JMLE GRQ JCLERF, QSAF RF?R GR GQ RFGAICP, R?SRCP, ?LB L?PPMUCP ?R RFC MSRQGBC MD RFC UFMPJ (UFCPC RFC AMAFJC? GQ J?PECQR), ?LB RFGLLCP, DJMNNGCP, ?LB @PM?BCP RMU?PB RFC ?NCV, MP ACLRCP, MD RFC UFMPJ (UFCPC RFC AMAFJC? GQ QK?JJCQR). %GDDCPCLR PCEGMLQ MD RFC @?QGJ?P KCK@P?LC TG@P?RC ?AAMPBGLE RM RFC DPCOSCLAW MD RFC QMSLB U?TC AMLBSARCB RFPMSEF RFC DJSGB GL RFC AMAFJC?. 'MP RFCQC PC?QMLQ, RFC DJSGB-DGJJCB AMAFJC? BCRCARQ BGDDCPCLR U?TC DPCOSCLAGCQ (NGRAFCQ) ?R BGDDCPCLR PCEGMLQ MD RFC KCK@P?LC. 8FCL RFC QMSLB U?TCQ GL RFC AMAFJC?P DJSGB AMLR?AR RFC @?QGJ?P KCK@P?LC, GR DJCVCQ @?AI ?LB DMPRF GL ? U?TC-JGIC D?QFGML. "@MTC RFC @?QGJ?P KCK@P?LC GQ RFC tectorial membrane.
#(!04%2 36 X 3%.3/29 3934%-3
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organ of Corti (QNGP?J MPE?L). *R GQ AMKNMQCB MD F?GP ACJJQ FCJB GL NJ?AC ?@MTC RFC @?QGJ?P KCK@P?LC JGIC DJMUCPQ NPMHCARGLE SN DPMK QMGJ, UGRF RFCGP CVNMQCB QFMPR, F?GPJGIC stereocilia AMLR?ARGLE MP CK@CBBCB GL RFC RCARMPG?J KCK@P?LC ?@MTC RFCK. 5FC GLLCP F?GP ACJJQ ?PC RFC NPGK?PW ?SBGRMPW PCACNRMPQ ?LB CVGQR GL ? QGLEJC PMU, LSK@CPGLE ?NNPMVGK?RCJW 3,500. 5FC QRCPCMAGJG? DPMK GLLCP F?GP ACJJQ CVRCLB GLRM QK?JJ BGKNJCQ ML RFC RCARMPG?J KCK@P?LCsQ JMUCP QSPD?AC. 5FC MSRCP F?GP ACJJQ ?PC ?PP?LECB GL RFPCC MP DMSP PMUQ. 5FCW LSK@CP ?NNPMVGK?RCJW 12,000, ?LB RFCW DSLARGML RM DGLC RSLC GLAMKGLE QMSLB U?TCQ. 5FC JMLECP QRCPCMAGJG? RF?R NPMHCAR DPMK RFC MSRCP F?GP ACJJQ ?ARS?JJW ?RR?AF RM RFC RCARMPG?J KCK@P?LC. "JJ MD RFC QRCPCMAGJG? ?PC KCAF?LMPCACNRMPQ, ?LB UFCL @CLR @W TG@P?RGMLQ RFCW
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#(!04%2 36 X 3%.3/29 3934%-3
PCQNMLB @W MNCLGLE ? E?RCB GML AF?LLCJ (PCDCP RM m10265 (http://cnx.org/content/m10265/latest/#figch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tQF?PNCLGLEu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tQF?PNCLGLEu @SGJR GL. 5FC @P?GL A?L QCLB QGEL?JQ @?AI RM RFC AMAFJC?, PCQSJRGLE GL ? AF?LEC MD JCLERF GL RFC MSRCP F?GP ACJJQ, QF?PNCLGLE MP B?KNCLGLE RFC F?GP ACJJQs PCQNMLQC RM ACPR?GL DPCOSCLAGCQ.
8?RAF ?L animation (http://openstaxcollege.org/l/hearing) MD QMSLB CLRCPGLE RFC MSRCP C?P, KMTGLE RFPMSEF RFC C?P QRPSARSPC, QRGKSJ?RGLE AMAFJC?P LCPTC GKNSJQCQ, ?LB CTCLRS?JJW QCLBGLE QGEL?JQ RM RFC RCKNMP?J JM@C. H/-.+7 P74)+88/35FC GLLCP F?GP ACJJQ ?PC KMQR GKNMPR?LR DMP AMLTCWGLE ?SBGRMPW GLDMPK?RGML RM RFC @P?GL. "@MSR 90 NCPACLR MD RFC ?DDCPCLR LCSPMLQ A?PPW GLDMPK?RGML DPMK GLLCP F?GP ACJJQ, UGRF C?AF F?GP ACJJ QWL?NQGLE UGRF 10 MP QM LCSPMLQ. 0SRCP F?GP ACJJQ AMLLCAR RM MLJW 10 NCPACLR MD RFC ?DDCPCLR LCSPMLQ, ?LB C?AF ?DDCPCLR LCSPML GLLCPT?RCQ K?LW F?GP ACJJQ. 5FC ?DDCPCLR, @GNMJ?P LCSPMLQ RF?R AMLTCW ?SBGRMPW GLDMPK?RGML RP?TCJ DPMK RFC AMAFJC? RM RFC KCBSJJ?, RFPMSEF RFC NMLQ ?LB KGB@P?GL GL RFC @P?GLQRCK, DGL?JJW PC?AFGLE RFC NPGK?PW ?SBGRMPW AMPRCV GL RFC RCKNMP?J JM@C.
1<JK@9LC8I $E=FID8K@FE
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sQ ILMUL ?Q RFC TCQRG@SJ?P J?@WPGLRF RF?R GQ QFMUL GL Figure 36.15. 5FC SRPGAJC ?LB Q?AASJC PCQNMLB RM ?AACJCP?RGML GL ? QRP?GEFR JGLC, QSAF ?Q EP?TGRW. 5FC PMSEFJW 30,000 F?GP ACJJQ GL RFC SRPGAJC ?LB 16,000 F?GP ACJJQ GL RFC Q?AASJC JGC @CJMU ? ECJ?RGLMSQ J?WCP, UGRF RFCGP QRCPCMAGJG? NPMHCARGLE GLRM RFC ECJ?RGL. &K@CBBCB GL RFGQ ECJ?RGL ?PC A?JAGSK A?P@ML?RC APWQR?JQqJGIC RGLW PMAIQ. 8FCL RFC FC?B GQ RGJRCB, RFC APWQR?JQ AMLRGLSC RM @C NSJJCB QRP?GEFR BMUL @W EP?TGRW, @SR RFC LCU ?LEJC MD RFC FC?B A?SQCQ RFC ECJ?RGL
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RM QFGDR, RFCPC@W @CLBGLE RFC QRCPCMAGJG?. 5FC @CLBGLE MD RFC QRCPCMAGJG? QRGKSJ?RCQ RFC LCSPMLQ, ?LB RFCW QGEL?J RM RFC @P?GL RF?R RFC FC?B GQ RGJRCB, ?JJMUGLE RFC K?GLRCL?LAC MD @?J?LAC. *R GQ RFC TCQRG@SJ?P @P?LAF MD RFC TCQRG@SJMAMAFJC?P AP?LG?J LCPTC RF?R BC?JQ UGRF @?J?LAC.
!@>LI< 36.15 4EB PQOR@QROB LC QEB SBPQF?RI>O I>?VOFKQE FP PELTK. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .)()
5FC DJSGB-DGJJCB semicircular canals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igure 36.15. 8FCL RFC @P?GL NPMACQQCQ GLNSR DPMK ?JJ RFPCC A?L?JQ RMECRFCP, GR A?L BCRCAR ?LESJ?P ?AACJCP?RGML MP BCACJCP?RGML GL RFPCC BGKCLQGMLQ. 8FCL RFC FC?B RSPLQ, RFC DJSGB GL RFC A?L?JQ QFGDRQ, RFCPC@W @CLBGLE QRCPCMAGJG? ?LB QCLBGLE QGEL?JQ RM RFC @P?GL. 6NML ACQQ?RGML ?AACJCP?RGLE MP BCACJCP?RGLEqMP HSQR KMTGLEqRFC KMTCKCLR MD RFC DJSGB UGRFGL RFC A?L?JQ QJMUQ MP QRMNQ. 'MP CV?KNJC, GK?EGLC FMJBGLE ? EJ?QQ MD U?RCP. 8FCL KMTGLE DMPU?PB, U?RCP K?W QNJ?QF @?AIU?PBQ MLRM RFC F?LB, ?LB UFCL KMRGML F?Q QRMNNCB, U?RCP K?W QNJ?QF DMPU?PB MLRM RFC DGLECPQ. 8FGJC GL KMRGML, RFC U?RCP QCRRJCQ GL RFC EJ?QQ ?LB BMCQ LMR QNJ?QF. /MRC RF?R RFC A?L?JQ ?PC LMR QCLQGRGTC RM TCJMAGRW GRQCJD, @SR RM AF?LECQ GL TCJMAGRW, QM KMTGLE DMPU?PB ?R 60KNF UGRF WMSP CWCQ AJMQCB UMSJB LMR EGTC RFC QCLQ?RGML MD KMTCKCLR, @SR QSBBCLJW ?AACJCP?RGLE MP @P?IGLE UMSJB QRGKSJ?RC RFC PCACNRMPQ. H/-.+7 P74)+88/3)?GP ACJJQ DPMK RFC SRPGAJC, Q?AASJC, ?LB QCKGAGPASJ?P A?L?JQ ?JQM AMKKSLGA?RC RFPMSEF @GNMJ?P LCSPMLQ RM RFC AMAFJC?P LSAJCSQ GL RFC KCBSJJ?. $MAFJC?P LCSPMLQ QCLB BCQACLBGLE NPMHCARGMLQ RM RFC QNGL?J AMPB ?LB ?QACLBGLE NPMHCARGMLQ RM RFC NMLQ, RF?J?KSQ, ?LB ACPC@CJJSK. $MLLCARGMLQ RM RFC ACPC@CJJSK ?PC GKNMPR?LR DMP AMMPBGL?RCB KMTCKCLRQ. 5FCPC ?PC ?JQM NPMHCARGMLQ RM RFC RCKNMP?J AMPRCV, UFGAF ?AAMSLR DMP DCCJGLEQ MD BGXXGLCQQ; NPMHCARGMLQ RM ?SRMLMKGA LCPTMSQ QWQRCK ?PC?Q GL RFC @P?GLQRCK, UFGAF ?AAMSLR DMP KMRGML QGAILCQQ; ?LB NPMHCARGMLQ RM RFC NPGK?PW QMK?RMQCLQMPW AMPRCV, UFGAF KMLGRMPQ QS@HCARGTC KC?QSPCKCLRQ MD RFC CVRCPL?J UMPJB ?LB QCJD-KMTCKCLR. 1CMNJC UGRF JCQGMLQ GL RFC TCQRG@SJ?P ?PC? MD RFC QMK?RMQCLQMPW AMPRCV QCC TCPRGA?J M@HCARQ GL RFC UMPJB ?Q @CGLE RGJRCB. 'GL?JJW, RFC TCQRG@SJ?P QGEL?JQ NPMHCAR RM ACPR?GL MNRGA KSQAJCQ RM AMMPBGL?RC CWC ?LB FC?B KMTCKCLRQ.
$JGAI RFPMSEF RFGQ interactive tutorial (http://openstaxcollege.org/l/ear_anatomy) RM PCTGCU RFC N?PRQ MD RFC C?P ?LB FMU RFCW DSLARGML RM NPMACQQ QMSLB.
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#(!04%2 36 X 3%.3/29 3934%-3
36.5 R 1@J@FE
#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL FMU CJCARPMK?ELCRGA U?TCQ BGDDCPQ DPMK QMSLB U?TCQ z 5P?AC RFC N?RF MD JGEFR RFPMSEF RFC CWC RM RFC NMGLR MD RFC MNRGA LCPTC z &VNJ?GL RMLGA ?ARGTGRW ?Q GR GQ K?LGDCQRCB GL NFMRMPCACNRMPQ GL RFC PCRGL? Vision GQ RFC ?@GJGRW RM BCRCAR JGEFR N?RRCPLQ DPMK RFC MSRQGBC CLTGPMLKCLR ?LB GLRCPNPCR RFCK GLRM GK?ECQ. "LGK?JQ ?PC @MK@?PBCB UGRF QCLQMPW GLDMPK?RGML, ?LB RFC QFCCP TMJSKC MD TGQS?J GLDMPK?RGML A?L @C NPM@JCK?RGA. 'MPRSL?RCJW, RFC TGQS?J QWQRCKQ MD QNCAGCQ F?TC CTMJTCB RM ?RRCLB RM RFC KMQR-GKNMPR?LR QRGKSJG. 5FC GKNMPR?LAC MD TGQGML RM FSK?LQ GQ DSPRFCP QS@QR?LRG?RCB @W RFC D?AR RF?R ?@MSR MLC-RFGPB MD RFC FSK?L ACPC@P?J AMPRCV GQ BCBGA?RCB RM ?L?JWXGLE ?LB NCPACGTGLE TGQS?J GLDMPK?RGML.
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"Q UGRF ?SBGRMPW QRGKSJG, JGEFR RP?TCJQ GL U?TCQ. 5FC AMKNPCQQGML U?TCQ RF?R AMKNMQC QMSLB KSQR RP?TCJ GL ? KCBGSKq? E?Q, ? JGOSGB, MP ? QMJGB. *L AMLRP?QR, JGEFR GQ AMKNMQCB MD CJCARPMK?ELCRGA U?TCQ ?LB LCCBQ LM KCBGSK; JGEFR A?L RP?TCJ GL ? T?ASSK (Figure 36.16). 5FC @CF?TGMP MD JGEFR A?L @C BGQASQQCB GL RCPKQ MD RFC @CF?TGMP MD U?TCQ ?LB ?JQM GL RCPKQ MD RFC @CF?TGMP MD RFC DSLB?KCLR?J SLGR MD JGEFRq? N?AICR MD CJCARPMK?ELCRGA P?BG?RGML A?JJCB ? NFMRML. " EJ?LAC ?R RFC CJCARPMK?ELCRGA QNCARPSK QFMUQ RF?R TGQG@JC JGEFR DMP FSK?LQ GQ HSQR ? QK?JJ QJGAC MD RFC CLRGPC QNCARPSK, UFGAF GLAJSBCQ P?BG?RGML RF?R UC A?LLMR QCC ?Q JGEFR @CA?SQC GR GQ @CJMU RFC DPCOSCLAW MD TGQG@JC PCB JGEFR ?LB ?@MTC RFC DPCOSCLAW MD TGQG@JC TGMJCR JGEFR. $CPR?GL T?PG?@JCQ ?PC GKNMPR?LR UFCL BGQASQQGLE NCPACNRGML MD JGEFR. 8?TCJCLERF (UFGAF T?PGCQ GLTCPQCJW UGRF DPCOSCLAW) K?LGDCQRQ GRQCJD ?Q FSC. -GEFR ?R RFC PCB CLB MD RFC TGQG@JC QNCARPSK F?Q JMLECP U?TCJCLERFQ (?LB GQ JMUCP DPCOSCLAW), UFGJC JGEFR ?R RFC TGMJCR CLB F?Q QFMPRCP U?TCJCLERFQ (?LB GQ FGEFCP DPCOSCLAW). 5FC U?TCJCLERF MD JGEFR GQ CVNPCQQCB GL L?LMKCRCPQ (LK); MLC L?LMKCRCP GQ MLC @GJJGMLRF MD ? KCRCP. )SK?LQ NCPACGTC JGEFR RF?R P?LECQ @CRUCCL ?NNPMVGK?RCJW 380 LK ?LB 740 LK. 4MKC MRFCP ?LGK?JQ, RFMSEF, A?L BCRCAR U?TCJCLERFQ MSRQGBC MD RFC FSK?L P?LEC. 'MP CV?KNJC, @CCQ QCC LC?P-SJRP?TGMJCR JGEFR GL MPBCP RM JMA?RC LCAR?P ESGBCQ ML DJMUCPQ, ?LB QMKC LML-?TG?L PCNRGJCQ QCLQC GLDP?PCB JGEFR (FC?R RF?R NPCW EGTCQ MDD).
!@>LI< 36.16 )K QEB BIB@QOLJ>DKBQF@ PMB@QORJ, SFPF?IB IFDEQ IFBP ?BQTBBK 380 KJ >KA 740 KJ. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .!3!)
8?TC ?KNJGRSBC GQ NCPACGTCB ?Q JSKGLMSQ GLRCLQGRW, MP @PGEFRLCQQ. 5FC QR?LB?PB SLGR MD GLRCLQGRW MD JGEFR GQ RFC candela, UFGAF GQ ?NNPMVGK?RCJW RFC JSKGLMSQ GLRCLQGRW MD ? MLC AMKKML A?LBJC. -GEFR U?TCQ RP?TCJ 299,792 IK NCP QCAMLB GL ? T?ASSK, (?LB QMKCUF?R QJMUCP GL T?PGMSQ KCBG? QSAF ?Q ?GP ?LB U?RCP), ?LB RFMQC U?TCQ ?PPGTC ?R RFC CWC ?Q JMLE (PCB), KCBGSK (EPCCL), ?LB QFMPR (@JSC) U?TCQ. 8F?R GQ RCPKCB tUFGRC JGEFRu GQ JGEFR RF?R GQ NCPACGTCB ?Q UFGRC @W RFC FSK?L CWC. 5FGQ CDDCAR GQ NPMBSACB @W JGEFR RF?R QRGKSJ?RCQ COS?JJW RFC AMJMP PCACNRMPQ GL RFC FSK?L CWC. 5FC ?NN?PCLR AMJMP MD ?L M@HCAR GQ RFC AMJMP (MP AMJMPQ) RF?R RFC M@HCAR PCDJCARQ. 5FSQ ? PCB M@HCAR PCDJCARQ RFC PCB U?TCJCLERFQ GL KGVCB (UFGRC) JGEFR ?LB ?@QMP@Q ?JJ MRFCP U?TCJCLERFQ MD JGEFR.
#(!04%2 36 X 3%.3/29 3934%-3
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5FC NFMRMPCACNRGTC ACJJQ MD RFC CWC, UFCPC RP?LQBSARGML MD JGEFR RM LCPTMSQ GKNSJQCQ MAASPQ, ?PC JMA?RCB GL RFC retina (QFMUL GL Figure 36.17) ML RFC GLLCP QSPD?AC MD RFC @?AI MD RFC CWC. #SR JGEFR BMCQ LMR GKNGLEC ML RFC PCRGL? SL?JRCPCB. *R N?QQCQ RFPMSEF MRFCP J?WCPQ RF?R NPMACQQ GR QM RF?R GR A?L @C GLRCPNPCRCB @W RFC PCRGL? (Figure 36.17b). 5FC cornea, RFC DPMLR RP?LQN?PCLR J?WCP MD RFC CWC, ?LB RFC APWQR?JJGLC lens, ? RP?LQN?PCLR AMLTCV QRPSARSPC @CFGLB RFC AMPLC?, @MRF PCDP?AR (@CLB) JGEFR RM DMASQ RFC GK?EC ML RFC PCRGL?. 5FC iris, UFGAF GQ AMLQNGASMSQ ?Q RFC AMJMPCB N?PR MD RFC CWC, GQ ? AGPASJ?P KSQASJ?P PGLE JWGLE @CRUCCL RFC JCLQ ?LB AMPLC? RF?R PCESJ?RCQ RFC ?KMSLR MD JGEFR CLRCPGLE RFC CWC. *L AMLBGRGMLQ MD FGEF ?K@GCLR JGEFR, RFC GPGQ AMLRP?ARQ, PCBSAGLE RFC QGXC MD RFC NSNGJ ?R GRQ ACLRCP. *L AMLBGRGMLQ MD JMU JGEFR, RFC GPGQ PCJ?VCQ ?LB RFC NSNGJ CLJ?PECQ.
!@>LI< 36.17 (>) 4EB ERJ>K BVB FP PELTK FK @OLPP PB@QFLK. (?) ! ?ILTRM PELTP QEB I>VBOP LC QEB OBQFK>.
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presbyopia PCQSJRQ. 1PCQ@WMNG? MAASPQ @CA?SQC RFC GK?EC DMASQCQ @CFGLB RFC PCRGL?. 1PCQ@WMNG? GQ ? BCDGAGR QGKGJ?P RM ? BGDDCPCLR RWNC MD D?PQGEFRCBLCQQ A?JJCB hyperopia A?SQCB @W ?L CWC@?JJ RF?R GQ RMM QFMPR. 'MP @MRF BCDCARQ, GK?ECQ GL RFC BGQR?LAC ?PC AJC?P @SR GK?ECQ LC?P@W ?PC @JSPPW. Myopia (LC?PQGEFRCBLCQQ) MAASPQ UFCL ?L CWC@?JJ GQ CJMLE?RCB ?LB RFC GK?EC DMASQ D?JJQ GL DPMLR MD RFC PCRGL?. *L RFGQ A?QC, GK?ECQ GL RFC BGQR?LAC ?PC @JSPPW @SR GK?ECQ LC?P@W ?PC AJC?P. 5FCPC ?PC RUM RWNCQ MD NFMRMPCACNRMPQ GL RFC PCRGL?: rods ?LB cones, L?KCB DMP RFCGP ECLCP?J ?NNC?P?LAC ?Q GJJSQRP?RCB GL Figure 36.18. 3MBQ ?PC QRPMLEJW NFMRMQCLQGRGTC ?LB ?PC JMA?RCB GL RFC MSRCP CBECQ MD RFC PCRGL?. 5FCW BCRCAR BGK JGEFR ?LB ?PC SQCB NPGK?PGJW DMP NCPGNFCP?J ?LB LGEFRRGKC TGQGML. $MLCQ ?PC UC?IJW NFMRMQCLQGRGTC ?LB ?PC JMA?RCB LC?P RFC ACLRCP MD RFC PCRGL?. 5FCW PCQNMLB RM @PGEFR JGEFR, ?LB RFCGP NPGK?PW PMJC GQ GL B?WRGKC, AMJMP TGQGML.
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#(!04%2 36 X 3%.3/29 3934%-3
!@>LI< 36.18 2LAP >KA @LKBP >OB MELQLOB@BMQLOP FK QEB OBQFK>. 2LAP OBPMLKA FK ILT IFDEQ >KA @>K ABQB@Q LKIV PE>ABP LC DO>V. #LKBP OBPMLKA FK FKQBKPB IFDEQ >KA >OB OBPMLKPF?IB CLO @LILO SFPFLK. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V 0FLQO 3IFT>)
5FC fovea GQ RFC PCEGML GL RFC ACLRCP @?AI MD RFC CWC RF?R GQ PCQNMLQG@JC DMP ?ASRC TGQGML. 5FC DMTC? F?Q ? FGEF BCLQGRW MD AMLCQ. 8FCL WMS @PGLE WMSP E?XC RM ?L M@HCAR RM CV?KGLC GR GLRCLRJW GL @PGEFR JGEFR, RFC CWCQ MPGCLR QM RF?R RFC M@HCARsQ GK?EC D?JJQ ML RFC DMTC?. )MUCTCP, UFCL JMMIGLE ?R ? QR?P GL RFC LGEFR QIW MP MRFCP M@HCAR GL BGK JGEFR, RFC M@HCAR A?L @C @CRRCP TGCUCB @W RFC NCPGNFCP?J TGQGML @CA?SQC GR GQ RFC PMBQ ?R RFC CBECQ MD RFC PCRGL?, P?RFCP RF?L RFC AMLCQ ?R RFC ACLRCP, RF?R MNCP?RC @CRRCP GL JMU JGEFR. *L FSK?LQ, AMLCQ D?P MSRLSK@CP PMBQ GL RFC DMTC?.
3CTGCU RFC anatomical structure (http://openstaxcollege.org/l/eye_diagram) MD RFC CWC, AJGAIGLE ML C?AF N?PR RM NP?ARGAC GBCLRGDGA?RGML.
/I8EJ;L:K@FE F= '@>?K
5FC PMBQ ?LB AMLCQ ?PC RFC QGRC MD RP?LQBSARGML MD JGEFR RM ? LCSP?J QGEL?J. #MRF PMBQ ?LB AMLCQ AMLR?GL NFMRMNGEKCLRQ. *L TCPRC@P?RCQ, RFC K?GL NFMRMNGEKCLR, rhodopsin, F?Q RUM K?GL N?PRQ Figure 36.19): ?L MNQGL, UFGAF GQ ? KCK@P?LC NPMRCGL (GL RFC DMPK MD ? AJSQRCP MD i-FCJGACQ RF?R QN?L RFC KCK@P?LC), ?LB PCRGL?Jq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igure 36.20).
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!@>LI< 36.19 (>) 2ELALMPFK, QEB MELQLOB@BMQLO FK SBOQB?O>QBP, E>P QTL M>OQP: QEB QO>KP-JBJ?O>KB MOLQBFK LMPFK, >KA OBQFK>I. 7EBK IFDEQ PQOFHBP OBQFK>I, FQ @E>KDBP PE>MB COLJ (?) > *0: QL > ;9(5: CLOJ. 4EB PFDK>I FP M>PPBA QL > '-MOLQBFK @>IIBA QO>KPAR@FK, QOFDDBOFKD > PBOFBP LC ALTKPQOB>J BSBKQP.
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#(!04%2 36 X 3%.3/29 3934%-3
!@>LI< 36.20 7EBK IFDEQ PQOFHBP OELALMPFK, QEB '-MOLQBFK QO>KPAR@FK FP >@QFS>QBA, TEF@E FK QROK >@QFS>QBP MELPMELAFBPQBO>PB. 0ELPMELAFBPQBO>PB @LKSBOQP @'-0 QL '-0, QEBOB?V @ILPFKD PLAFRJ @E>KKBIP. !P > OBPRIQ, QEB JBJ?O>KB ?B@LJBP EVMBOMLI>OFWBA. 4EB EVMBOMLI>OFWBA JBJ?O>KB ALBP KLQ OBIB>PB DIRQ>J>QB QL QEB ?FMLI>O @BII.
"7/).742'9/) C4*/35FCPC ?PC RFPCC RWNCQ MD AMLCQ (UGRF BGDDCPCLR NFMRMNQGLQ), ?LB RFCW BGDDCP GL RFC U?TCJCLERF RM UFGAF RFCW ?PC KMQR PCQNMLQGTC, ?Q QFMUL GL Figure 36.21. 4MKC AMLCQ ?PC K?VGK?JJW PCQNMLQGTC RM QFMPR JGEFR U?TCQ MD 420 LK, QM RFCW ?PC A?JJCB 4 AMLCQ (t4u DMP tQFMPRu); MRFCPQ PCQNMLB K?VGK?JJW RM U?TCQ MD 530 LK (. AMLCQ, DMP tKCBGSKu); ? RFGPB EPMSN PCQNMLBQ K?VGK?JJW RM JGEFR MD JMLECP U?TCJCLERFQ, ?R 560 LK (-, MP tJMLEu AMLCQ). 8GRF MLJW MLC RWNC MD AMLC, AMJMP TGQGML UMSJB LMR @C NMQQG@JC, ?LB ? RUM-AMLC (BGAFPMK?RGA) QWQRCK F?Q JGKGR?RGMLQ. 1PGK?RCQ SQC ? RFPCC-AMLC (RPGAFPMK?RGA) QWQRCK, PCQSJRGLE GL DSJJ AMJMP TGQGML. 5FC AMJMP UC NCPACGTC GQ ? PCQSJR MD RFC P?RGM MD ?ARGTGRW MD MSP RFPCC RWNCQ MD AMLCQ. 5FC AMJMPQ MD RFC TGQS?J QNCARPSK, PSLLGLE DPMK JMLE-U?TCJCLERF JGEFR RM QFMPR, ?PC PCB (700 LK), MP?LEC (600 LK), WCJJMU (565 LK), EPCCL (497 LK), @JSC (470 LK), GLBGEM (450 LK), ?LB TGMJCR (425 LK). )SK?LQ F?TC TCPW QCLQGRGTC NCPACNRGML MD AMJMP ?LB A?L BGQRGLESGQF ?@MSR 500 JCTCJQ MD @PGEFRLCQQ, 200 BGDDCPCLR FSCQ, ?LB 20 QRCNQ MD Q?RSP?RGML, MP ?@MSR 2 KGJJGML BGQRGLAR AMJMPQ.
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!@>LI< 36.21 (RJ>K OLA @BIIP >KA QEB AFCCBOBKQ QVMBP LC @LKB @BIIP B>@E E>SB >K LMQFJ>I T>SBIBKDQE. (LTBSBO, QEBOB FP @LKPFABO>?IB LSBOI>M FK QEB T>SBIBKDQEP LC IFDEQ ABQB@QBA.
+9/3'1 P74)+88/37GQS?J QGEL?JQ JC?TC RFC AMLCQ ?LB PMBQ, RP?TCJ RM RFC @GNMJ?P ACJJQ, ?LB RFCL RM E?LEJGML ACJJQ. " J?PEC BCEPCC MD NPMACQQGLE MD TGQS?J GLDMPK?RGML MAASPQ GL RFC PCRGL? GRQCJD, @CDMPC TGQS?J GLDMPK?RGML GQ QCLR RM RFC @P?GL. 1FMRMPCACNRMPQ GL RFC PCRGL? AMLRGLSMSQJW SLBCPEM tonic activity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tRPGAIu WMSP PCRGL? ?LB @P?GL ?@MSR RFC AMJMPQ WMS ?PC M@QCPTGLE GL WMSP TGQS?J DGCJB. -MMI DGVCBJW ?R Figure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tMNNMLCLRu AMJMPQ: PCB, UFGRC, ?LB @JSC, PCQNCARGTCJW, GL RFC TGQS?J DGCJB. 0LAC RFC E?LEJGML ACJJQ PCRSPL RM RFCGP @?QCJGLC ?ARGTGRW QR?RC, RFC D?JQC NCPACNRGML MD AMJMP UGJJ BGQ?NNC?P.
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!@>LI< 36.22 6FBT QEFP CI>D QL RKABOPQ>KA ELT OBQFK>I MOL@BPPFKD TLOHP. 3Q>OB >Q QEB @BKQBO LC QEB CI>D (FKAF@>QBA ?V QEB TEFQB ALQ) CLO 45 PB@LKAP, >KA QEBK NRF@HIV ILLH >Q > TEFQB ?>@HDOLRKA, KLQF@FKD ELT @LILOP >MMB>O.
#@>?
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tAPMQQGLEu) DMSLB ?R RFC @?QC MD RFC @P?GL ?LB ?JJMUQ SQ RM AMMPBGL?RC GLDMPK?RGML DPMK @MRF CWCQ. 0LAC GL RFC @P?GL, TGQS?J GLDMPK?RGML GQ NPMACQQCB GL QCTCP?J NJ?ACQ, ?LB GRQ PMSRCQ PCDJCAR RFC AMKNJCVGRW ?LB GKNMPR?LAC MD TGQS?J GLDMPK?RGML RM FSK?LQ ?LB MRFCP ?LGK?JQ. 0LC PMSRC R?ICQ RFC QGEL?JQ RM RFC RF?J?KSQ, UFGAF QCPTCQ ?Q RFC PMSRGLE QR?RGML DMP ?JJ GLAMKGLE QCLQMPW GKNSJQCQ CVACNR MJD?ARGML. *L RFC RF?J?KSQ, RFC K?ELMACJJSJ?P ?LB N?PTMACJJSJ?P BGQRGLARGMLQ PCK?GL GLR?AR, ?LB RFCPC ?PC BGDDCPCLR J?WCPQ MD RFC RF?J?KSQ BCBGA?RCB RM C?AF. 8FCL TGQS?J QGEL?JQ JC?TC RFC RF?J?KSQ, RFCW RP?TCJ RM RFC NPGK?PW TGQS?J AMPRCV ?R RFC PC?P MD RFC @P?GL. 'PMK RFC TGQS?J AMPRCV, RFC TGQS?J QGEL?JQ RP?TCJ GL RUM BGPCARGMLQ. 0LC QRPC?K RF?R NPMHCARQ RM RFC N?PGCR?J JM@C, GL RFC QGBC MD RFC @P?GL, A?PPGCQ K?ELMACJJSJ?P (tUFCPCu) GLDMPK?RGML. " QCAMLB QRPC?K NPMHCARQ RM RFC RCKNMP?J JM@C ?LB A?PPGCQ @MRF K?ELMACJJSJ?P (tUFCPCu) ?LB N?PTMACJJSJ?P (tUF?Ru) GLDMPK?RGML. "LMRFCP GKNMPR?LR TGQS?J PMSRC GQ ? N?RFU?W DPMK RFC PCRGL? RM RFC superior colliculus GL RFC KGB@P?GL, UFCPC CWC KMTCKCLRQ ?PC AMMPBGL?RCB ?LB GLRCEP?RCB UGRF ?SBGRMPW GLDMPK?RGML. 'GL?JJW, RFCPC GQ RFC N?RFU?W DPMK RFC PCRGL? RM RFC suprachiasmatic nucleus (4$/) MD RFC FWNMRF?J?KSQ. 5FC 4$/ GQ ? AJSQRCP MD ACJJQ RF?R GQ AMLQGBCPCB RM @C RFC @MBWsQ GLRCPL?J AJMAI, UFGAF AMLRPMJQ MSP circadian (B?WJMLE) AWAJC. 5FC 4$/ QCLBQ GLDMPK?RGML RM RFC NGLC?J EJ?LB, UFGAF GQ GKNMPR?LR GL QJCCN/U?IC N?RRCPLQ ?LB ?LLS?J AWAJCQ.
7GCU RFGQ interactive presentation (http://openstaxcollege.org/l/sense_of_sight) RM PCTGCU UF?R WMS F?TC JC?PLCB ?@MSR FMU TGQGML DSLARGMLQ.
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& 4 / -(.
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
#(!04%2 36 X 3%.3/29 3934%-3
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"L ?LGK?JsQ CLBMAPGLC QWQRCK AMLRPMJQ @MBW NPMACQQCQ RFPMSEF RFC NPMBSARGML, QCAPCRGML, ?LB PCESJ?RGML MD FMPKMLCQ, UFGAF QCPTC ?Q AFCKGA?J tKCQQCLECPQu DSLARGMLGLE GL ACJJSJ?P ?LB MPE?L ?ARGTGRW ?LB, SJRGK?RCJW, K?GLR?GLGLE RFC @MBWsQ FMKCMQR?QGQ. 5FC CLBMAPGLC QWQRCK NJ?WQ ? PMJC GL EPMURF, KCR?@MJGQK, ?LB QCVS?J BCTCJMNKCLR. *L FSK?LQ, AMKKML CLBMAPGLC QWQRCK BGQC?QCQ GLAJSBC RFWPMGB BGQC?QC ?LB BG?@CRCQ KCJJGRSQ. *L MPE?LGQKQ RF?R SLBCPEM KCR?KMPNFMQGQ, RFC NPMACQQ GQ AMLRPMJJCB @W RFC CLBMAPGLC QWQRCK. 5FC RP?LQDMPK?RGML DPMK R?BNMJC RM DPME, DMP CV?KNJC, GQ AMKNJCV ?LB LS?LACB RM ?B?NR RM QNCAGDGA CLTGPMLKCLRQ ?LB CAMJMEGA?J AGPASKQR?LACQ.
37.1 R /PG<J F= #FIDFE<J

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z -GQR RFC BGDDCPCLR RWNCQ MD FMPKMLCQ z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
1064
#(!04%2 37 X 4(% %.$/#2).% 3934%-
NPMRCGLQ) FMPKMLCQ. 0LC MD RFC ICW BGQRGLESGQFGLE DC?RSPCQ MD JGNGB-BCPGTCB FMPKMLCQ GQ RF?R RFCW A?L BGDDSQC ?APMQQ NJ?QK? KCK@P?LCQ UFCPC?Q RFC ?KGLM ?AGB-BCPGTCB ?LB NCNRGBC FMPKMLCQ A?LLMR.
'@G@;-D<I@M<; #FIDFE<J (FI '@G@;-JFCL9C< #FIDFE<J)

.MQR lipid hormones ?PC BCPGTCB DPMK AFMJCQRCPMJ ?LB RFSQ ?PC QRPSARSP?JJW QGKGJ?P RM GR, ?Q GJJSQRP?RCB GL Figure 37.2. 5FC NPGK?PW AJ?QQ MD JGNGB FMPKMLCQ GL FSK?LQ GQ RFC QRCPMGB FMPKMLCQ. $FCKGA?JJW, RFCQC FMPKMLCQ ?PC SQS?JJW ICRMLCQ MP ?JAMFMJQ; RFCGP AFCKGA?J L?KCQ UGJJ CLB GL t-MJu DMP ?JAMFMJQ MP t-MLCu DMP ICRMLCQ. &V?KNJCQ MD QRCPMGB FMPKMLCQ GLAJSBC CQRP?BGMJ, UFGAF GQ ?L estrogen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
!@>LI< 37.2 4EB PQOR@QROBP PELTK EBOB OBMOBPBKQ (>) @ELIBPQBOLI, MIRP QEB PQBOLFA ELOJLKBP (?) QBPQLPQBOLKB >KA (@) BPQO>AFLI.
AD@EF A:@;-D<I@M<; #FIDFE<J

5FC amino acid-derived hormones ?PC PCJ?RGTCJW QK?JJ KMJCASJCQ RF?R ?PC BCPGTCB DPMK RFC ?KGLM ?AGBQ RWPMQGLC ?LB RPWNRMNF?L, QFMUL GL Figure 37.3. *D ? FMPKMLC GQ ?KGLM ?AGB-BCPGTCB, GRQ AFCKGA?J L?KC UGJJ CLB GL t-GLCu. &V?KNJCQ MD ?KGLM ?AGB-BCPGTCB FMPKMLCQ GLAJSBC CNGLCNFPGLC ?LB LMPCNGLCNFPGLC, UFGAF ?PC QWLRFCQGXCB GL RFC KCBSJJ? MD RFC ?BPCL?J EJ?LBQ, ?LB RFWPMVGLC, UFGAF GQ NPMBSACB @W RFC RFWPMGB EJ?LB. 5FC NGLC?J EJ?LB GL RFC @P?GL K?ICQ ?LB QCAPCRCQ KCJ?RMLGL UFGAF PCESJ?RCQ QJCCN AWAJCQ.
#(!04%2 37 X 4(% %.$/#2).% 3934%-
1065
!@>LI< 37.3 (>) 4EB ELOJLKB BMFKBMEOFKB, TEF@E QOFDDBOP QEB CFDEQ-LO-CIFDEQ OBPMLKPB, FP ABOFSBA COLJ QEB >JFKL >@FA QVOLPFKB. (?) 4EB ELOJLKB JBI>QLKFK, TEF@E OBDRI>QBP @FO@>AF>K OEVQEJP, FP ABOFSBA COLJ QEB >JFKL >@FA QOVMQLME>K.
+<GK@;< #FIDFE<J
5FC QRPSARSPC MD peptide hormones GQ RF?R MD ? NMJWNCNRGBC AF?GL (AF?GL MD ?KGLM ?AGBQ). 5FC NCNRGBC FMPKMLCQ GLAJSBC KMJCASJCQ RF?R ?PC QFMPR NMJWNCNRGBC AF?GLQ, QSAF ?Q ?LRGBGSPCRGA FMPKMLC ?LB MVWRMAGL NPMBSACB GL RFC @P?GL ?LB PCJC?QCB GLRM RFC @JMMB GL RFC NMQRCPGMP NGRSGR?PW EJ?LB. 5FGQ AJ?QQ ?JQM GLAJSBCQ QK?JJ NPMRCGLQ, JGIC EPMURF FMPKMLCQ NPMBSACB @W RFC NGRSGR?PW, ?LB J?PEC EJWAMNPMRCGLQ QSAF ?Q DMJJGAJCQRGKSJ?RGLE FMPKMLC NPMBSACB @W RFC NGRSGR?PW. Figure 37.4 GJJSQRP?RCQ RFCQC NCNRGBC FMPKMLCQ. 4CAPCRCB NCNRGBCQ JGIC GLQSJGL ?PC QRMPCB UGRFGL TCQGAJCQ GL RFC ACJJQ RF?R QWLRFCQGXC RFCK. 5FCW ?PC RFCL PCJC?QCB GL PCQNMLQC RM QRGKSJG QSAF ?Q FGEF @JMMB EJSAMQC JCTCJQ GL RFC A?QC MD GLQSJGL. "KGLM ?AGBBCPGTCB ?LB NMJWNCNRGBC FMPKMLCQ ?PC U?RCP-QMJS@JC ?LB GLQMJS@JC GL JGNGBQ. 5FCQC FMPKMLCQ A?LLMR N?QQ RFPMSEF NJ?QK? KCK@P?LCQ MD ACJJQ; RFCPCDMPC, RFCGP PCACNRMPQ ?PC DMSLB ML RFC QSPD?AC MD RFC R?PECR ACJJQ.
!@>LI< 37.4 4EB PQOR@QROBP LC MBMQFAB ELOJLKBP (>) LUVQL@FK, (?) DOLTQE ELOJLKB, >KA (@) CLIIF@IBPQFJRI>QFKD ELOJLKB >OB PELTK. 4EBPB MBMQFAB ELOJLKBP >OB JR@E I>ODBO QE>K QELPB ABOFSBA COLJ @ELIBPQBOLI LO >JFKL >@FAP.
1066
#(!04%2 37 X 4(% %.$/#2).% 3934%-
E;F:I@EFCF>@JK
!K BKAL@OFKLILDFPQ FP > JBAF@>I AL@QLO TEL PMB@F>IFWBP FK QOB>QFKD AFPLOABOP LC QEB BKAL@OFKB DI>KAP, ELOJLKB PVPQBJP, >KA DIR@LPB >KA IFMFA JBQ>?LIF@ M>QET>VP. !K BKAL@OFKB PRODBLK PMB@F>IFWBP FK QEB PRODF@>I QOB>QJBKQ LC BKAL@OFKB AFPB>PBP >KA DI>KAP. 3LJB LC QEB AFPB>PBP QE>Q >OB J>K>DBA ?V BKAL@OFKLILDFPQP: AFPLOABOP LC QEB M>K@OB>P (AF>?BQBP JBIIFQRP), AFPLOABOP LC QEB MFQRFQ>OV (DFD>KQFPJ, >@OLJBD>IV, >KA MFQRFQ>OV AT>OCFPJ), AFPLOABOP LC QEB QEVOLFA DI>KA (DLFQBO >KA 'O>SBPw AFPB>PB), >KA AFPLOABOP LC QEB >AOBK>I DI>KAP (#RPEFKDwP AFPB>PB >KA !AAFPLKw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
37.2 R #FN #FIDFE<J 2FIB

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL FMU FMPKMLCQ UMPI z %GQASQQ RFC PMJC MD BGDDCPCLR RWNCQ MD FMPKMLC PCACNRMPQ )MPKMLCQ KCBG?RC AF?LECQ GL R?PECR ACJJQ @W @GLBGLE RM QNCAGDGA hormone receptors. *L RFGQ U?W, CTCL RFMSEF FMPKMLCQ AGPASJ?RC RFPMSEFMSR RFC @MBW ?LB AMKC GLRM AMLR?AR UGRF K?LW BGDDCPCLR ACJJ RWNCQ, RFCW MLJW ?DDCAR ACJJQ RF?R NMQQCQQ RFC LCACQQ?PW PCACNRMPQ. 3CACNRMPQ DMP ? QNCAGDGA FMPKMLC K?W @C DMSLB ML K?LW BGDDCPCLR ACJJQ MP K?W @C JGKGRCB RM ? QK?JJ LSK@CP MD QNCAG?JGXCB ACJJQ. 'MP CV?KNJC, RFWPMGB FMPKMLCQ ?AR ML K?LW BGDDCPCLR RGQQSC RWNCQ, QRGKSJ?RGLE KCR?@MJGA ?ARGTGRW RFPMSEFMSR RFC @MBW. $CJJQ A?L F?TC K?LW PCACNRMPQ DMP RFC Q?KC FMPKMLC @SR MDRCL ?JQM NMQQCQQ PCACNRMPQ DMP BGDDCPCLR RWNCQ MD FMPKMLCQ. 5FC LSK@CP MD PCACNRMPQ RF?R PCQNMLB RM ? FMPKMLC BCRCPKGLCQ RFC ACJJsQ QCLQGRGTGRW RM RF?R FMPKMLC, ?LB RFC PCQSJRGLE ACJJSJ?P PCQNMLQC. "BBGRGML?JJW, RFC LSK@CP MD PCACNRMPQ RF?R PCQNMLB RM ? FMPKMLC A?L AF?LEC MTCP RGKC, PCQSJRGLE GL GLAPC?QCB MP BCAPC?QCB ACJJ QCLQGRGTGRW. *L up-regulation, RFC LSK@CP MD PCACNRMPQ GLAPC?QCQ GL PCQNMLQC RM PGQGLE FMPKMLC JCTCJQ, K?IGLE RFC ACJJ KMPC QCLQGRGTC RM RFC FMPKMLC ?LB ?JJMUGLE DMP KMPC ACJJSJ?P ?ARGTGRW. 8FCL RFC LSK@CP MD PCACNRMPQ BCAPC?QCQ GL PCQNMLQC RM PGQGLE FMPKMLC JCTCJQ, A?JJCB down-regulation, ACJJSJ?P ?ARGTGRW GQ PCBSACB. 3CACNRMP @GLBGLE ?JRCPQ ACJJSJ?P ?ARGTGRW ?LB PCQSJRQ GL ?L GLAPC?QC MP BCAPC?QC GL LMPK?J @MBW NPMACQQCQ. %CNCLBGLE ML RFC JMA?RGML MD RFC NPMRCGL PCACNRMP ML RFC R?PECR ACJJ ?LB RFC AFCKGA?J QRPSARSPC MD RFC FMPKMLC, FMPKMLCQ A?L KCBG?RC AF?LECQ BGPCARJW @W @GLBGLE RM intracellular hormone receptors ?LB KMBSJ?RGLE ECLC RP?LQAPGNRGML, MP GLBGPCARJW @W @GLBGLE RM ACJJ QSPD?AC PCACNRMPQ ?LB QRGKSJ?RGLE QGEL?JGLE N?RFU?WQ.
#(!04%2 37 X 4(% %.$/#2).% 3934%-
1067
$EKI8:<CCLC8I #FIDFE< -<:<GKFIJ

-GNGB-BCPGTCB (QMJS@JC) FMPKMLCQ QSAF ?Q QRCPMGB FMPKMLCQ BGDDSQC ?APMQQ RFC KCK@P?LCQ MD RFC CLBMAPGLC ACJJ. 0LAC MSRQGBC RFC ACJJ, RFCW @GLB RM RP?LQNMPR NPMRCGLQ RF?R ICCN RFCK QMJS@JC GL RFC @JMMBQRPC?K. "R RFC R?PECR ACJJ, RFC FMPKMLCQ ?PC PCJC?QCB DPMK RFC A?PPGCP NPMRCGL ?LB BGDDSQC ?APMQQ RFC JGNGB @GJ?WCP MD RFC NJ?QK? KCK@P?LC MD ACJJQ. 5FC QRCPMGB FMPKMLCQ N?QQ RFPMSEF RFC NJ?QK? KCK@P?LC MD ? R?PECR ACJJ ?LB ?BFCPC RM GLRP?ACJJSJ?P PCACNRMPQ PCQGBGLE GL RFC AWRMNJ?QK MP GL RFC LSAJCSQ. 5FC ACJJ QGEL?JGLE N?RFU?WQ GLBSACB @W RFC QRCPMGB FMPKMLCQ PCESJ?RC QNCAGDGA ECLCQ ML RFC ACJJ'Q %/". 5FC FMPKMLCQ ?LB PCACNRMP AMKNJCV ?AR ?Q RP?LQAPGNRGML PCESJ?RMPQ @W GLAPC?QGLE MP BCAPC?QGLE RFC QWLRFCQGQ MD K3/" KMJCASJCQ MD QNCAGDGA ECLCQ. 5FGQ, GL RSPL, BCRCPKGLCQ RFC ?KMSLR MD AMPPCQNMLBGLE NPMRCGL RF?R GQ QWLRFCQGXCB @W ?JRCPGLE ECLC CVNPCQQGML. 5FGQ NPMRCGL A?L @C SQCB CGRFCP RM AF?LEC RFC QRPSARSPC MD RFC ACJJ MP RM NPMBSAC CLXWKCQ RF?R A?R?JWXC AFCKGA?J PC?ARGMLQ. *L RFGQ U?W, RFC QRCPMGB FMPKMLC PCESJ?RCQ QNCAGDGA ACJJ NPMACQQCQ ?Q GJJSQRP?RCB GL Figure 37.5.
!@>LI< 37.5 !K FKQO>@BIIRI>O KR@IB>O OB@BMQLO (.2) FP IL@>QBA FK QEB @VQLMI>PJ ?LRKA QL > EB>Q PEL@H MOLQBFK ((30). 5MLK ELOJLKB ?FKAFKD, QEB OB@BMQLO AFPPL@F>QBP COLJ QEB EB>Q PEL@H MOLQBFK >KA QO>KPIL@>QBP QL QEB KR@IBRP. )K QEB KR@IBRP, QEB ELOJLKB-OB@BMQLO @LJMIBU ?FKAP QL > $.! PBNRBK@B @>IIBA > ELOJLKB OBPMLKPB BIBJBKQ ((2%), TEF@E QOFDDBOP DBKB QO>KP@OFMQFLK >KA QO>KPI>QFLK. 4EB @LOOBPMLKAFKD MOLQBFK MOLAR@Q @>K QEBK JBAF>QB @E>KDBP FK @BII CRK@QFLK.
(B>Q PEL@H MOLQBFKP ((30) >OB PL K>JBA ?B@>RPB QEBV EBIM OBCLIA JFPCLIABA MOLQBFKP. )K OBPMLKPB QL FK@OB>PBA QBJMBO>QROB (> xEB>Q PEL@Hy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
+C8JD8 (<D9I8E< #FIDFE< -<:<GKFIJ

"KGLM ?AGB BCPGTCB FMPKMLCQ ?LB NMJWNCNRGBC FMPKMLCQ ?PC LMR JGNGB-BCPGTCB (JGNGB-QMJS@JC) ?LB RFCPCDMPC A?LLMR BGDDSQC RFPMSEF RFC NJ?QK? KCK@P?LC MD ACJJQ. -GNGB GLQMJS@JC FMPKMLCQ @GLB RM PCACNRMPQ ML RFC MSRCP QSPD?AC MD RFC NJ?QK? KCK@P?LC, TG? plasma membrane hormone receptors. 6LJGIC QRCPMGB FMPKMLCQ, JGNGB GLQMJS@JC FMPKMLCQ BM LMR BGPCARJW ?DDCAR RFC R?PECR ACJJ @CA?SQC RFCW A?LLMR CLRCP RFC ACJJ ?LB ?AR BGPCARJW ML %/". #GLBGLE MD RFCQC FMPKMLCQ RM ? ACJJ QSPD?AC PCACNRMP
1068
#(!04%2 37 X 4(% %.$/#2).% 3934%-
PCQSJRQ GL ?ARGT?RGML MD ? QGEL?JGLE N?RFU?W; RFGQ RPGEECPQ GLRP?ACJJSJ?P ?ARGTGRW ?LB A?PPGCQ MSR RFC QNCAGDGA CDDCARQ ?QQMAG?RCB UGRF RFC FMPKMLC. *L RFGQ U?W, LMRFGLE N?QQCQ RFPMSEF RFC ACJJ KCK@P?LC; RFC FMPKMLC RF?R @GLBQ ?R RFC QSPD?AC PCK?GLQ ?R RFC QSPD?AC MD RFC ACJJ UFGJC RFC GLRP?ACJJSJ?P NPMBSAR PCK?GLQ GLQGBC RFC ACJJ. 5FC FMPKMLC RF?R GLGRG?RCQ RFC QGEL?JGLE N?RFU?W GQ A?JJCB ? first messenger, UFGAF ?ARGT?RCQ ? QCAMLB KCQQCLECP GL RFC AWRMNJ?QK, ?Q GJJSQRP?RCB GL Figure 37.6.
!@>LI< 37.6 4EB >JFKL >@FA-ABOFSBA ELOJLKBP BMFKBMEOFKB >KA KLOBMFKBMEOFKB ?FKA QL ?BQ>>AOBKBODF@ OB@BMQLOP LK QEB MI>PJ> JBJ?O>KB LC @BIIP. (LOJLKB ?FKAFKD QL OB@BMQLO >@QFS>QBP > '-MOLQBFK, TEF@E FK QROK >@QFS>QBP >ABKVIVI @V@I>PB, @LKSBOQFKD !40 QL @!-0. @!-0 FP > PB@LKA JBPPBKDBO QE>Q JBAF>QBP > @BII-PMB@FCF@ OBPMLKPB. !K BKWVJB @>IIBA MELPMELAFBPQBO>PB ?OB>HP ALTK @!-0, QBOJFK>QFKD QEB PFDK>I.
0LC TCPW GKNMPR?LR QCAMLB KCQQCLECP GQ AWAJGA ".1 (A".1). 8FCL ? FMPKMLC @GLBQ RM GRQ KCK@P?LC PCACNRMP, ? G-protein RF?R GQ ?QQMAG?RCB UGRF RFC PCACNRMP GQ ?ARGT?RCB; (-NPMRCGLQ ?PC NPMRCGLQ QCN?P?RC DPMK PCACNRMPQ RF?R ?PC DMSLB GL RFC ACJJ KCK@P?LC. 8FCL ? FMPKMLC GQ LMR @MSLB RM RFC PCACNRMP, RFC (-NPMRCGL GQ GL?ARGTC ?LB GQ @MSLB RM ES?LMQGLC BGNFMQNF?RC, MP (%1. 8FCL ? FMPKMLC @GLBQ RM RFC PCACNRMP, RFC (-NPMRCGL GQ ?ARGT?RCB @W @GLBGLE ES?LMQGLC RPGNFMQNF?RC, MP (51, GL NJ?AC MD (%1. "DRCP @GLBGLE, (51 GQ FWBPMJWQCB @W RFC (-NPMRCGL GLRM (%1 ?LB @CAMKCQ GL?ARGTC. 5FC ?ARGT?RCB (-NPMRCGL GL RSPL ?ARGT?RCQ ? KCK@P?LC-@MSLB CLXWKC A?JJCB adenylyl cyclase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phosphodiesterase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
#(!04%2 37 X 4(% %.$/#2).% 3934%-
1069
37.3 R -<>LC8K@FE F= BF;P +IF:<JJ<J

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL FMU FMPKMLCQ PCESJ?RC RFC CVAPCRMPW QWQRCK z %GQASQQ RFC PMJC MD FMPKMLCQ GL RFC PCNPMBSARGTC QWQRCK z %CQAPG@C FMU FMPKMLCQ PCESJ?RC KCR?@MJGQK z &VNJ?GL RFC PMJC MD FMPKMLCQ GL BGDDCPCLR BGQC?QCQ )MPKMLCQ F?TC ? UGBC P?LEC MD CDDCARQ ?LB KMBSJ?RC K?LW BGDDCPCLR @MBW NPMACQQCQ. 5FC ICW PCESJ?RMPW NPMACQQCQ RF?R UGJJ @C CV?KGLCB FCPC ?PC RFMQC ?DDCARGLE RFC CVAPCRMPW QWQRCK, RFC PCNPMBSARGTC QWQRCK, KCR?@MJGQK, @JMMB A?JAGSK AMLACLRP?RGMLQ, EPMURF, ?LB RFC QRPCQQ PCQNMLQC.
#FIDFE8C -<>LC8K@FE F= K?< O:I<KFIP .PJK<D

.?GLR?GLGLE ? NPMNCP U?RCP @?J?LAC GL RFC @MBW GQ GKNMPR?LR RM ?TMGB BCFWBP?RGML MP MTCP-FWBP?RGML (FWNML?RPCKG?). 5FC U?RCP AMLACLRP?RGML MD RFC @MBW GQ KMLGRMPCB @W osmoreceptors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antidiuretic hormone (ADH), UFGAF GQ RP?LQNMPRCB RM ?LB PCJC?QCB DPMK RFC NMQRCPGMP NGRSGR?PW EJ?LB. 5FC NPGLAGN?J ?ARGML MD "%) GQ RM PCESJ?RC RFC ?KMSLR MD U?RCP CVAPCRCB @W RFC IGBLCWQ. "Q "%) (UFGAF GQ ?JQM ILMUL ?Q T?QMNPCQQGL) A?SQCQ BGPCAR U?RCP PC?@QMPNRGML DPMK RFC IGBLCW RS@SJCQ, Q?JRQ ?LB U?QRCQ ?PC AMLACLRP?RCB GL UF?R UGJJ CTCLRS?JJW @C CVAPCRCB ?Q SPGLC. 5FC FWNMRF?J?KSQ AMLRPMJQ RFC KCAF?LGQKQ MD "%) QCAPCRGML, CGRFCP @W PCESJ?RGLE @JMMB TMJSKC MP RFC AMLACLRP?RGML MD U?RCP GL RFC @JMMB. %CFWBP?RGML MP NFWQGMJMEGA?J QRPCQQ A?L A?SQC ?L GLAPC?QC MD MQKMJ?PGRW ?@MTC 300 K0QK/-, UFGAF GL RSPL, P?GQCQ "%) QCAPCRGML ?LB U?RCP UGJJ @C PCR?GLCB, A?SQGLE ?L GLAPC?QC GL @JMMB NPCQQSPC. "%) RP?TCJQ GL RFC @JMMBQRPC?K RM RFC IGBLCWQ. 0LAC ?R RFC IGBLCWQ, "%) AF?LECQ RFC IGBLCWQ RM @CAMKC KMPC NCPKC?@JC RM U?RCP @W RCKNMP?PGJW GLQCPRGLE U?RCP AF?LLCJQ, ?OS?NMPGLQ, GLRM RFC IGBLCW RS@SJCQ. 8?RCP KMTCQ MSR MD RFC IGBLCW RS@SJCQ RFPMSEF RFC ?OS?NMPGLQ, PCBSAGLE SPGLC TMJSKC. 5FC U?RCP GQ PC?@QMP@CB GLRM RFC A?NGJJ?PGCQ JMUCPGLE @JMMB MQKMJ?PGRW @?AI RMU?PB LMPK?J. "Q @JMMB MQKMJ?PGRW BCAPC?QCQ, ? LCE?RGTC DCCB@?AI KCAF?LGQK PCBSACQ MQKMPCACNRMP ?ARGTGRW GL RFC FWNMRF?J?KSQ, ?LB "%) QCAPCRGML GQ PCBSACB. "%) PCJC?QC A?L @C PCBSACB @W ACPR?GL QS@QR?LACQ, GLAJSBGLE ?JAMFMJ, UFGAF A?L A?SQC GLAPC?QCB SPGLC NPMBSARGML ?LB BCFWBP?RGML. $FPMLGA SLBCPNPMBSARGML MD "%) MP ? KSR?RGML GL RFC "%) PCACNRMP PCQSJRQ GL diabetes insipidus. *D RFC NMQRCPGMP NGRSGR?PW BMCQ LMR PCJC?QC CLMSEF "%), U?RCP A?LLMR @C PCR?GLCB @W RFC IGBLCWQ ?LB GQ JMQR ?Q SPGLC. 5FGQ A?SQCQ GLAPC?QCB RFGPQR, @SR U?RCP R?ICL GL GQ JMQR ?E?GL ?LB KSQR @C AMLRGLS?JJW AMLQSKCB. *D RFC AMLBGRGML GQ LMR QCTCPC, BCFWBP?RGML K?W LMR MAASP, @SR QCTCPC A?QCQ A?L JC?B RM CJCARPMJWRC GK@?J?LACQ BSC RM BCFWBP?RGML. "LMRFCP FMPKMLC PCQNMLQG@JC DMP K?GLR?GLGLE CJCARPMJWRC AMLACLRP?RGMLQ GL CVRP?ACJJSJ?P DJSGBQ GQ aldosterone, ? QRCPMGB FMPKMLC RF?R GQ NPMBSACB @W RFC ?BPCL?J AMPRCV. *L AMLRP?QR RM "%), UFGAF NPMKMRCQ RFC PC?@QMPNRGML MD U?RCP RM K?GLR?GL NPMNCP U?RCP @?J?LAC, ?JBMQRCPMLC K?GLR?GLQ NPMNCP U?RCP @?J?LAC @W CLF?LAGLE /?+ PC?@QMPNRGML ?LB ,+ QCAPCRGML DPMK CVRP?ACJJSJ?P DJSGB MD RFC ACJJQ GL IGBLCW RS@SJCQ. #CA?SQC GR GQ NPMBSACB GL RFC AMPRCV MD RFC ?BPCL?J EJ?LB ?LB ?DDCARQ RFC AMLACLRP?RGMLQ MD KGLCP?JQ /?+ ?LB ,+, ?JBMQRCPMLC GQ PCDCPPCB RM ?Q ? mineralocorticoid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igure 37.7. 8FCL @JMMB NPCQQSPC BPMNQ, RFC PCLGL-?LEGMRCLQGL?JBMQRCPMLC QWQRCK (3""4) GQ ?ARGT?RCB. $CJJQ GL RFC HSVR?EJMKCPSJ?P ?NN?P?RSQ, UFGAF PCESJ?RCQ RFC DSLARGMLQ MD RFC LCNFPMLQ MD RFC IGBLCW, BCRCAR RFGQ ?LB PCJC?QC renin. 3CLGL, ?L CLXWKC, AGPASJ?RCQ GL RFC @JMMB ?LB PC?ARQ UGRF ? NJ?QK? NPMRCGL NPMBSACB @W RFC JGTCP A?JJCB ?LEGMRCLQGLMECL. 8FCL ?LEGMRCLQGLMECL GQ AJC?TCB @W PCLGL, GR NPMBSACQ ?LEGMRCLQGL *, UFGAF GQ RFCL AMLTCPRCB GLRM ?LEGMRCLQGL
1070
#(!04%2 37 X 4(% %.$/#2).% 3934%-
** GL RFC JSLEQ. "LEGMRCLQGL ** DSLARGMLQ ?Q ? FMPKMLC ?LB RFCL A?SQCQ RFC PCJC?QC MD RFC FMPKMLC ?JBMQRCPMLC @W RFC ?BPCL?J AMPRCV, PCQSJRGLE GL GLAPC?QCB /?+ PC?@QMPNRGML, U?RCP PCRCLRGML, ?LB ?L GLAPC?QC GL @JMMB NPCQQSPC. "LEGMRCLQGL ** GL ?BBGRGML RM @CGLE ? NMRCLR T?QMAMLQRPGARMP ?JQM A?SQCQ ?L GLAPC?QC GL "%) ?LB GLAPC?QCB RFGPQR, @MRF MD UFGAF FCJN RM P?GQC @JMMB NPCQQSPC.
!@>LI< 37.7 !$( >KA >IALPQBOLKB FK@OB>PB ?ILLA MOBPPROB >KA SLIRJB. !KDFLQBKPFK )) PQFJRI>QBP OBIB>PB LC QEBPB ELOJLKBP. !KDFLQBKPFK )), FK QROK, FP CLOJBA TEBK OBKFK @IB>SBP >KDFLQBKPFK. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V -FH>BI (aDDPQOhJ)
#FIDFE8C -<>LC8K@FE F= K?< -<GIF;L:K@M< .PJK<D

3CESJ?RGML MD RFC PCNPMBSARGTC QWQRCK GQ ? NPMACQQ RF?R PCOSGPCQ RFC ?ARGML MD FMPKMLCQ DPMK RFC NGRSGR?PW EJ?LB, RFC ?BPCL?J AMPRCV, ?LB RFC EML?BQ. %SPGLE NS@CPRW GL @MRF K?JCQ ?LB DCK?JCQ, RFC FWNMRF?J?KSQ NPMBSACQ EML?BMRPMNGL-PCJC?QGLE FMPKMLC ((L3)), UFGAF QRGKSJ?RCQ RFC NPMBSARGML ?LB PCJC?QC MD follicle-stimulating hormone (FSH) ?LB JSRCGLGXGLE FMPKMLC (-)) DPMK RFC ?LRCPGMP NGRSGR?PW EJ?LB. 5FCQC FMPKMLCQ PCESJ?RC RFC EML?BQ (RCQRCQ GL K?JCQ ?LB MT?PGCQ GL DCK?JCQ) ?LB RFCPCDMPC ?PC A?JJCB gonadotropins. *L @MRF K?JCQ ?LB DCK?JCQ, '4) QRGKSJ?RCQ E?KCRC NPMBSARGML ?LB -) QRGKSJ?RCQ NPMBSARGML MD FMPKMLCQ @W RFC EML?BQ. "L GLAPC?QC GL EML?B FMPKMLC JCTCJQ GLFG@GRQ (L3) NPMBSARGML RFPMSEF ? LCE?RGTC DCCB@?AI JMMN. +-:1'9/43 4, 9.+ M'1+ +574*:)9/;+ !>89+2 *L K?JCQ, '4) QRGKSJ?RCQ RFC K?RSP?RGML MD QNCPK ACJJQ. '4) NPMBSARGML GQ GLFG@GRCB @W RFC FMPKMLC GLFG@GL, UFGAF GQ PCJC?QCB @W RFC RCQRCQ. -) QRGKSJ?RCQ NPMBSARGML MD RFC QCV FMPKMLCQ ( androgens) @W RFC GLRCPQRGRG?J ACJJQ MD RFC RCQRCQ ?LB RFCPCDMPC GQ ?JQM A?JJCB GLRCPQRGRG?J ACJJ-QRGKSJ?RGLE FMPKMLC. 5FC KMQR UGBCJW ILMUL ?LBPMECL GL K?JCQ GQ RCQRMQRCPMLC. 5CQRMQRCPMLC NPMKMRCQ RFC NPMBSARGML MD QNCPK ?LB K?QASJGLC AF?P?ARCPGQRGAQ. 5FC ?BPCL?J AMPRCV ?JQM NPMBSACQ QK?JJ ?KMSLRQ MD RCQRMQRCPMLC NPCASPQMP, ?JRFMSEF RFC PMJC MD RFGQ ?BBGRGML?J FMPKMLC NPMBSARGML GQ LMR DSJJW SLBCPQRMMB.
#(!04%2 37 X 4(% %.$/#2).% 3934%-
1071
/?< D8E><IJ F= .PEK?<K@: #FIDFE<J
!@>LI< 37.8 0OLCBPPFLK>I ?>PB?>II MI>VBO *>PLK 'F>J?F MR?IF@>IIV >AJFQQBA QL, >KA >MLILDFWBA CLO, EFP RPB LC >K>?LIF@ PQBOLFAP PRMMIFBA ?V > QO>FKBO. (@OBAFQ: "OV@B %AT>OAP)
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igure 37.9. &QRP?BGMJ ?LB NPMECQRCPMLC ?PC QRCPMGB FMPKMLCQ RF?R NPCN?PC RFC @MBW DMP NPCEL?LAW. &QRP?BGMJ NPMBSACQ QCAMLB?PW QCV AF?P?ARCPGQRGAQ GL DCK?JCQ, UFGJC @MRF CQRP?BGMJ ?LB NPMECQRCPMLC PCESJ?RC RFC KCLQRPS?J AWAJC.
1072
#(!04%2 37 X 4(% %.$/#2).% 3934%-
!@>LI< 37.9 (LOJLK>I OBDRI>QFLK LC QEB CBJ>IB OBMOLAR@QFSB PVPQBJ FKSLISBP ELOJLKBP COLJ QEB EVMLQE>I>JRP, MFQRFQ>OV, >KA LS>OFBP.
*L ?BBGRGML RM NPMBSAGLE '4) ?LB -), RFC ?LRCPGMP NMPRGML MD RFC NGRSGR?PW EJ?LB ?JQM NPMBSACQ RFC FMPKMLC prolactin (PRL) GL DCK?JCQ. 1PMJ?ARGL QRGKSJ?RCQ RFC NPMBSARGML MD KGJI @W RFC K?KK?PW EJ?LBQ DMJJMUGLE AFGJB@GPRF. 1PMJ?ARGL JCTCJQ ?PC PCESJ?RCB @W RFC FWNMRF?J?KGA FMPKMLCQ prolactinreleasing hormone (PRH) ?LB prolactin-inhibiting hormone (PIH), UFGAF GQ LMU ILMUL RM @C BMN?KGLC. 13) QRGKSJ?RCQ RFC PCJC?QC MD NPMJ?ARGL ?LB 1*) GLFG@GRQ GR. 5FC NMQRCPGMP NGRSGR?PW PCJC?QCQ RFC FMPKMLC oxytocin, UFGAF QRGKSJ?RCQ SRCPGLC AMLRP?ARGMLQ BSPGLE AFGJB@GPRF. 5FC SRCPGLC QKMMRF KSQAJCQ ?PC LMR TCPW QCLQGRGTC RM MVWRMAGL SLRGJ J?RC GL NPCEL?LAW UFCL RFC LSK@CP MD MVWRMAGL PCACNRMPQ GL RFC SRCPSQ NC?IQ. 4RPCRAFGLE MD RGQQSCQ GL RFC SRCPSQ ?LB ACPTGV QRGKSJ?RCQ MVWRMAGL PCJC?QC BSPGLE AFGJB@GPRF. $MLRP?ARGMLQ GLAPC?QC GL GLRCLQGRW ?Q @JMMB JCTCJQ MD MVWRMAGL PGQC TG? ? NMQGRGTC DCCB@?AI KCAF?LGQK SLRGJ RFC @GPRF GQ AMKNJCRC. 0VWRMAGL ?JQM QRGKSJ?RCQ RFC AMLRP?ARGML MD KWMCNGRFCJG?J ACJJQ ?PMSLB RFC KGJI-NPMBSAGLE K?KK?PW EJ?LBQ. "Q RFCQC ACJJQ AMLRP?AR, KGJI GQ DMPACB DPMK RFC QCAPCRMPW ?JTCMJG GLRM KGJI BSARQ ?LB GQ CHCARCB DPMK RFC @PC?QRQ GL KGJI CHCARGML (tJCRBMULu) PCDJCV. 0VWRMAGL PCJC?QC GQ QRGKSJ?RCB @W RFC QSAIJGLE MD ?L GLD?LR, UFGAF RPGEECPQ RFC QWLRFCQGQ MD MVWRMAGL GL RFC FWNMRF?J?KSQ ?LB GRQ PCJC?QC GLRM AGPASJ?RGML ?R RFC NMQRCPGMP NGRSGR?PW.
#FIDFE8C -<>LC8K@FE F= (<K89FC@JD

#JMMB EJSAMQC JCTCJQ T?PW UGBCJW MTCP RFC AMSPQC MD ? B?W ?Q NCPGMBQ MD DMMB AMLQSKNRGML ?JRCPL?RC UGRF NCPGMBQ MD D?QRGLE. *LQSJGL ?LB EJSA?EML ?PC RFC RUM FMPKMLCQ NPGK?PGJW PCQNMLQG@JC DMP K?GLR?GLGLE FMKCMQR?QGQ MD @JMMB EJSAMQC JCTCJQ. "BBGRGML?J PCESJ?RGML GQ KCBG?RCB @W RFC RFWPMGB FMPKMLCQ.
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+-:1'9/43 4, B144* G1:)48+ L+;+18 (> I38:1/3 '3* G1:)'-43 $CJJQ MD RFC @MBW PCOSGPC LSRPGCLRQ GL MPBCP RM DSLARGML, ?LB RFCQC LSRPGCLRQ ?PC M@R?GLCB RFPMSEF DCCBGLE. *L MPBCP RM K?L?EC LSRPGCLR GLR?IC, QRMPGLE CVACQQ GLR?IC ?LB SRGJGXGLE PCQCPTCQ UFCL LCACQQ?PW, RFC @MBW SQCQ FMPKMLCQ RM KMBCP?RC CLCPEW QRMPCQ. Insulin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tJMU QSE?Pu CDDCAR, UFGAF GLFG@GRQ DSPRFCP GLQSJGL PCJC?QC DPMK @CR? ACJJQ RFPMSEF ? LCE?RGTC DCCB@?AI JMMN.
5FGQ animation (http://openstaxcollege.org/l/insulin) BCQAPG@C RFC PMJC MD GLQSJGL ?LB RFC N?LAPC?Q GL BG?@CRCQ. *KN?GPCB GLQSJGL DSLARGML A?L JC?B RM ? AMLBGRGML A?JJCB diabetes mellitus, RFC K?GL QWKNRMKQ MD UFGAF ?PC GJJSQRP?RCB GL Figure 37.10. 5FGQ A?L @C A?SQCB @W JMU JCTCJQ MD GLQSJGL NPMBSARGML @W RFC @CR? ACJJQ MD RFC N?LAPC?Q, MP @W PCBSACB QCLQGRGTGRW MD RGQQSC ACJJQ RM GLQSJGL. 5FGQ NPCTCLRQ EJSAMQC DPMK @CGLE ?@QMP@CB @W ACJJQ, A?SQGLE FGEF JCTCJQ MD @JMMB EJSAMQC, MP hyperglycemia (FGEF QSE?P). )GEF @JMMB EJSAMQC JCTCJQ K?IC GR BGDDGASJR DMP RFC IGBLCWQ RM PCAMTCP ?JJ RFC EJSAMQC DPMK L?QACLR SPGLC, PCQSJRGLE GL EJSAMQC @CGLE JMQR GL SPGLC. )GEF EJSAMQC JCTCJQ ?JQM PCQSJR GL JCQQ U?RCP @CGLE PC?@QMP@CB @W RFC IGBLCWQ, A?SQGLE FGEF ?KMSLRQ MD SPGLC RM @C NPMBSACB; RFGQ K?W PCQSJR GL BCFWBP?RGML. 0TCP RGKC, FGEF @JMMB EJSAMQC JCTCJQ A?L A?SQC LCPTC B?K?EC RM RFC CWCQ ?LB NCPGNFCP?J @MBW RGQQSCQ, ?Q UCJJ ?Q B?K?EC RM RFC IGBLCWQ ?LB A?PBGMT?QASJ?P QWQRCK. 0TCPQCAPCRGML MD GLQSJGL A?L A?SQC hypoglycemia, JMU @JMMB EJSAMQC JCTCJQ. 5FGQ A?SQCQ GLQSDDGAGCLR EJSAMQC ?T?GJ?@GJGRW RM ACJJQ, MDRCL JC?BGLE RM KSQAJC UC?ILCQQ, ?LB A?L QMKCRGKCQ A?SQC SLAMLQAGMSQLCQQ MP BC?RF GD JCDR SLRPC?RCB.
!@>LI< 37.10 4EB J>FK PVJMQLJP LC AF>?BQBP >OB PELTK. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V -FH>BI (aDDPQOhJ)
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#(!04%2 37 X 4(% %.$/#2).% 3934%-
8FCL @JMMB EJSAMQC JCTCJQ BCAJGLC @CJMU LMPK?J JCTCJQ, DMP CV?KNJC @CRUCCL KC?JQ MP UFCL EJSAMQC GQ SRGJGXCB P?NGBJW BSPGLE CVCPAGQC, RFC FMPKMLC glucagon GQ PCJC?QCB DPMK RFC ?JNF? ACJJQ MD RFC N?LAPC?Q. (JSA?EML P?GQCQ @JMMB EJSAMQC JCTCJQ, CJGAGRGLE UF?R GQ A?JJCB ? FWNCPEJWACKGA CDDCAR, @W QRGKSJ?RGLE RFC @PC?IBMUL MD EJWAMECL RM EJSAMQC GL QICJCR?J KSQAJC ACJJQ ?LB JGTCP ACJJQ GL ? NPMACQQ A?JJCB glycogenolysis. (JSAMQC A?L RFCL @C SRGJGXCB ?Q CLCPEW @W KSQAJC ACJJQ ?LB PCJC?QCB GLRM AGPASJ?RGML @W RFC JGTCP ACJJQ. (JSA?EML ?JQM QRGKSJ?RCQ ?@QMPNRGML MD ?KGLM ?AGBQ DPMK RFC @JMMB @W RFC JGTCP, UFGAF RFCL AMLTCPRQ RFCK RM EJSAMQC. 5FGQ NPMACQQ MD EJSAMQC QWLRFCQGQ GQ A?JJCB gluconeogenesis. (JSA?EML ?JQM QRGKSJ?RCQ ?BGNMQC ACJJQ RM PCJC?QC D?RRW ?AGBQ GLRM RFC @JMMB. 5FCQC ?ARGMLQ KCBG?RCB @W EJSA?EML PCQSJR GL ?L GLAPC?QC GL @JMMB EJSAMQC JCTCJQ RM LMPK?J FMKCMQR?RGA JCTCJQ. 3GQGLE @JMMB EJSAMQC JCTCJQ GLFG@GR DSPRFCP EJSA?EML PCJC?QC @W RFC N?LAPC?Q TG? ? LCE?RGTC DCCB@?AI KCAF?LGQK. *L RFGQ U?W, GLQSJGL ?LB EJSA?EML UMPI RMECRFCP RM K?GLR?GL FMKCMQR?RGA EJSAMQC JCTCJQ, ?Q QFMUL GL Figure 37.11.
!@>LI< 37.11 )KPRIFK >KA DIR@>DLK OBDRI>QB ?ILLA DIR@LPB IBSBIP.
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thyroxine, ?JQM ILMUL ?Q RCRP?GMBMRFWPMLGLC MP 54, ?LB triiodothyronine, ?JQM ILMUL ?Q 53. 5FCQC FMPKMLCQ ?DDCAR LC?PJW CTCPW ACJJ GL RFC @MBW CVACNR DMP RFC ?BSJR @P?GL, SRCPSQ, RCQRCQ, @JMMB ACJJQ, ?LB QNJCCL. 5FCW ?PC RP?LQNMPRCB ?APMQQ RFC NJ?QK? KCK@P?LC MD R?PECR ACJJQ ?LB @GLB RM PCACNRMPQ ML RFC KGRMAFMLBPG? PCQSJRGLE GL GLAPC?QCB "51 NPMBSARGML. *L RFC LSAJCSQ, 53 ?LB 54 ?ARGT?RC ECLCQ GLTMJTCB GL CLCPEW NPMBSARGML ?LB EJSAMQC MVGB?RGML. 5FGQ PCQSJRQ GL GLAPC?QCB P?RCQ MD KCR?@MJGQK ?LB @MBW FC?R NPMBSARGML, UFGAF GQ ILMUL ?Q RFC FMPKMLCsQ A?JMPGECLGA CDDCAR. 53 ?LB 54 PCJC?QC DPMK RFC RFWPMGB EJ?LB GQ QRGKSJ?RCB @W thyroid-stimulating hormone (TSH), UFGAF GQ NPMBSACB @W RFC ?LRCPGMP NGRSGR?PW. 54) @GLBGLE ?R RFC PCACNRMPQ MD RFC DMJJGAJC MD RFC RFWPMGB RPGEECPQ RFC NPMBSARGML MD 53 ?LB 54 DPMK ? EJWAMNPMRCGL A?JJCB thyroglobulin. 5FWPMEJM@SJGL GQ NPCQCLR GL RFC DMJJGAJCQ MD RFC RFWPMGB, ?LB GQ AMLTCPRCB GLRM RFWPMGB FMPKMLCQ UGRF RFC ?BBGRGML MD GMBGLC. *MBGLC GQ DMPKCB DPMK GMBGBC GMLQ RF?R ?PC ?ARGTCJW RP?LQNMPRCB GLRM RFC RFWPMGB DMJJGAJC DPMK RFC @JMMBQRPC?K.
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s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goiter, UFGAF GQ A?SQCB @W MTCPNPMBSARGML MD 54) UGRFMSR RFC DMPK?RGML MD RFWPMGB FMPKMLC. 5FWPMEJM@SJGL GQ AMLR?GLCB GL ? DJSGB A?JJCB AMJJMGB, ?LB 54) QRGKSJ?RGML PCQSJRQ GL FGEFCP JCTCJQ MD AMJJMGB ?AASKSJ?RGML GL RFC RFWPMGB. *L RFC ?@QCLAC MD GMBGLC, RFGQ GQ LMR AMLTCPRCB RM RFWPMGB FMPKMLC, ?LB AMJJMGB @CEGLQ RM ?AASKSJ?RC KMPC ?LB KMPC GL RFC RFWPMGB EJ?LB, JC?BGLE RM EMGRCP. %GQMPBCPQ A?L ?PGQC DPMK @MRF RFC SLBCPNPMBSARGML ?LB MTCPNPMBSARGML MD RFWPMGB FMPKMLCQ. Hypothyroidism, SLBCPNPMBSARGML MD RFC RFWPMGB FMPKMLCQ, A?L A?SQC ? JMU KCR?@MJGA P?RC JC?BGLE RM UCGEFR E?GL, QCLQGRGTGRW RM AMJB, ?LB PCBSACB KCLR?J ?ARGTGRW, ?KMLE MRFCP QWKNRMKQ. *L AFGJBPCL, FWNMRFWPMGBGQK A?L A?SQC APCRGLGQK, UFGAF A?L JC?B RM KCLR?J PCR?PB?RGML ?LB EPMURF BCDCARQ. Hyperthyroidism, RFC MTCPNPMBSARGML MD RFWPMGB FMPKMLCQ, A?L JC?B RM ?L GLAPC?QCB KCR?@MJGA P?RC ?LB GRQ CDDCARQ: UCGEFR JMQQ, CVACQQ FC?R NPMBSARGML, QUC?RGLE, ?LB ?L GLAPC?QCB FC?PR P?RC. (P?TCQs BGQC?QC GQ MLC CV?KNJC MD ? FWNCPRFWPMGB AMLBGRGML.
#FIDFE8C CFEKIFC F= BCFF; C8C:@LD '<M<CJ

3CESJ?RGML MD @JMMB A?JAGSK AMLACLRP?RGMLQ GQ GKNMPR?LR DMP ECLCP?RGML MD KSQAJC AMLRP?ARGMLQ ?LB LCPTC GKNSJQCQ, UFGAF ?PC CJCARPGA?JJW QRGKSJ?RCB. *D A?JAGSK JCTCJQ ECR RMM FGEF, KCK@P?LC NCPKC?@GJGRW RM QMBGSK BCAPC?QCQ ?LB KCK@P?LCQ @CAMKC JCQQ PCQNMLQGTC. *D A?JAGSK JCTCJQ ECR RMM JMU, KCK@P?LC NCPKC?@GJGRW RM QMBGSK GLAPC?QCQ ?LB AMLTSJQGMLQ MP KSQAJC QN?QKQ A?L PCQSJR. #JMMB A?JAGSK JCTCJQ ?PC PCESJ?RCB @W parathyroid hormone (PTH), UFGAF GQ NPMBSACB @W RFC N?P?RFWPMGB EJ?LBQ, ?Q GJJSQRP?RCB GL Figure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calcitonin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
#FIDFE8C -<>LC8K@FE F= "IFNK?

)MPKML?J PCESJ?RGML GQ PCOSGPCB DMP RFC EPMURF ?LB PCNJGA?RGML MD KMQR ACJJQ GL RFC @MBW. Growth hormone (GH), NPMBSACB @W RFC ?LRCPGMP NMPRGML MD RFC NGRSGR?PW EJ?LB, ?AACJCP?RCQ RFC P?RC MD NPMRCGL QWLRFCQGQ, N?PRGASJ?PJW GL QICJCR?J KSQAJC ?LB @MLCQ. (PMURF FMPKMLC F?Q BGPCAR ?LB GLBGPCAR KCAF?LGQKQ MD ?ARGML. 5FC DGPQR BGPCAR ?ARGML MD () GQ QRGKSJ?RGML MD RPGEJWACPGBC @PC?IBMUL (JGNMJWQGQ) ?LB PCJC?QC GLRM RFC @JMMB @W ?BGNMAWRCQ. 5FGQ PCQSJRQ GL ? QUGRAF @W KMQR RGQQSCQ DPMK SRGJGXGLE EJSAMQC ?Q ?L CLCPEW QMSPAC RM SRGJGXGLE D?RRW ?AGBQ. 5FGQ NPMACQQ GQ A?JJCB ? glucose-sparing effect. *L ?LMRFCP BGPCAR KCAF?LGQK, () QRGKSJ?RCQ EJWAMECL @PC?IBMUL GL RFC JGTCP; RFC EJWAMECL GQ RFCL PCJC?QCB GLRM RFC @JMMB ?Q EJSAMQC. #JMMB EJSAMQC JCTCJQ GLAPC?QC ?Q KMQR RGQQSCQ ?PC SRGJGXGLE D?RRW ?AGBQ GLQRC?B MD EJSAMQC DMP RFCGP CLCPEW LCCBQ. 5FC () KCBG?RCB GLAPC?QC GL @JMMB EJSAMQC JCTCJQ GQ A?JJCB ? diabetogenic effect @CA?SQC GR GQ QGKGJ?P RM RFC FGEF @JMMB EJSAMQC JCTCJQ QCCL GL BG?@CRCQ KCJJGRSQ. 5FC GLBGPCAR KCAF?LGQK MD () ?ARGML GQ KCBG?RCB @W insulin-like growth factors (IGFs) MP QMK?RMKCBGLQ, UFGAF ?PC ? D?KGJW MD EPMURF-NPMKMRGLE NPMRCGLQ NPMBSACB @W RFC JGTCP, UFGAF QRGKSJ?RCQ RGQQSC EPMURF. *('Q QRGKSJ?RC RFC SNR?IC MD ?KGLM ?AGBQ DPMK RFC @JMMB, ?JJMUGLE RFC DMPK?RGML MD LCU NPMRCGLQ, N?PRGASJ?PJW GL QICJCR?J KSQAJC ACJJQ, A?PRGJ?EC ACJJQ, ?LB MRFCP R?PECR ACJJQ, ?Q QFMUL GL Figure 37.13. 5FGQ GQ CQNCAG?JJW GKNMPR?LR ?DRCP ? KC?J, UFCL EJSAMQC ?LB ?KGLM ?AGB AMLACLRP?RGML JCTCJQ ?PC FGEF GL RFC @JMMB. () JCTCJQ ?PC PCESJ?RCB @W RUM FMPKMLCQ NPMBSACB @W RFC FWNMRF?J?KSQ. () PCJC?QC GQ QRGKSJ?RCB @W growth hormone-releasing hormone (GHRH) ?LB GQ GLFG@GRCB @W growth hormoneinhibiting hormone (GHIH), ?JQM A?JJCB QMK?RMQR?RGL.
#(!04%2 37 X 4(% %.$/#2).% 3934%-
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" @?J?LACB NPMBSARGML MD EPMURF FMPKMLC GQ APGRGA?J DMP NPMNCP BCTCJMNKCLR. 6LBCPNPMBSARGML MD () GL ?BSJRQ BMCQ LMR ?NNC?P RM A?SQC ?LW ?@LMPK?JGRGCQ, @SR GL AFGJBPCL GR A?L PCQSJR GL pituitary dwarfism, GL UFGAF EPMURF GQ PCBSACB. 1GRSGR?PW BU?PDGQK GQ AF?P?ARCPGXCB @W QWKKCRPGA @MBW DMPK?RGML. *L QMKC A?QCQ, GLBGTGBS?JQ ?PC SLBCP 30 GLAFCQ GL FCGEFR. 0TCPQCAPCRGML MD EPMURF FMPKMLC A?L JC?B RM gigantism GL AFGJBPCL, A?SQGLE CVACQQGTC EPMURF. *L QMKC BMASKCLRCB A?QCQ, GLBGTGBS?JQ A?L PC?AF FCGEFRQ MD MTCP CGEFR DCCR. *L ?BSJRQ, CVACQQGTC () A?L JC?B RM acromegaly, ? AMLBGRGML GL UFGAF RFCPC GQ CLJ?PECKCLR MD @MLCQ GL RFC D?AC, F?LBQ, ?LB DCCR RF?R ?PC QRGJJ A?N?@JC MD EPMURF.
#FIDFE8C -<>LC8K@FE F= .KI<JJ

8FCL ? RFPC?R MP B?LECP GQ NCPACGTCB, RFC @MBW PCQNMLBQ @W PCJC?QGLE FMPKMLCQ RF?R UGJJ PC?BW GR DMP RFC tDGEFR-MP-DJGEFRu PCQNMLQC. 5FC CDDCARQ MD RFGQ PCQNMLQC ?PC D?KGJG?P RM ?LWMLC UFM F?Q @CCL GL ? QRPCQQDSJ QGRS?RGML: GLAPC?QCB FC?PR P?RC, BPW KMSRF, ?LB F?GP QR?LBGLE SN.
!@>?K-FI-!C@>?K -<JGFEJ<
)KQBO>@QFLKP LC QEB BKAL@OFKB ELOJLKBP E>SB BSLISBA QL BKPROB QEB ?LAVw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w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
1078
#(!04%2 37 X 4(% %.$/#2).% 3934%-
PCQNMLQCQ) TG? LCPTC GKNSJQCQ, ?LB RFC ?BPCL?J AMPRCV, UFGAF KCBG?RCQ JMLE-RCPK QRPCQQ PCQNMLQCQ, TG? RFC FMPKMLC adrenocorticotropic hormone (ACTH), UFGAF GQ NPMBSACB @W RFC ?LRCPGMP NGRSGR?PW. !.479-9+72 !97+88 +85438+ 8FCL NPCQCLRCB UGRF ? QRPCQQDSJ QGRS?RGML, RFC @MBW PCQNMLBQ @W A?JJGLE DMP RFC PCJC?QC MD FMPKMLCQ RF?R NPMTGBC ? @SPQR MD CLCPEW. 5FC FMPKMLCQ epinephrine (?JQM ILMUL ?Q ?BPCL?JGLC) ?LB norepinephrine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
8?RAF RFGQ Discovery Channel animation (http://openstaxcollege.org/l/adrenaline) BCQAPG@GLE RFC DJGEFR-MP-DJGEFR PCQNMLQC. L43--9+72 !97+88 +85438+ -MLE-RCPK QRPCQQ PCQNMLQC BGDDCPQ DPMK QFMPR-RCPK QRPCQQ PCQNMLQC. 5FC @MBW A?LLMR QSQR?GL RFC @SPQRQ MD CLCPEW KCBG?RCB @W CNGLCNFPGLC ?LB LMPCNGLCNFPGLC DMP JMLE RGKCQ. *LQRC?B, MRFCP FMPKMLCQ AMKC GLRM NJ?W. *L ? JMLE-RCPK QRPCQQ PCQNMLQC, RFC FWNMRF?J?KSQ RPGEECPQ RFC PCJC?QC MD "$5) DPMK RFC ?LRCPGMP NGRSGR?PW EJ?LB. 5FC ?BPCL?J AMPRCV GQ QRGKSJ?RCB @W "$5) RM PCJC?QC QRCPMGB FMPKMLCQ A?JJCB corticosteroids. $MPRGAMQRCPMGBQ RSPL ML RP?LQAPGNRGML MD ACPR?GL ECLCQ GL RFC LSAJCG MD R?PECR ACJJQ. 5FCW AF?LEC CLXWKC AMLACLRP?RGMLQ GL RFC AWRMNJ?QK ?LB ?DDCAR ACJJSJ?P KCR?@MJGQK. 5FCPC ?PC RUM K?GL AMPRGAMQRCPMGBQ: EJSAMAMPRGAMGBQ QSAF ?Q cortisol, ?LB KGLCP?JMAMPRGAMGBQ QSAF ?Q ?JBMQRCPMLC. 5FCQC FMPKMLCQ R?PECR RFC @PC?IBMUL MD D?R GLRM D?RRW ?AGBQ GL RFC ?BGNMQC RGQQSC. 5FC D?RRW ?AGBQ ?PC PCJC?QCB GLRM RFC @JMMBQRPC?K DMP MRFCP RGQQSCQ RM SQC DMP "51 NPMBSARGML. 5FC glucocorticoids NPGK?PGJW ?DDCAR EJSAMQC KCR?@MJGQK @W QRGKSJ?RGLE EJSAMQC QWLRFCQGQ. (JSAMAMPRGAMGBQ ?JQM F?TC ?LRGGLDJ?KK?RMPW NPMNCPRGCQ RFPMSEF GLFG@GRGML MD RFC GKKSLC QWQRCK. 'MP CV?KNJC, AMPRGQMLC GQ SQCB ?Q ?L ?LRG-GLDJ?KK?RMPW KCBGA?RGML; FMUCTCP, GR A?LLMR @C SQCB JMLE RCPK ?Q GR GLAPC?QCQ QSQACNRG@GJGRW RM BGQC?QC BSC RM GRQ GKKSLC-QSNNPCQQGLE CDDCARQ. .GLCP?JMAMPRGAMGBQ DSLARGML RM PCESJ?RC GML ?LB U?RCP @?J?LAC MD RFC @MBW. 5FC FMPKMLC ?JBMQRCPMLC QRGKSJ?RCQ RFC PC?@QMPNRGML MD U?RCP ?LB QMBGSK GMLQ GL RFC IGBLCW, UFGAF PCQSJRQ GL GLAPC?QCB @JMMB NPCQQSPC ?LB TMJSKC. )WNCPQCAPCRGML MD EJSAMAMPRGAMGBQ A?L A?SQC ? AMLBGRGML ILMUL ?Q Cushings disease, AF?P?ARCPGXCB @W ? QFGDRGLE MD D?R QRMP?EC ?PC?Q MD RFC @MBW. 5FGQ A?L A?SQC RFC ?AASKSJ?RGML MD ?BGNMQC RGQQSC GL RFC D?AC ?LB LCAI, ?LB CVACQQGTC EJSAMQC GL RFC @JMMB. )WNMQCAPCRGML MD RFC AMPRGAMQRCPMGBQ A?L A?SQC Addisons disease, UFGAF K?W PCQSJR GL @PMLXGLE MD RFC QIGL, FWNMEJWACKG?, ?LB JMU CJCARPMJWRC JCTCJQ GL RFC @JMMB.
37.4 R -<>LC8K@FE F= #FIDFE< +IF;L:K@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL FMU FMPKMLC NPMBSARGML GQ PCESJ?RCB z %GQASQQ RFC BGDDCPCLR QRGKSJG RF?R AMLRPMJ FMPKMLC JCTCJQ GL RFC @MBW )MPKMLC NPMBSARGML ?LB PCJC?QC ?PC NPGK?PGJW AMLRPMJJCB @W LCE?RGTC DCCB@?AI. *L LCE?RGTC DCCB@?AI QWQRCKQ, ? QRGKSJSQ CJGAGRQ RFC PCJC?QC MD ? QS@QR?LAC; MLAC RFC QS@QR?LAC PC?AFCQ ? ACPR?GL JCTCJ, GR QCLBQ ? QGEL?J RF?R QRMNQ DSPRFCP PCJC?QC MD RFC QS@QR?LAC. *L RFGQ U?W, RFC AMLACLRP?RGML MD FMPKMLCQ GL @JMMB GQ K?GLR?GLCB UGRFGL ? L?PPMU P?LEC. 'MP CV?KNJC, RFC ?LRCPGMP NGRSGR?PW QGEL?JQ RFC RFWPMGB RM
#(!04%2 37 X 4(% %.$/#2).% 3934%-
1079
PCJC?QC RFWPMGB FMPKMLCQ. *LAPC?QGLE JCTCJQ MD RFCQC FMPKMLCQ GL RFC @JMMB RFCL EGTC DCCB@?AI RM RFC FWNMRF?J?KSQ ?LB ?LRCPGMP NGRSGR?PW RM GLFG@GR DSPRFCP QGEL?JGLE RM RFC RFWPMGB EJ?LB, ?Q GJJSQRP?RCB GL Figure 37.14. 5FCPC ?PC RFPCC KCAF?LGQKQ @W UFGAF CLBMAPGLC EJ?LBQ ?PC QRGKSJ?RCB RM QWLRFCQGXC ?LB PCJC?QC FMPKMLCQ: FSKMP?J QRGKSJG, FMPKML?J QRGKSJG, ?LB LCSP?J QRGKSJG.
!@>LI< 37.14 4EB >KQBOFLO MFQRFQ>OV PQFJRI>QBP QEB QEVOLFA DI>KA QL OBIB>PB QEVOLFA ELOJLKBP 43 >KA 44. )K@OB>PFKD IBSBIP LC QEBPB ELOJLKBP FK QEB ?ILLA OBPRIQP FK CBBA?>@H QL QEB EVMLQE>I>JRP >KA >KQBOFLO MFQRFQ>OV QL FKEF?FQ CROQEBO PFDK>IFKD QL QEB QEVOLFA DI>KA. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V -FH>BI (aDDPQOhJ)
(VMBOQEVOLFAFPJ FP > @LKAFQFLK FK TEF@E QEB QEVOLFA DI>KA FP LSBO>@QFSB. (VMLQEVOLFAFPJ FP > @LKAFQFLK FK TEF@E QEB QEVOLFA DI>KA FP RKABO>@QFSB. 7EF@E LC QEB @LKAFQFLKP >OB QEB CLIILTFKD QTL M>QFBKQP JLPQ IFHBIV QL E>SB 0>QFBKQ ! E>P PVJMQLJP FK@IRAFKD TBFDEQ D>FK, @LIA PBKPFQFSFQV, ILT EB>OQ O>QB >KA C>QFDRB. 0>QFBKQ " E>P PVJMQLJP FK@IRAFKD TBFDEQ ILPP, MOLCRPB PTB>QFKD, FK@OB>PBA EB>OQ O>QB >KA AFCCF@RIQV PIBBMFKD.
#LDFI8C .K@DLC@
5FC RCPK tFSKMP?Ju GQ BCPGTCB DPMK RFC RCPK tFSKMP,u UFGAF PCDCPQ RM @MBGJW DJSGBQ QSAF ?Q @JMMB. " humoral stimulus PCDCPQ RM RFC AMLRPMJ MD FMPKMLC PCJC?QC GL PCQNMLQC RM AF?LECQ GL CVRP?ACJJSJ?P DJSGBQ QSAF ?Q @JMMB MP RFC GML AMLACLRP?RGML GL RFC @JMMB. 'MP CV?KNJC, ? PGQC GL @JMMB EJSAMQC JCTCJQ RPGEECPQ RFC N?LAPC?RGA PCJC?QC MD GLQSJGL. *LQSJGL A?SQCQ @JMMB EJSAMQC JCTCJQ RM BPMN, UFGAF QGEL?JQ RFC N?LAPC?Q RM QRMN NPMBSAGLE GLQSJGL GL ? LCE?RGTC DCCB@?AI JMMN.
#FIDFE8C .K@DLC@
Hormonal stimuli PCDCPQ RM RFC PCJC?QC MD ? FMPKMLC GL PCQNMLQC RM ?LMRFCP FMPKMLC. " LSK@CP MD CLBMAPGLC EJ?LBQ PCJC?QC FMPKMLCQ UFCL QRGKSJ?RCB @W FMPKMLCQ PCJC?QCB @W MRFCP CLBMAPGLC EJ?LBQ. 'MP CV?KNJC, RFC FWNMRF?J?KSQ NPMBSACQ FMPKMLCQ RF?R QRGKSJ?RC RFC ?LRCPGMP NMPRGML MD RFC NGRSGR?PW EJ?LB. 5FC ?LRCPGMP NGRSGR?PW GL RSPL PCJC?QCQ FMPKMLCQ RF?R PCESJ?RC FMPKMLC NPMBSARGML @W MRFCP CLBMAPGLC EJ?LBQ. 5FC ?LRCPGMP NGRSGR?PW PCJC?QCQ RFC RFWPMGB-QRGKSJ?RGLE FMPKMLC, UFGAF RFCL QRGKSJ?RCQ
1080
#(!04%2 37 X 4(% %.$/#2).% 3934%-
RFC RFWPMGB EJ?LB RM NPMBSAC RFC FMPKMLCQ 53 ?LB 54. "Q @JMMB AMLACLRP?RGMLQ MD 53 ?LB 54 PGQC, RFCW GLFG@GR @MRF RFC NGRSGR?PW ?LB RFC FWNMRF?J?KSQ GL ? LCE?RGTC DCCB@?AI JMMN.
)<LI8C .K@DLC@
*L QMKC A?QCQ, RFC LCPTMSQ QWQRCK BGPCARJW QRGKSJ?RCQ CLBMAPGLC EJ?LBQ RM PCJC?QC FMPKMLCQ, UFGAF GQ PCDCPPCB RM ?Q neural stimuli. 3CA?JJ RF?R GL ? QFMPR-RCPK QRPCQQ PCQNMLQC, RFC FMPKMLCQ CNGLCNFPGLC ?LB LMPCNGLCNFPGLC ?PC GKNMPR?LR DMP NPMTGBGLE RFC @SPQRQ MD CLCPEW PCOSGPCB DMP RFC @MBW RM PCQNMLB. )CPC, LCSPML?J QGEL?JGLE DPMK RFC QWKN?RFCRGA LCPTMSQ QWQRCK BGPCARJW QRGKSJ?RCQ RFC ?BPCL?J KCBSJJ? RM PCJC?QC RFC FMPKMLCQ CNGLCNFPGLC ?LB LMPCNGLCNFPGLC GL PCQNMLQC RM QRPCQQ.
37.5 R E;F:I@E< "C8E;J

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC PMJC MD BGDDCPCLR EJ?LBQ GL RFC CLBMAPGLC QWQRCK z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
#PGFK?8C8D@:-+@KL@K8IP AO@J
5FC hypothalamus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pituitary gland, QMKCRGKCQ A?JJCB RFC FWNMNFWQGQ MP tK?QRCP EJ?LBu GQ JMA?RCB ?R RFC @?QC MD RFC @P?GL GL RFC QCJJ? RSPAGA?, ? EPMMTC MD RFC QNFCLMGB @MLC MD RFC QISJJ, GJJSQRP?RCB GL Figure 37.15. *R GQ ?RR?AFCB RM RFC FWNMRF?J?KSQ TG? ? QR?JI A?JJCB RFC pituitary stalk (MP GLDSLBG@SJSK). 5FC ?LRCPGMP NMPRGML MD RFC NGRSGR?PW EJ?LB GQ PCESJ?RCB @W PCJC?QGLE MP PCJC?QC-GLFG@GRGLE FMPKMLCQ NPMBSACB @W RFC FWNMRF?J?KSQ, ?LB RFC NMQRCPGMP NGRSGR?PW PCACGTCQ QGEL?JQ TG? LCSPMQCAPCRMPW ACJJQ RM PCJC?QC FMPKMLCQ NPMBSACB @W RFC FWNMRF?J?KSQ. 5FC NGRSGR?PW F?Q RUM BGQRGLAR PCEGMLQqRFC ?LRCPGMP NGRSGR?PW ?LB RFC NMQRCPGMP NGRSGR?PWqUFGAF @CRUCCL RFCK QCAPCRC LGLC BGDDCPCLR NCNRGBC MP NPMRCGL FMPKMLCQ. 5FC NMQRCPGMP JM@C MD RFC NGRSGR?PW EJ?LB AMLR?GLQ ?VMLQ MD RFC FWNMRF?J?KGA LCSPMLQ.
!@>LI< 37.15 4EB MFQRFQ>OV DI>KA FP IL@>QBA >Q (>) QEB ?>PB LC QEB ?O>FK >KA (?) @LKKB@QBA QL QEB EVMLQE>I>JRP ?V QEB MFQRFQ>OV PQ>IH. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V .#); @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V 'O>VwP !K>QLJV)
A39+7/47 P/9:/9'7> 5FC anterior pituitary EJ?LB, MP ?BCLMFWNMNFWQGQ, GQ QSPPMSLBCB @W ? A?NGJJ?PW LCRUMPI RF?R CVRCLBQ DPMK RFC FWNMRF?J?KSQ, BMUL ?JMLE RFC GLDSLBG@SJSK, ?LB RM RFC ?LRCPGMP NGRSGR?PW. 5FGQ A?NGJJ?PW
#(!04%2 37 X 4(% %.$/#2).% 3934%-
1081
LCRUMPI GQ ? N?PR MD RFC hypophyseal portal system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posterior pituitary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
/?PIF@; "C8E;
5FC thyroid gland GQ JMA?RCB GL RFC LCAI, HSQR @CJMU RFC J?PWLV ?LB GL DPMLR MD RFC RP?AFC?, ?Q QFMUL GL Figure 37.16. *R GQ ? @SRRCPDJW-QF?NCB EJ?LB UGRF RUM JM@CQ RF?R ?PC AMLLCARCB @W RFC isthmus. *R F?Q ? B?PI PCB AMJMP BSC RM GRQ CVRCLQGTC T?QASJ?P QWQRCK. 8FCL RFC RFWPMGB QUCJJQ BSC RM BWQDSLARGML, GR A?L @C DCJR SLBCP RFC QIGL MD RFC LCAI.
!@>LI< 37.16 4EFP FIIRPQO>QFLK PELTP QEB IL@>QFLK LC QEB QEVOLFA DI>KA.
5FC RFWPMGB EJ?LB GQ K?BC SN MD K?LW QNFCPGA?J RFWPMGB DMJJGAJCQ, UFGAF ?PC JGLCB UGRF ? QGKNJC AS@MGB?J CNGRFCJGSK. 5FCQC DMJJGAJCQ AMLR?GL ? TGQAMSQ DJSGB, A?JJCB colloid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
1082
#(!04%2 37 X 4(% %.$/#2).% 3934%-
UFGAF GQ NPMBSACB @W RFC ?LRCPGMP NGRSGR?PW. 5FCQC RFWPMGB FMPKMLCQ GLAPC?QC RFC P?RCQ MD KGRMAFMLBPG?J "51 NPMBSARGML. " RFGPB FMPKMLC, A?JAGRMLGL, GQ NPMBSACB @W parafollicular cells MD RFC RFWPMGB CGRFCP PCJC?QGLE FMPKMLCQ MP GLFG@GRGLE FMPKMLCQ. $?JAGRMLGL PCJC?QC GQ LMR AMLRPMJJCB @W 54), @SR GLQRC?B GQ PCJC?QCB UFCL A?JAGSK GML AMLACLRP?RGMLQ GL RFC @JMMB PGQC. $?JAGRMLGL DSLARGMLQ RM FCJN PCESJ?RC A?JAGSK AMLACLRP?RGMLQ GL @MBW DJSGBQ. *R ?ARQ GL RFC @MLCQ RM GLFG@GR MQRCMAJ?QR ?ARGTGRW ?LB GL RFC IGBLCWQ RM QRGKSJ?RC CVAPCRGML MD A?JAGSK. 5FC AMK@GL?RGML MD RFCQC RUM CTCLRQ JMUCPQ @MBW DJSGB JCTCJQ MD A?JAGSK.
+8I8K?PIF@; "C8E;J
.MQR NCMNJC F?TC DMSP parathyroid glands; FMUCTCP, RFC LSK@CP A?L T?PW DPMK RUM RM QGV. 5FCQC EJ?LBQ ?PC JMA?RCB ML RFC NMQRCPGMP QSPD?AC MD RFC RFWPMGB EJ?LB, ?Q QFMUL GL Figure 37.17. /MPK?JJW, RFCPC GQ ? QSNCPGMP EJ?LB ?LB ?L GLDCPGMP EJ?LB ?QQMAG?RCB UGRF C?AF MD RFC RFWPMGBsQ RUM JM@CQ. &?AF N?P?RFWPMGB EJ?LB GQ AMTCPCB @W AMLLCARGTC RGQQSC ?LB AMLR?GLQ K?LW QCAPCRMPW ACJJQ RF?R ?PC ?QQMAG?RCB UGRF ? A?NGJJ?PW LCRUMPI.
!@>LI< 37.17 4EB M>O>QEVOLFA DI>KAP >OB IL@>QBA LK QEB MLPQBOFLO LC QEB QEVOLFA DI>KA. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .#))
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5FC adrenal glands ?PC ?QQMAG?RCB UGRF RFC IGBLCWQ; MLC EJ?LB GQ JMA?RCB ML RMN MD C?AF IGBLCW ?Q GJJSQRP?RCB GL Figure 37.18. 5FC ?BPCL?J EJ?LBQ AMLQGQR MD ?L MSRCP ?BPCL?J AMPRCV ?LB ?L GLLCP ?BPCL?J KCBSJJ?. 5FCQC PCEGMLQ QCAPCRC BGDDCPCLR FMPKMLCQ.
#(!04%2 37 X 4(% %.$/#2).% 3934%-
1083
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A*7+3'1 C479+= 5FC adrenal cortex GQ K?BC SN MD J?WCPQ MD CNGRFCJG?J ACJJQ ?LB ?QQMAG?RCB A?NGJJ?PW LCRUMPIQ. 5FCQC J?WCPQ DMPK RFPCC BGQRGLAR PCEGMLQ: ?L MSRCP XML? EJMKCPSJMQ? RF?R NPMBSACQ KGLCP?JMAMPRGAMGBQ, ? KGBBJC XML? D?QAGASJ?R? RF?R NPMBSACQ EJSAMAMPRGAMGBQ, ?LB ?L GLLCP XML? PCRGASJ?PGQ RF?R NPMBSACQ ?LBPMECLQ. 5FC K?GL KGLCP?JMAMPRGAMGB GQ ?JBMQRCPMLC, UFGAF PCESJ?RCQ RFC AMLACLRP?RGML MD /?+ GMLQ GL SPGLC, QUC?R, N?LAPC?Q, ?LB Q?JGT?. "JBMQRCPMLC PCJC?QC DPMK RFC ?BPCL?J AMPRCV GQ QRGKSJ?RCB @W ? BCAPC?QC GL @JMMB AMLACLRP?RGMLQ MD QMBGSK GMLQ, @JMMB TMJSKC, MP @JMMB NPCQQSPC, MP @W ?L GLAPC?QC GL @JMMB NMR?QQGSK JCTCJQ. 5FC RFPCC K?GL EJSAMAMPRGAMGBQ ?PC AMPRGQMJ, AMPRGAMQRCPMLC, ?LB AMPRGQMLC. 5FC EJSAMAMPRGAMGBQ QRGKSJ?RC RFC QWLRFCQGQ MD EJSAMQC ?LB EJSAMLCMECLCQGQ (AMLTCPRGLE ? LML-A?P@MFWBP?RC RM EJSAMQC) @W JGTCP ACJJQ ?LB RFCW NPMKMRC RFC PCJC?QC MD D?RRW ?AGBQ DPMK ?BGNMQC RGQQSC. 5FCQC FMPKMLCQ GLAPC?QC @JMMB EJSAMQC JCTCJQ RM K?GLR?GL JCTCJQ UGRFGL ? LMPK?J P?LEC @CRUCCL KC?JQ. 5FCQC FMPKMLCQ ?PC QCAPCRCB GL PCQNMLQC RM "$5) ?LB JCTCJQ ?PC PCESJ?RCB @W LCE?RGTC DCCB@?AI. "LBPMECLQ ?PC QCV FMPKMLCQ RF?R NPMKMRC K?QASJGLGRW. 5FCW ?PC NPMBSACB GL QK?JJ ?KMSLRQ @W RFC ?BPCL?J AMPRCV GL @MRF K?JCQ ?LB DCK?JCQ. 5FCW BM LMR ?DDCAR QCVS?J AF?P?ARCPGQRGAQ ?LB K?W QSNNJCKCLR QCV FMPKMLCQ PCJC?QCB DPMK RFC EML?BQ. A*7+3'1 M+*:11' 5FC adrenal medulla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5FC pancreas, GJJSQRP?RCB GL Figure 37.19, GQ ?L CJMLE?RCB MPE?L RF?R GQ JMA?RCB @CRUCCL RFC QRMK?AF ?LB RFC NPMVGK?J NMPRGML MD RFC QK?JJ GLRCQRGLC. *R AMLR?GLQ @MRF CVMAPGLC ACJJQ RF?R CVAPCRC BGECQRGTC CLXWKCQ ?LB CLBMAPGLC ACJJQ RF?R PCJC?QC FMPKMLCQ. *R GQ QMKCRGKCQ PCDCPPCB RM ?Q ? FCRCPMAPGLC EJ?LB @CA?SQC GR F?Q @MRF CLBMAPGLC ?LB CVMAPGLC DSLARGMLQ.
1084
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5FC CLBMAPGLC ACJJQ MD RFC N?LAPC?Q DMPK AJSQRCPQ A?JJCB N?LAPC?RGA GQJCRQ MP RFC islets of Langerhans, ?Q TGQG@JC GL RFC KGAPMEP?NF QFMUL GL Figure 37.20. 5FC N?LAPC?RGA GQJCRQ AMLR?GL RUM NPGK?PW ACJJ RWNCQ: alpha cells, UFGAF NPMBSAC RFC FMPKMLC EJSA?EML, ?LB beta cells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#(!04%2 37 X 4(% %.$/#2).% 3934%-
1085
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1088
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1090
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37.2 #FN #FIDFE<J 2FIB

)MPKMLCQ A?SQC ACJJSJ?P AF?LECQ @W @GLBGLE RM PCACNRMPQ ML R?PECR ACJJQ. 5FC LSK@CP MD PCACNRMPQ ML ? R?PECR ACJJ A?L GLAPC?QC MP BCAPC?QC GL PCQNMLQC RM FMPKMLC ?ARGTGRW. )MPKMLCQ A?L ?DDCAR ACJJQ BGPCARJW RFPMSEF GLRP?ACJJSJ?P FMPKMLC PCACNRMPQ MP GLBGPCARJW RFPMSEF NJ?QK? KCK@P?LC FMPKMLC PCACNRMPQ. -GNGB-BCPGTCB (QMJS@JC) FMPKMLCQ A?L CLRCP RFC ACJJ @W BGDDSQGLE ?APMQQ RFC NJ?QK? KCK@P?LC ?LB @GLBGLE RM %/" RM PCESJ?RC ECLC RP?LQAPGNRGML ?LB RM AF?LEC RFC ACJJsQ ?ARGTGRGCQ @W GLBSAGLE NPMBSARGML MD NPMRCGLQ RF?R ?DDCAR, GL ECLCP?J, RFC JMLE-RCPK QRPSARSPC ?LB DSLARGML MD RFC ACJJ. -GNGB GLQMJS@JC FMPKMLCQ @GLB RM PCACNRMPQ ML RFC NJ?QK? KCK@P?LC QSPD?AC ?LB RPGEECP ? QGEL?JGLE N?RFU?W RM AF?LEC RFC ACJJsQ ?ARGTGRGCQ @W GLBSAGLE NPMBSARGML MD T?PGMSQ ACJJ NPMBSARQ RF?R ?DDCAR RFC ACJJ GL RFC QFMPR-RCPK. 5FC FMPKMLC GQ A?JJCB ? DGPQR KCQQCLECP ?LB RFC ACJJSJ?P AMKNMLCLR GQ A?JJCB ? QCAMLB KCQQCLECP. (NPMRCGLQ ?ARGT?RC RFC QCAMLB KCQQCLECP (AWAJGA ".1), RPGEECPGLE RFC ACJJSJ?P PCQNMLQC. 3CQNMLQC RM FMPKMLC @GLBGLE GQ ?KNJGDGCB ?Q RFC QGEL?JGLE N?RFU?W NPMEPCQQCQ. $CJJSJ?P PCQNMLQCQ RM FMPKMLCQ GLAJSBC RFC NPMBSARGML MD NPMRCGLQ ?LB CLXWKCQ ?LB ?JRCPCB KCK@P?LC NCPKC?@GJGRW.
37.3 -<>LC8K@FE F= BF;P +IF:<JJ<J

8?RCP JCTCJQ GL RFC @MBW ?PC AMLRPMJJCB @W ?LRGBGSPCRGA FMPKMLC ("%)), UFGAF GQ NPMBSACB GL RFC FWNMRF?J?KSQ ?LB RPGEECPQ RFC PC?@QMPNRGML MD U?RCP @W RFC IGBLCWQ. 6LBCPNPMBSARGML MD "%) A?L A?SQC BG?@CRCQ GLQGNGBSQ. "JBMQRCPMLC, ? FMPKMLC NPMBSACB @W RFC ?BPCL?J AMPRCV MD RFC IGBLCWQ, CLF?LACQ /?+ PC?@QMPNRGML DPMK RFC CVRP?ACJJSJ?P DJSGBQ ?LB QS@QCOSCLR U?RCP PC?@QMPNRGML @W BGDDSQGML. 5FC PCLGL-?LEGMRCLQGL-?JBMQRCPMLC QWQRCK GQ MLC U?W RF?R ?JBMQRCPMLC PCJC?QC GQ AMLRPMJJCB. 5FC PCNPMBSARGTC QWQRCK GQ AMLRPMJJCB @W RFC EML?BMRPMNGLQ DMJJGAJC-QRGKSJ?RGLE FMPKMLC ('4)) ?LB JSRCGLGXGLE FMPKMLC (-)), UFGAF ?PC NPMBSACB @W RFC NGRSGR?PW EJ?LB. (ML?BMRPMNGL PCJC?QC GQ AMLRPMJJCB @W RFC FWNMRF?J?KGA FMPKMLC EML?BMRPMNGL-PCJC?QGLE FMPKMLC ((L3)). '4) QRGKSJ?RCQ RFC K?RSP?RGML MD QNCPK ACJJQ GL K?JCQ ?LB GQ GLFG@GRCB @W RFC FMPKMLC GLFG@GL, UFGJC -) QRGKSJ?RCQ RFC NPMBSARGML MD RFC ?LBPMECL RCQRMQRCPMLC. '4) QRGKSJ?RCQ CEE K?RSP?RGML GL DCK?JCQ, UFGJC -) QRGKSJ?RCQ RFC NPMBSARGML MD CQRPMECLQ ?LB NPMECQRCPMLC. Estrogens ?PC ? EPMSN MD QRCPMGB FMPKMLCQ NPMBSACB @W RFC MT?PGCQ RF?R RPGEECP RFC BCTCJMNKCLR MD QCAMLB?PW QCV AF?P?ARCPGQRGAQ GL DCK?JCQ ?Q UCJJ ?Q AMLRPMJ RFC K?RSP?RGML MD RFC MT?. *L DCK?JCQ, RFC NGRSGR?PW ?JQM NPMBSACQ NPMJ?ARGL, UFGAF QRGKSJ?RCQ KGJI NPMBSARGML ?DRCP AFGJB@GPRF, ?LB MVWRMAGL, UFGAF QRGKSJ?RCQ SRCPGLC AMLRP?ARGML BSPGLE AFGJB@GPRF ?LB KGJI JCR-BMUL BSPGLE QSAIJGLE. *LQSJGL GQ NPMBSACB @W RFC N?LAPC?Q GL PCQNMLQC RM PGQGLE @JMMB EJSAMQC JCTCJQ ?LB ?JJMUQ ACJJQ RM SRGJGXC @JMMB EJSAMQC ?LB QRMPC CVACQQ EJSAMQC DMP J?RCP SQC. %G?@CRCQ KCJJGRSQ GQ A?SQCB @W PCBSACB GLQSJGL ?ARGTGRW ?LB A?SQCQ FGEF @JMMB EJSAMQC JCTCJQ, MP FWNCPEJWACKG?. (JSA?EML GQ PCJC?QCB @W RFC N?LAPC?Q GL PCQNMLQC RM JMU @JMMB EJSAMQC JCTCJQ ?LB QRGKSJ?RCQ RFC @PC?IBMUL MD EJWAMECL GLRM EJSAMQC, UFGAF A?L @C SQCB @W RFC @MBW. 5FC @MBWs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
#(!04%2 37 X 4(% %.$/#2).% 3934%-
1091
RPGEECP RFC DGEFR MP DJGEFR PCQNMLQC. -MLE-RCPK QRPCQQ A?SQCQ RFC FWNMRF?J?KSQ RM RPGEECP RFC ?LRCPGMP NGRSGR?PW RM PCJC?QC ?BPCLMAMPRGAMRPMNGA FMPKMLC ("$5)), UFGAF A?SQCQ RFC PCJC?QC MD AMPRGAMQRCPMGBQ, EJSAMAMPRGAMGBQ, ?LB KGLCP?JMAMPRGAMGBQ, DPMK RFC ?BPCL?J AMPRCV.
37.4 -<>LC8K@FE F= #FIDFE< +IF;L:K@FE

)MPKMLC JCTCJQ ?PC NPGK?PGJW AMLRPMJJCB RFPMSEF LCE?RGTC DCCB@?AI, GL UFGAF PGQGLE JCTCJQ MD ? FMPKMLC GLFG@GR GRQ DSPRFCP PCJC?QC. 5FC RFPCC KCAF?LGQKQ MD FMPKML?J PCJC?QC ?PC FSKMP?J QRGKSJG, FMPKML?J QRGKSJG, ?LB LCSP?J QRGKSJG. )SKMP?J QRGKSJG PCDCPQ RM RFC AMLRPMJ MD FMPKML?J PCJC?QC GL PCQNMLQC RM AF?LECQ GL CVRP?ACJJSJ?P DJSGB JCTCJQ MP GML JCTCJQ. )MPKML?J QRGKSJG PCDCPQ RM RFC PCJC?QC MD FMPKMLCQ GL PCQNMLQC RM FMPKMLCQ PCJC?QCB @W MRFCP CLBMAPGLC EJ?LBQ. /CSP?J QRGKSJG PCDCPQ RM RFC PCJC?QC MD FMPKMLCQ GL PCQNMLQC RM LCSP?J QRGKSJ?RGML.
37.5 E;F:I@E< "C8E;J

5FC NGRSGR?PW EJ?LB GQ JMA?RCB ?R RFC @?QC MD RFC @P?GL ?LB GQ ?RR?AFCB RM RFC FWNMRF?J?KSQ @W RFC GLDSLBG@SJSK. 5FC ?LRCPGMP NGRSGR?PW PCACGTCQ NPMBSARQ DPMK RFC FWNMRF?J?KSQ @W RFC FWNMNFWQC?J NMPR?J QWQRCK ?LB NPMBSACQ QGV FMPKMLCQ. 5FC NMQRCPGMP NGRSGR?PW GQ ?L CVRCLQGML MD RFC @P?GL ?LB PCJC?QCQ FMPKMLCQ (?LRGBGSPCRGA FMPKMLC ?LB MVWRMAGL) NPMBSACB @W RFC FWNMRF?J?KSQ. 5FC RFWPMGB EJ?LB GQ JMA?RCB GL RFC LCAI ?LB GQ AMKNMQCB MD RUM JM@CQ AMLLCARCB @W RFC GQRFKSQ. 5FC RFWPMGB GQ K?BC SN MD DMJJGAJC ACJJQ RF?R NPMBSAC RFC FMPKMLCQ RFWPMVGLC ?LB RPGGMBMRFWPMLGLC. 1?P?DMJJGASJ?P ACJJQ MD RFC RFWPMGB NPMBSAC A?JAGRMLGL. 5FC N?P?RFWPMGB EJ?LBQ JGC ML RFC NMQRCPGMP QSPD?AC MD RFC RFWPMGB EJ?LB ?LB NPMBSAC N?P?RFWPMGB FMPKMLC. 5FC ?BPCL?J EJ?LBQ ?PC JMA?RCB ML RMN MD RFC IGBLCWQ ?LB AMLQGQR MD RFC PCL?J AMPRCV ?LB PCL?J KCBSJJ?. 5FC ?BPCL?J AMPRCV GQ RFC MSRCP N?PR MD RFC ?BPCL?J EJ?LB ?LB NPMBSACQ RFC AMPRGAMQRCPMGBQ, EJSAMAMPRGAMGBQ, ?LB KGLCP?JMAMPRGAMGBQ. 5FC ?BPCL?J KCBSJJ? GQ RFC GLLCP N?PR MD RFC ?BPCL?J EJ?LB ?LB NPMBSACQ RFC A?RCAFMJ?KGLCQ CNGLCNFPGLC ?LB LMPCNGLCNFPGLC. 5FC N?LAPC?Q JGCQ GL RFC ?@BMKCL @CRUCCL RFC QRMK?AF ?LB RFC QK?JJ GLRCQRGLC. $JSQRCPQ MD CLBMAPGLC ACJJQ GL RFC N?LAPC?Q DMPK RFC GQJCRQ MD -?LECPF?LQ, UFGAF ?PC AMKNMQCB MD ?JNF? ACJJQ RF?R PCJC?QC EJSA?EML ?LB @CR? ACJJQ RF?R PCJC?QC GLQSJGL. 4MKC MPE?LQ NMQQCQQ CLBMAPGLC ?ARGTGRW ?Q ? QCAMLB?PW DSLARGML @SR F?TC ?LMRFCP NPGK?PW DSLARGML. 5FC FC?PR NPMBSACQ RFC FMPKMLC ?RPG?J L?RPGSPCRGA NCNRGBC, UFGAF DSLARGMLQ RM PCBSAC @JMMB TMJSKC, NPCQQSPC, ?LB /?+ AMLACLRP?RGML. 5FC E?QRPMGLRCQRGL?J RP?AR NPMBSACQ T?PGMSQ FMPKMLCQ RF?R ?GB GL BGECQRGML. 5FC IGBLCWQ NPMBSAC PCLGL, A?JAGRPGMJ, ?LB CPWRFPMNMGCRGL. "BGNMQC RGQQSC NPMBSACQ JCNRGL, UFGAF NPMKMRCQ Q?RGCRW QGEL?JQ GL RFC @P?GL.
A-/ C*)) C/$*) ,0 ./$*).

1. Figure 37.5 )C?R QFMAI NPMRCGLQ ()41) ?PC QM L?KCB @CA?SQC RFCW FCJN PCDMJB KGQ-DMJBCB NPMRCGLQ. *L PCQNMLQC RM GLAPC?QCB RCKNCP?RSPC (? tFC?R QFMAIu), FC?R QFMAI NPMRCGLQ ?PC ?ARGT?RCB @W PCJC?QC DPMK RFC /3/)41 AMKNJCV. "R RFC Q?KC RGKC, RP?LQAPGNRGML MD )41 ECLCQ GQ ?ARGT?RCB. 8FW BM WMS RFGLI RFC ACJJ PCQNMLBQ RM ? FC?R QFMAI @W GLAPC?QGLE RFC ?ARGTGRW MD NPMRCGLQ RF?R FCJN PCDMJB KGQDMJBCB NPMRCGLQ 2. Figure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igure 37.14 )WNCPRFWPMGBGQK GQ ? AMLBGRGML GL UFGAF RFC RFWPMGB EJ?LB GQ MTCP?ARGTC. )WNMRFWPMGBGQK GQ ? AMLBGRGML GL UFGAF RFC RFWPMGB EJ?LB GQ SLBCP?ARGTC. 8FGAF MD RFC AMLBGRGMLQ ?PC RFC DMJJMUGLE RUM N?RGCLRQ KMQR JGICJW RM F?TC 1?RGCLR " F?Q QWKNRMKQ GLAJSBGLE UCGEFR E?GL, AMJB QCLQGRGTGRW, JMU FC?PR P?RC ?LB D?RGESC. 1?RGCLR # F?Q QWKNRMKQ GLAJSBGLE UCGEFR JMQQ, NPMDSQC QUC?RGLE, GLAPC?QCB FC?PR P?RC ?LB BGDDGASJRW QJCCNGLE.
- 1$ 2 ,0 ./$*).
4. " LCUJW BGQAMTCPCB FMPKMLC AMLR?GLQ DMSP ?KGLM ?AGBQ JGLICB RMECRFCP. 6LBCP UFGAF AFCKGA?J AJ?QQ UMSJB RFGQ FMPKMLC @C AJ?QQGDGCB >. JGNGB-BCPGTCB FMPKMLC
1092
#(!04%2 37 X 4(% %.$/#2).% 3934%-
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
14. 8FGAF CLBMAPGLC EJ?LBQ ?PC ?QQMAG?RCB UGRF RFC IGBLCWQ >. RFWPMGB EJ?LBQ ?. NGRSGR?PW EJ?LBQ @. ?BPCL?J EJ?LBQ A. EML?BQ 15. 8FGAF MD RFC DMJJMUGLE FMPKMLCQ GQ LMR NPMBSACB @W RFC ?LRCPGMP NGRSGR?PW >. MVWRMAGL ?. EPMURF FMPKMLC @. NPMJ?ARGL A. RFWPMGB-QRGKSJ?RGLE FMPKMLC
C-$/$CA' /#$)&$)" ,0 ./$*).

16. "JRFMSEF RFCPC ?PC K?LW BGDDCPCLR FMPKMLCQ GL RFC FSK?L @MBW, RFCW A?L @C BGTGBCB GLRM RFPCC AJ?QQCQ @?QCB ML RFCGP AFCKGA?J QRPSARSPC. 8F?R ?PC RFCQC AJ?QQCQ ?LB UF?R GQ MLC D?ARMP RF?R BGQRGLESGQFCQ RFCK 17. 8FCPC GQ GLQSJGL QRMPCB, ?LB UFW UMSJB GR @C PCJC?QCB 18. /?KC RUM GKNMPR?LR DSLARGMLQ MD FMPKMLC PCACNRMPQ. 19. )MU A?L FMPKMLCQ KCBG?RC AF?LECQ 20. /?KC ?LB BCQAPG@C ? DSLARGML MD MLC FMPKMLC NPMBSACB @W RFC ?LRCPGMP NGRSGR?PW ?LB MLC FMPKMLC NPMBSACB @W RFC NMQRCPGMP NGRSGR?PW. 21. %CQAPG@C MLC BGPCAR ?ARGML MD EPMURF FMPKMLC (()). 22. )MU GQ FMPKMLC NPMBSARGML ?LB PCJC?QC NPGK?PGJW AMLRPMJJCB 23. $MKN?PC ?LB AMLRP?QR FMPKML?J ?LB FSKMP?J QRGKSJG. 24. 8F?R BMCQ ?JBMQRCPMLC PCESJ?RC, ?LB FMU GQ GR QRGKSJ?RCB 25. 5FC ?BPCL?J KCBSJJ? AMLR?GLQ RUM RWNCQ MD QCAPCRMPW ACJJQ, UF?R ?PC RFCW ?LB UF?R ?PC RFCGP DSLARGMLQ
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1093
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38.1: /PG<J F= .B<C<K8C .PJK<DJ 38.2: BFE< 38.3: %F@EKJ 8E; .B<C<K8C (FM<D<EK 38.4: (LJ:C< CFEKI8:K@FE 8E; 'F:FDFK@FE
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5FC KSQASJ?P ?LB QICJCR?J QWQRCKQ NPMTGBC QSNNMPR RM RFC @MBW ?LB ?JJMU DMP ? UGBC P?LEC MD KMTCKCLR. 5FC @MLCQ MD RFC QICJCR?J QWQRCK NPMRCAR RFC @MBWsQ GLRCPL?J MPE?LQ ?LB QSNNMPR RFC UCGEFR MD RFC @MBW. 5FC KSQAJCQ MD RFC KSQASJ?P QWQRCK AMLRP?AR ?LB NSJJ ML RFC @MLCQ, ?JJMUGLE DMP KMTCKCLRQ ?Q BGTCPQC ?Q QR?LBGLE, U?JIGLE, PSLLGLE, ?LB EP?QNGLE GRCKQ. *LHSPW MP BGQC?QC ?DDCARGLE RFC KSQASJMQICJCR?J QWQRCK A?L @C TCPW BC@GJGR?RGLE. *L FSK?LQ, RFC KMQR AMKKML KSQASJMQICJCR?J BGQC?QCQ UMPJBUGBC ?PC A?SQCB @W K?JLSRPGRGML. "GJKCLRQ RF?R ?DDCAR RFC HMGLRQ ?PC ?JQM UGBCQNPC?B, QSAF ?Q ?PRFPGRGQ, UFGAF A?L K?IC KMTCKCLR BGDDGASJR ?LBqGL ?BT?LACB A?QCQqAMKNJCRCJW GKN?GP KM@GJGRW. *L QCTCPC A?QCQ GL UFGAF RFC HMGLR F?Q QSDDCPCB CVRCLQGTC B?K?EC, HMGLR PCNJ?ACKCLR QSPECPW K?W @C LCCBCB. 1PMEPCQQ GL RFC QAGCLAC MD NPMQRFCQGQ BCQGEL F?Q PCQSJRCB GL RFC BCTCJMNKCLR MD ?PRGDGAG?J HMGLRQ, UGRF HMGLR PCNJ?ACKCLR QSPECPW GL RFC FGNQ ?LB ILCCQ @CGLE RFC KMQR AMKKML. 3CNJ?ACKCLR HMGLRQ DMP QFMSJBCPQ, CJ@MUQ, ?LB DGLECPQ ?PC ?JQM ?T?GJ?@JC. &TCL UGRF RFGQ NPMEPCQQ, RFCPC GQ QRGJJ PMMK DMP
1094
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
GKNPMTCKCLR GL RFC BCQGEL MD NPMQRFCQCQ. 5FC QR?RC-MD-RFC-?PR NPMQRFCQCQ F?TC JGKGRCB BSP?@GJGRW ?LB RFCPCDMPC UC?P MSR OSGAIJW, N?PRGASJ?PJW GL WMSLE MP ?ARGTC GLBGTGBS?JQ. $SPPCLR PCQC?PAF GQ DMASQCB ML RFC SQC MD LCU K?RCPG?JQ, QSAF ?Q A?P@ML DG@CP, RF?R K?W K?IC NPMQRFCQCQ KMPC BSP?@JC.
38.1 R /PG<J F= .B<C<K8C .PJK<DJ

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %GQASQQ RFC BGDDCPCLR RWNCQ MD QICJCR?J QWQRCKQ z &VNJ?GL RFC PMJC MD RFC FSK?L QICJCR?J QWQRCK z $MKN?PC ?LB AMLRP?QR BGDDCPCLR QICJCR?J QWQRCKQ " QICJCR?J QWQRCK GQ LCACQQ?PW RM QSNNMPR RFC @MBW, NPMRCAR GLRCPL?J MPE?LQ, ?LB ?JJMU DMP RFC KMTCKCLR MD ?L MPE?LGQK. 5FCPC ?PC RFPCC BGDDCPCLR QICJCRML BCQGELQ RF?R DSJDGJJ RFCQC DSLARGMLQ: FWBPMQR?RGA QICJCRML, CVMQICJCRML, ?LB CLBMQICJCRML.
#P;IFJK8K@: .B<C<KFE
" hydrostatic skeleton GQ ? QICJCRML DMPKCB @W ? DJSGB-DGJJCB AMKN?PRKCLR UGRFGL RFC @MBW, A?JJCB RFC AMCJMK. 5FC MPE?LQ MD RFC AMCJMK ?PC QSNNMPRCB @W RFC ?OSCMSQ DJSGB, UFGAF ?JQM PCQGQRQ CVRCPL?J AMKNPCQQGML. 5FGQ AMKN?PRKCLR GQ SLBCP FWBPMQR?RGA NPCQQSPC @CA?SQC MD RFC DJSGB ?LB QSNNMPRQ RFC MRFCP MPE?LQ MD RFC MPE?LGQK. 5FGQ RWNC MD QICJCR?J QWQRCK GQ DMSLB GL QMDR-@MBGCB ?LGK?JQ QSAF ?Q QC? ?LCKMLCQ, C?PRFUMPKQ, $LGB?PG?, ?LB MRFCP GLTCPRC@P?RCQ (Figure 38.2).
!@>LI< 38.2 4EB PHBIBQLK LC QEB OBA-HKL??BA PB> PQ>O (P96;69,(:;,9 305*200) FP >K BU>JMIB LC > EVAOLPQ>QF@ PHBIBQLK. (@OBAFQ: x!J>A>44y/7FHFJBAF> #LJJLKP)
.MTCKCLR GL ? FWBPMQR?RGA QICJCRML GQ NPMTGBCB @W KSQAJCQ RF?R QSPPMSLB RFC AMCJMK. 5FC KSQAJCQ GL ? FWBPMQR?RGA QICJCRML AMLRP?AR RM AF?LEC RFC QF?NC MD RFC AMCJMK; RFC NPCQQSPC MD RFC DJSGB GL RFC AMCJMK NPMBSACQ KMTCKCLR. 'MP CV?KNJC, C?PRFUMPKQ KMTC @W U?TCQ MD KSQASJ?P AMLRP?ARGMLQ MD RFC QICJCR?J KSQAJC MD RFC @MBW U?JJ FWBPMQR?RGA QICJCRML, A?JJCB NCPGQR?JQGQ, UFGAF ?JRCPL?RCJW QFMPRCL ?LB JCLERFCL RFC @MBW. -CLERFCLGLE RFC @MBW CVRCLBQ RFC ?LRCPGMP CLB MD RFC MPE?LGQK. .MQR MPE?LGQKQ F?TC ? KCAF?LGQK RM DGV RFCKQCJTCQ GL RFC QS@QRP?RC. 4FMPRCLGLE RFC KSQAJCQ RFCL BP?UQ RFC NMQRCPGMP NMPRGML MD RFC @MBW DMPU?PB. "JRFMSEF ? FWBPMQR?RGA QICJCRML GQ UCJJ-QSGRCB RM GLTCPRC@P?RC MPE?LGQKQ QSAF ?Q C?PRFUMPKQ ?LB QMKC ?OS?RGA MPE?LGQKQ, GR GQ LMR ?L CDDGAGCLR QICJCRML DMP RCPPCQRPG?J ?LGK?JQ.
OFJB<C<KFE
"L exoskeleton GQ ?L CVRCPL?J QICJCRML RF?R AMLQGQRQ MD ? F?PB CLA?QCKCLR ML RFC QSPD?AC MD ?L MPE?LGQK. 'MP CV?KNJC, RFC QFCJJQ MD AP?@Q ?LB GLQCARQ ?PC CVMQICJCRMLQ (Figure 38.3). 5FGQ QICJCRML RWNC NPMTGBCQ BCDCLAC ?E?GLQR NPCB?RMPQ, QSNNMPRQ RFC @MBW, ?LB ?JJMUQ DMP KMTCKCLR RFPMSEF RFC AMLRP?ARGML MD ?RR?AFCB KSQAJCQ. "Q UGRF TCPRC@P?RCQ, KSQAJCQ KSQR APMQQ ? HMGLR GLQGBC RFC CVMQICJCRML. 4FMPRCLGLE MD RFC KSQAJC AF?LECQ RFC PCJ?RGMLQFGN MD RFC RUM QCEKCLRQ MD RFC CVMQICJCRML. "PRFPMNMBQ QSAF ?Q AP?@Q ?LB JM@QRCPQ F?TC CVMQICJCRMLQ RF?R AMLQGQR MD 30p50 NCPACLR AFGRGL, ? NMJWQ?AAF?PGBC BCPGT?RGTC MD EJSAMQC RF?R GQ ? QRPMLE @SR DJCVG@JC K?RCPG?J. $FGRGL GQ QCAPCRCB @W RFC CNGBCPK?J ACJJQ. 5FC CVMQICJCRML GQ DSPRFCP QRPCLERFCLCB @W RFC ?BBGRGML MD A?JAGSK A?P@ML?RC GL MPE?LGQKQ QSAF ?Q RFC JM@QRCP. #CA?SQC RFC CVMQICJCRML GQ ?ACJJSJ?P, ?PRFPMNMBQ KSQR NCPGMBGA?JJW QFCB RFCGP CVMQICJCRMLQ @CA?SQC RFC CVMQICJCRML BMCQ LMR EPMU ?Q RFC MPE?LGQK EPMUQ.
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1095
!@>LI< 38.3 -RP@IBP >QQ>@EBA QL QEB BULPHBIBQLK LC QEB (>IILTBBK @O>? (G,*(9*05<: 8<(+9(;<:) >IILT FQ QL JLSB.
E;FJB<C<KFE
"L endoskeleton GQ ? QICJCRML RF?R AMLQGQRQ MD F?PB, KGLCP?JGXCB QRPSARSPCQ JMA?RCB UGRFGL RFC QMDR RGQQSC MD MPE?LGQKQ. "L CV?KNJC MD ? NPGKGRGTC CLBMQICJCR?J QRPSARSPC GQ RFC QNGASJCQ MD QNMLECQ. 5FC @MLCQ MD TCPRC@P?RCQ ?PC AMKNMQCB MD RGQQSCQ, UFCPC?Q QNMLECQ F?TC LM RPSC RGQQSCQ (Figure 38.4). &LBMQICJCRMLQ NPMTGBC QSNNMPR DMP RFC @MBW, NPMRCAR GLRCPL?J MPE?LQ, ?LB ?JJMU DMP KMTCKCLR RFPMSEF AMLRP?ARGML MD KSQAJCQ ?RR?AFCB RM RFC QICJCRML.
!@>LI< 38.4 4EB PHBIBQLKP LC ERJ>KP >KA ELOPBP >OB BU>JMIBP LC BKALPHBIBQLKP. (@OBAFQ: 2LPP -ROMEV)
5FC FSK?L QICJCRML GQ ?L CLBMQICJCRML RF?R AMLQGQRQ MD 206 @MLCQ GL RFC ?BSJR. *R F?Q DGTC K?GL DSLARGMLQ: NPMTGBGLE QSNNMPR RM RFC @MBW, QRMPGLE KGLCP?JQ ?LB JGNGBQ, NPMBSAGLE @JMMB ACJJQ, NPMRCARGLE GLRCPL?J MPE?LQ, ?LB ?JJMUGLE DMP KMTCKCLR. 5FC QICJCR?J QWQRCK GL TCPRC@P?RCQ GQ BGTGBCB GLRM RFC ?VG?J QICJCRML (UFGAF AMLQGQRQ MD RFC QISJJ, TCPRC@P?J AMJSKL, ?LB PG@ A?EC), ?LB RFC ?NNCLBGASJ?P QICJCRML (UFGAF AMLQGQRQ MD RFC QFMSJBCPQ, JGK@ @MLCQ, RFC NCARMP?J EGPBJC, ?LB RFC NCJTGA EGPBJC).
1096
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
7GQGR RFC interactive body (http://openstaxcollege.org/l/virt_skeleton) QGRC RM @SGJB ? TGPRS?J QICJCRML: QCJCAR "QICJCRML" ?LB AJGAI RFPMSEF RFC ?ARGTGRW RM NJ?AC C?AF @MLC.
#LD8E AO@8C .B<C<KFE

5FC axial skeleton DMPKQ RFC ACLRP?J ?VGQ MD RFC @MBW ?LB GLAJSBCQ RFC @MLCQ MD RFC QISJJ, MQQGAJCQ MD RFC KGBBJC C?P, FWMGB @MLC MD RFC RFPM?R, TCPRC@P?J AMJSKL, ?LB RFC RFMP?AGA A?EC (PG@A?EC) (Figure 38.5). 5FC DSLARGML MD RFC ?VG?J QICJCRML GQ RM NPMTGBC QSNNMPR ?LB NPMRCARGML DMP RFC @P?GL, RFC QNGL?J AMPB, ?LB RFC MPE?LQ GL RFC TCLRP?J @MBW A?TGRW. *R NPMTGBCQ ? QSPD?AC DMP RFC ?RR?AFKCLR MD KSQAJCQ RF?R KMTC RFC FC?B, LCAI, ?LB RPSLI, NCPDMPKQ PCQNGP?RMPW KMTCKCLRQ, ?LB QR?@GJGXCQ N?PRQ MD RFC ?NNCLBGASJ?P QICJCRML.
!@>LI< 38.5 4EB >UF>I PHBIBQLK @LKPFPQP LC QEB ?LKBP LC QEB PHRII, LPPF@IBP LC QEB JFAAIB B>O, EVLFA ?LKB, SBOQB?O>I @LIRJK, >KA OF? @>DB. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
".+ !0:11 5FC @MLCQ MD RFC skull QSNNMPR RFC QRPSARSPCQ MD RFC D?AC ?LB NPMRCAR RFC @P?GL. 5FC QISJJ AMLQGQRQ MD 22 @MLCQ, UFGAF ?PC BGTGBCB GLRM RUM A?RCEMPGCQ: AP?LG?J @MLCQ ?LB D?AG?J @MLCQ. 5FC cranial bones ?PC CGEFR @MLCQ RF?R DMPK RFC AP?LG?J A?TGRW, UFGAF CLAJMQCQ RFC @P?GL ?LB QCPTCQ ?Q ?L ?RR?AFKCLR QGRC DMP RFC KSQAJCQ MD RFC FC?B ?LB LCAI. 5FC CGEFR AP?LG?J @MLCQ ?PC RFC DPMLR?J @MLC, RUM N?PGCR?J @MLCQ, RUM RCKNMP?J @MLCQ, MAAGNGR?J @MLC, QNFCLMGB @MLC, ?LB RFC CRFKMGB @MLC. "JRFMSEF RFC @MLCQ BCTCJMNCB QCN?P?RCJW GL RFC CK@PWM ?LB DCRSQ, GL RFC ?BSJR, RFCW ?PC RGEFRJW DSQCB UGRF AMLLCARGTC RGQQSC ?LB ?BHMGLGLE @MLCQ BM LMR KMTC (Figure 38.6).
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1097
!@>LI< 38.6 4EB ?LKBP LC QEB PHRII PRMMLOQ QEB PQOR@QROBP LC QEB C>@B >KA MOLQB@Q QEB ?O>FK. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
5FC auditory ossicles MD RFC KGBBJC C?P RP?LQKGR QMSLBQ DPMK RFC ?GP ?Q TG@P?RGMLQ RM RFC DJSGB-DGJJCB AMAFJC?. 5FC ?SBGRMPW MQQGAJCQ AMLQGQR MD QGV @MLCQ: RUM K?JJCSQ @MLCQ, RUM GLASQ @MLCQ, ?LB RUM QR?NCQ ML C?AF QGBC. 5FCQC ?PC RFC QK?JJCQR @MLCQ GL RFC @MBW ?LB ?PC SLGOSC RM K?KK?JQ. 'MSPRCCL facial bones DMPK RFC D?AC, NPMTGBC A?TGRGCQ DMP RFC QCLQC MPE?LQ (CWCQ, KMSRF, ?LB LMQC), NPMRCAR RFC CLRP?LACQ RM RFC BGECQRGTC ?LB PCQNGP?RMPW RP?ARQ, ?LB QCPTC ?Q ?RR?AFKCLR NMGLRQ DMP D?AG?J KSQAJCQ. 5FC 14 D?AG?J @MLCQ ?PC RFC L?Q?J @MLCQ, RFC K?VGJJ?PW @MLCQ, XWEMK?RGA @MLCQ, N?J?RGLC, TMKCP, J?APGK?J @MLCQ, RFC GLDCPGMP L?Q?J AMLAF?C, ?LB RFC K?LBG@JC. "JJ MD RFCQC @MLCQ MAASP GL N?GPQ CVACNR DMP RFC K?LBG@JC ?LB RFC TMKCP (Figure 38.7).
!@>LI< 38.7 4EB @O>KF>I ?LKBP, FK@IRAFKD QEB COLKQ>I, M>OFBQ>I, >KA PMEBKLFA ?LKBP, @LSBO QEB QLM LC QEB EB>A. 4EB C>@F>I ?LKBP LC QEB PHRII CLOJ QEB C>@B >KA MOLSFAB @>SFQFBP CLO QEB BVBP, KLPB, >KA JLRQE.
"JRFMSEF GR GQ LMR DMSLB GL RFC QISJJ, RFC FWMGB @MLC GQ AMLQGBCPCB ? AMKNMLCLR MD RFC ?VG?J QICJCRML. 5FC hyoid bone JGCQ @CJMU RFC K?LBG@JC GL RFC DPMLR MD RFC LCAI. *R ?ARQ ?Q ? KMT?@JC @?QC DMP RFC RMLESC
1098
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
?LB GQ AMLLCARCB RM KSQAJCQ MD RFC H?U, J?PWLV, ?LB RMLESC. 5FC K?LBG@JC ?PRGASJ?RCQ UGRF RFC @?QC MD RFC QISJJ. 5FC K?LBG@JC AMLRPMJQ RFC MNCLGLE RM RFC ?GPU?W ?LB ESR. *L ?LGK?JQ UGRF RCCRF, RFC K?LBG@JC @PGLEQ RFC QSPD?ACQ MD RFC RCCRF GL AMLR?AR UGRF RFC K?VGJJ?PW RCCRF. ".+ $+79+(7'1 C41:23 5FC vertebral column, MP QNGL?J AMJSKL, QSPPMSLBQ ?LB NPMRCARQ RFC QNGL?J AMPB, QSNNMPRQ RFC FC?B, ?LB ?ARQ ?Q ?L ?RR?AFKCLR NMGLR DMP RFC PG@Q ?LB KSQAJCQ MD RFC @?AI ?LB LCAI. 5FC ?BSJR TCPRC@P?J AMJSKL AMKNPGQCQ 26 @MLCQ: RFC 24 TCPRC@P?C, RFC Q?APSK, ?LB RFC AMAAWV @MLCQ. *L RFC ?BSJR, RFC Q?APSK GQ RWNGA?JJW AMKNMQCB MD DGTC TCPRC@P?C RF?R DSQC GLRM MLC. 5FC AMAAWV GQ RWNGA?JJW 3p4 TCPRC@P?C RF?R DSQC GLRM MLC. "PMSLB RFC ?EC MD 70, RFC Q?APSK ?LB RFC AMAAWV K?W DSQC RMECRFCP. 8C @CEGL JGDC UGRF ?NNPMVGK?RCJW 33 TCPRC@P?C, @SR ?Q UC EPMU, QCTCP?J TCPRC@P?C DSQC RMECRFCP. 5FC ?BSJR TCPRC@P?C ?PC DSPRFCP BGTGBCB GLRM RFC 7 ACPTGA?J TCPRC@P?C, 12 RFMP?AGA TCPRC@P?C, ?LB 5 JSK@?P TCPRC@P?C (Figure 38.8).
!@>LI< 38.8 (>) 4EB SBOQB?O>I @LIRJK @LKPFPQP LC PBSBK @BOSF@>I SBOQB?O>B (#1t7) QTBISB QELO>@F@ SBOQB?O>B (4E1t12), CFSB IRJ?>O SBOQB?O>B (,1t5), QEB LP P>@ORJ, >KA QEB @L@@VU. (?) 3MFK>I @ROSBP FK@OB>PB QEB PQOBKDQE >KA CIBUF?FIFQV LC QEB PMFKB. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V 5TB 'FIIB ?>PBA LK LOFDFK>I TLOH ?V 'O>V'P !K>QLJV; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V .#), .)()
&?AF TCPRC@P?J @MBW F?Q ? J?PEC FMJC GL RFC ACLRCP RFPMSEF UFGAF RFC LCPTCQ MD RFC QNGL?J AMPB N?QQ. 5FCPC GQ ?JQM ? LMRAF ML C?AF QGBC RFPMSEF UFGAF RFC QNGL?J LCPTCQ, UFGAF QCPTC RFC @MBW ?R RF?R JCTCJ, A?L CVGR DPMK RFC QNGL?J AMPB. 5FC TCPRC@P?J AMJSKL GQ ?NNPMVGK?RCJW 71 AK (28 GLAFCQ) GL ?BSJR K?JC FSK?LQ ?LB GQ ASPTCB, UFGAF A?L @C QCCL DPMK ? QGBC TGCU. 5FC L?KCQ MD RFC QNGL?J ASPTCQ AMPPCQNMLB RM RFC PCEGML MD RFC QNGLC GL UFGAF RFCW MAASP. 5FC RFMP?AGA ?LB Q?AP?J ASPTCQ ?PC AMLA?TC (ASPTC GLU?PBQ PCJ?RGTC RM RFC DPMLR MD RFC @MBW) ?LB RFC ACPTGA?J ?LB JSK@?P ASPTCQ ?PC AMLTCV (ASPTC MSRU?PBQ PCJ?RGTC RM RFC DPMLR MD RFC @MBW). 5FC ?PAFCB ASPT?RSPC MD RFC TCPRC@P?J AMJSKL GLAPC?QCQ GRQ QRPCLERF ?LB DJCVG@GJGRW, ?JJMUGLE GR RM ?@QMP@ QFMAIQ JGIC ? QNPGLE (Figure 38.8). Intervertebral discs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thoracic cage, ?JQM ILMUL ?Q RFC PG@A?EC, GQ RFC QICJCRML MD RFC AFCQR, ?LB AMLQGQRQ MD RFC PG@Q, QRCPLSK, RFMP?AGA TCPRC@P?C, ?LB AMQR?J A?PRGJ?ECQ (Figure 38.9). 5FC RFMP?AGA A?EC CLAJMQCQ ?LB NPMRCARQ
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1099
RFC MPE?LQ MD RFC RFMP?AGA A?TGRW, GLAJSBGLE RFC FC?PR ?LB JSLEQ. *R ?JQM NPMTGBCQ QSNNMPR DMP RFC QFMSJBCP EGPBJCQ ?LB SNNCP JGK@Q, ?LB QCPTCQ ?Q RFC ?RR?AFKCLR NMGLR DMP RFC BG?NFP?EK, KSQAJCQ MD RFC @?AI, AFCQR, LCAI, ?LB QFMSJBCPQ. $F?LECQ GL RFC TMJSKC MD RFC RFMP?V CL?@JC @PC?RFGLE. 5FC sternum, MP @PC?QR@MLC, GQ ? JMLE, DJ?R @MLC JMA?RCB ?R RFC ?LRCPGMP MD RFC AFCQR. *R GQ DMPKCB DPMK RFPCC @MLCQ RF?R DSQC GL RFC ?BSJR. 5FC ribs ?PC 12 N?GPQ MD JMLE, ASPTCB @MLCQ RF?R ?RR?AF RM RFC RFMP?AGA TCPRC@P?C ?LB ASPTC RMU?PB RFC DPMLR MD RFC @MBW, DMPKGLE RFC PG@A?EC. $MQR?J A?PRGJ?ECQ AMLLCAR RFC ?LRCPGMP CLBQ MD RFC PG@Q RM RFC QRCPLSK, UGRF RFC CVACNRGML MD PG@ N?GPQ 11 ?LB 12, UFGAF ?PC DPCCDJM?RGLE PG@Q.
!@>LI< 38.9 4EB QELO>@F@ @>DB, LO OF? @>DB, MOLQB@QP QEB EB>OQ >KA QEB IRKDP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .#), .)()
#LD8E AGG<E;@:LC8I .B<C<KFE

5FC appendicular skeleton GQ AMKNMQCB MD RFC @MLCQ MD RFC SNNCP JGK@Q (UFGAF DSLARGML RM EP?QN ?LB K?LGNSJ?RC M@HCARQ) ?LB RFC JMUCP JGK@Q (UFGAF NCPKGR JMAMKMRGML). *R ?JQM GLAJSBCQ RFC NCARMP?J EGPBJC, MP QFMSJBCP EGPBJC, RF?R ?RR?AFCQ RFC SNNCP JGK@Q RM RFC @MBW, ?LB RFC NCJTGA EGPBJC RF?R ?RR?AFCQ RFC JMUCP JGK@Q RM RFC @MBW (Figure 38.10).
1100
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
!@>LI< 38.10 4EB >MMBKAF@RI>O PHBIBQLK FP @LJMLPBA LC QEB ?LKBP LC QEB MB@QLO>I IFJ?P (>OJ, CLOB>OJ, E>KA), QEB MBISF@ IFJ?P (QEFDE, IBD, CLLQ), QEB MB@QLO>I DFOAIB, >KA QEB MBISF@ DFOAIB. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
".+ P+)947'1 G/7*1+ 5FC pectoral girdle @MLCQ NPMTGBC RFC NMGLRQ MD ?RR?AFKCLR MD RFC SNNCP JGK@Q RM RFC ?VG?J QICJCRML. 5FC FSK?L NCARMP?J EGPBJC AMLQGQRQ MD RFC AJ?TGAJC (MP AMJJ?P@MLC) GL RFC ?LRCPGMP, ?LB RFC QA?NSJ? (MP QFMSJBCP @J?BCQ) GL RFC NMQRCPGMP (Figure 38.11).
!@>LI< 38.11 (>) 4EB MB@QLO>I DFOAIB FK MOFJ>QBP @LKPFPQP LC QEB @I>SF@IBP >KA P@>MRI>B. (?) 4EB MLPQBOFLO SFBT OBSB>IP QEB PMFKB LC QEB P@>MRI> QL TEF@E JRP@IB >QQ>@EBP.
5FC clavicles ?PC 4-QF?NCB @MLCQ RF?R NMQGRGML RFC ?PKQ ML RFC @MBW. 5FC AJ?TGAJCQ JGC FMPGXMLR?JJW ?APMQQ RFC DPMLR MD RFC RFMP?V (AFCQR) HSQR ?@MTC RFC DGPQR PG@. 5FCQC @MLCQ ?PC D?GPJW DP?EGJC ?LB ?PC QSQACNRG@JC RM DP?ARSPCQ. 'MP CV?KNJC, ? D?JJ UGRF RFC ?PKQ MSRQRPCRAFCB A?SQCQ RFC DMPAC RM @C RP?LQKGRRCB
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1101
RM RFC AJ?TGAJCQ, UFGAF A?L @PC?I GD RFC DMPAC GQ CVACQQGTC. 5FC AJ?TGAJC ?PRGASJ?RCQ UGRF RFC QRCPLSK ?LB RFC QA?NSJ?. 5FC scapulae ?PC DJ?R, RPG?LESJ?P @MLCQ RF?R ?PC JMA?RCB ?R RFC @?AI MD RFC NCARMP?J EGPBJC. 5FCW QSNNMPR RFC KSQAJCQ APMQQGLE RFC QFMSJBCP HMGLR. " PGBEC, A?JJCB RFC QNGLC, PSLQ ?APMQQ RFC @?AI MD RFC QA?NSJ? ?LB A?L C?QGJW @C DCJR RFPMSEF RFC QIGL (Figure 38.11). 5FC QNGLC MD RFC QA?NSJ? GQ ? EMMB CV?KNJC MD ? @MLW NPMRPSQGML RF?R D?AGJGR?RCQ ? @PM?B ?PC? MD ?RR?AFKCLR DMP KSQAJCQ RM @MLC. ".+ #55+7 L/2( 5FC SNNCP JGK@ AMLR?GLQ 30 @MLCQ GL RFPCC PCEGMLQ: RFC ?PK (QFMSJBCP RM CJ@MU), RFC DMPC?PK (SJL? ?LB P?BGSQ), ?LB RFC UPGQR ?LB F?LB (Figure 38.12).
!@>LI< 38.12 4EB RMMBO IFJ? @LKPFPQP LC QEB ERJBORP LC QEB RMMBO >OJ, QEB O>AFRP >KA RIK> LC QEB CLOB>OJ, BFDEQ ?LKBP LC QEB @>OMRP, CFSB ?LKBP LC QEB JBQ>@>OMRP, >KA 14 ?LKBP LC QEB ME>I>KDBP.
"L articulation GQ ?LW NJ?AC ?R UFGAF RUM @MLCQ ?PC HMGLCB. 5FC humerus GQ RFC J?PECQR ?LB JMLECQR @MLC MD RFC SNNCP JGK@ ?LB RFC MLJW @MLC MD RFC ?PK. *R ?PRGASJ?RCQ UGRF RFC QA?NSJ? ?R RFC QFMSJBCP ?LB UGRF RFC DMPC?PK ?R RFC CJ@MU. 5FC forearm CVRCLBQ DPMK RFC CJ@MU RM RFC UPGQR ?LB AMLQGQRQ MD RUM @MLCQ: RFC SJL? ?LB RFC P?BGSQ. 5FC radius GQ JMA?RCB ?JMLE RFC J?RCP?J (RFSK@) QGBC MD RFC DMPC?PK ?LB ?PRGASJ?RCQ UGRF RFC FSKCPSQ ?R RFC CJ@MU. 5FC ulna GQ JMA?RCB ML RFC KCBG?J ?QNCAR (NGLIW-DGLECP QGBC) MD RFC DMPC?PK. *R GQ JMLECP RF?L RFC P?BGSQ. 5FC SJL? ?PRGASJ?RCQ UGRF RFC FSKCPSQ ?R RFC CJ@MU. 5FC P?BGSQ ?LB SJL? ?JQM ?PRGASJ?RC UGRF RFC A?PN?J @MLCQ ?LB UGRF C?AF MRFCP, UFGAF GL TCPRC@P?RCQ CL?@JCQ ? T?PG?@JC BCEPCC MD PMR?RGML MD RFC A?PNSQ UGRF PCQNCAR RM RFC JMLE ?VGQ MD RFC JGK@. 5FC F?LB GLAJSBCQ RFC CGEFR @MLCQ MD RFC carpus (UPGQR), RFC DGTC @MLCQ MD RFC metacarpus (N?JK), ?LB RFC 14 @MLCQ MD RFC phalanges (BGEGRQ). &?AF BGEGR AMLQGQRQ MD RFPCC NF?J?LECQ, CVACNR DMP RFC RFSK@, UFCL NPCQCLR, UFGAF F?Q MLJW RUM. ".+ P+1;/) G/7*1+ 5FC pelvic girdle ?RR?AFCQ RM RFC JMUCP JGK@Q MD RFC ?VG?J QICJCRML. #CA?SQC GR GQ PCQNMLQG@JC DMP @C?PGLE RFC UCGEFR MD RFC @MBW ?LB DMP JMAMKMRGML, RFC NCJTGA EGPBJC GQ QCASPCJW ?RR?AFCB RM RFC ?VG?J QICJCRML @W QRPMLE JGE?KCLRQ. *R ?JQM F?Q BCCN QMAICRQ UGRF PM@SQR JGE?KCLRQ RM QCASPCJW ?RR?AF RFC DCKSP RM RFC @MBW. 5FC NCJTGA EGPBJC GQ DSPRFCP QRPCLERFCLCB @W RUM J?PEC FGN @MLCQ. *L ?BSJRQ, RFC FGN @MLCQ, MP coxal bones, ?PC DMPKCB @W RFC DSQGML MD RFPCC N?GPQ MD @MLCQ: RFC GJGSK, GQAFGSK, ?LB NS@GQ. 5FC NCJTGQ HMGLQ RMECRFCP GL RFC ?LRCPGMP MD RFC @MBW ?R ? HMGLR A?JJCB RFC NS@GA QWKNFWQGQ ?LB UGRF RFC @MLCQ MD RFC Q?APSK ?R RFC NMQRCPGMP MD RFC @MBW. 5FC DCK?JC NCJTGQ GQ QJGEFRJW BGDDCPCLR DPMK RFC K?JC NCJTGQ. 0TCP ECLCP?RGMLQ MD CTMJSRGML, DCK?JCQ UGRF ? UGBCP NS@GA ?LEJC ?LB J?PECP BG?KCRCP NCJTGA A?L?J PCNPMBSACB KMPC QSAACQQDSJJW. 5FCPCDMPC, RFCGP MDDQNPGLE ?JQM F?B NCJTGA ?L?RMKW RF?R CL?@JCB QSAACQQDSJ AFGJB@GPRF (Figure 38.13).
1102
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
!@>LI< 38.13 4L >A>MQ QL OBMOLAR@QFSB CFQKBPP, QEB (>) CBJ>IB MBISFP FP IFDEQBO, TFABO, PE>IILTBO, >KA E>P > ?OL>ABO >KDIB ?BQTBBK QEB MR?F@ ?LKBP QE>K (?) QEB J>IB MBISFP.
".+ L4<+7 L/2( 5FC lower limb AMLQGQRQ MD RFC RFGEF, RFC JCE, ?LB RFC DMMR. 5FC @MLCQ MD RFC JMUCP JGK@ ?PC RFC DCKSP (RFGEF @MLC), N?RCJJ? (ILCCA?N), RG@G? ?LB DG@SJ? (@MLCQ MD RFC JCE), R?PQ?JQ (@MLCQ MD RFC ?LIJC), ?LB KCR?R?PQ?JQ ?LB NF?J?LECQ (@MLCQ MD RFC DMMR) (Figure 38.14). 5FC @MLCQ MD RFC JMUCP JGK@Q ?PC RFGAICP ?LB QRPMLECP RF?L RFC @MLCQ MD RFC SNNCP JGK@Q @CA?SQC MD RFC LCCB RM QSNNMPR RFC CLRGPC UCGEFR MD RFC @MBW ?LB RFC PCQSJRGLE DMPACQ DPMK JMAMKMRGML. *L ?BBGRGML RM CTMJSRGML?PW DGRLCQQ, RFC @MLCQ MD ?L GLBGTGBS?J UGJJ PCQNMLB RM DMPACQ CVCPRCB SNML RFCK.
!@>LI< 38.14 4EB ILTBO IFJ? @LKPFPQP LC QEB QEFDE (CBJRO), HKBB@>M (M>QBII>), IBD (QF?F> >KA CF?RI>), >KHIB (Q>OP>IP), >KA CLLQ (JBQ>Q>OP>IP >KA ME>I>KDBP) ?LKBP.
5FC femur, MP RFGEF@MLC, GQ RFC JMLECQR, FC?TGCQR, ?LB QRPMLECQR @MLC GL RFC @MBW. 5FC DCKSP ?LB NCJTGQ DMPK RFC FGN HMGLR ?R RFC NPMVGK?J CLB. "R RFC BGQR?J CLB, RFC DCKSP, RG@G?, ?LB N?RCJJ? DMPK RFC ILCC HMGLR. 5FC patella, MP ILCCA?N, GQ ? RPG?LESJ?P @MLC RF?R JGCQ ?LRCPGMP RM RFC ILCC HMGLR. 5FC N?RCJJ? GQ CK@CBBCB GL RFC RCLBML MD RFC DCKMP?J CVRCLQMPQ (OS?BPGACNQ). *R GKNPMTCQ ILCC CVRCLQGML @W PCBSAGLE DPGARGML. 5FC tibia, MP QFGL@MLC, GQ ? J?PEC @MLC MD RFC JCE RF?R GQ JMA?RCB BGPCARJW @CJMU RFC ILCC. 5FC RG@G? ?PRGASJ?RCQ UGRF RFC DCKSP ?R GRQ NPMVGK?J CLB, UGRF RFC DG@SJ? ?LB RFC R?PQ?J @MLCQ ?R GRQ BGQR?J CLB. *R GQ RFC QCAMLB J?PECQR @MLC GL RFC FSK?L @MBW ?LB GQ PCQNMLQG@JC DMP RP?LQKGRRGLE RFC UCGEFR MD RFC @MBW DPMK RFC DCKSP RM RFC DMMR. 5FC fibula, MP A?JD @MLC, N?P?JJCJQ ?LB ?PRGASJ?RCQ UGRF RFC RG@G?. *R BMCQ LMR ?PRGASJ?RC UGRF RFC DCKSP ?LB BMCQ LMR @C?P UCGEFR. 5FC DG@SJ? ?ARQ ?Q ? QGRC DMP KSQAJC ?RR?AFKCLR ?LB DMPKQ RFC J?RCP?J N?PR MD RFC ?LIJC HMGLR. 5FC tarsals ?PC RFC QCTCL @MLCQ MD RFC ?LIJC. 5FC ?LIJC RP?LQKGRQ RFC UCGEFR MD RFC @MBW DPMK RFC RG@G? ?LB RFC DG@SJ? RM RFC DMMR. 5FC metatarsals ?PC RFC DGTC @MLCQ MD RFC DMMR. 5FC NF?J?LECQ ?PC RFC 14 @MLCQ MD RFC RMCQ. &?AF RMC AMLQGQRQ MD RFPCC NF?J?LECQ, CVACNR DMP RFC @GE RMC RF?R F?Q MLJW RUM
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1103
(Figure 38.15). 7?PG?RGMLQ CVGQR GL MRFCP QNCAGCQ; DMP CV?KNJC, RFC FMPQCsQ KCR?A?PN?JQ ?LB KCR?R?PQ?JQ ?PC MPGCLRCB TCPRGA?JJW ?LB BM LMR K?IC AMLR?AR UGRF RFC QS@QRP?RC.
!@>LI< 38.15 4EFP AO>TFKD PELTP QEB ?LKBP LC QEB ERJ>K CLLQ >KA >KHIB, FK@IRAFKD QEB JBQ>Q>OP>IP >KA QEB ME>I>KDBP.
MFCLK@FE F= BF;P D<J@>E =FI 'F:FDFK@FE FE '8E;

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w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xMRPE-RMP.y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
1104
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
38.2 R BFE<
#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z $J?QQGDW RFC BGDDCPCLR RWNCQ MD @MLCQ GL RFC QICJCRML z &VNJ?GL RFC PMJC MD RFC BGDDCPCLR ACJJ RWNCQ GL @MLC z &VNJ?GL FMU @MLC DMPKQ BSPGLE BCTCJMNKCLR Bone, MP osseous tissue, GQ ? AMLLCARGTC RGQQSC RF?R AMLQRGRSRCQ RFC CLBMQICJCRML. *R AMLR?GLQ QNCAG?JGXCB ACJJQ ?LB ? K?RPGV MD KGLCP?J Q?JRQ ?LB AMJJ?ECL DG@CPQ. 5FC KGLCP?J Q?JRQ NPGK?PGJW GLAJSBC FWBPMVW?N?RGRC, ? KGLCP?J DMPKCB DPMK A?JAGSK NFMQNF?RC. Calcification GQ RFC NPMACQQ MD BCNMQGRGML MD KGLCP?J Q?JRQ ML RFC AMJJ?ECL DG@CP K?RPGV RF?R APWQR?JJGXCQ ?LB F?PBCLQ RFC RGQQSC. 5FC NPMACQQ MD A?JAGDGA?RGML MLJW MAASPQ GL RFC NPCQCLAC MD AMJJ?ECL DG@CPQ. 5FC @MLCQ MD RFC FSK?L QICJCRML ?PC AJ?QQGDGCB @W RFCGP QF?NC: JMLE @MLCQ, QFMPR @MLCQ, DJ?R @MLCQ, QSRSP?J @MLCQ, QCQ?KMGB @MLCQ, ?LB GPPCESJ?P @MLCQ (Figure 38.16).
!@>LI< 38.16 3ELTK >OB AFCCBOBKQ QVMBP LC ?LKBP: CI>Q, FOOBDRI>O, ILKD, PELOQ, >KA PBP>JLFA.
Long bones ?PC JMLECP RF?L RFCW ?PC UGBC ?LB F?TC ? QF?DR ?LB RUM CLBQ. 5FC diaphysis, MP ACLRP?J QF?DR, AMLR?GLQ @MLC K?PPMU GL ? K?PPMU A?TGRW. 5FC PMSLBCB CLBQ, RFC epiphyses, ?PC AMTCPCB UGRF ?PRGASJ?P A?PRGJ?EC ?LB ?PC DGJJCB UGRF PCB @MLC K?PPMU, UFGAF NPMBSACQ @JMMB ACJJQ (Figure 38.17). .MQR MD RFC JGK@ @MLCQ ?PC JMLE @MLCQqDMP CV?KNJC, RFC DCKSP, RG@G?, SJL?, ?LB P?BGSQ. &VACNRGMLQ RM RFGQ GLAJSBC RFC N?RCJJ? ?LB RFC @MLCQ MD RFC UPGQR ?LB ?LIJC.
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1105
!@>LI< 38.17 4EB ILKD ?LKB FP @LSBOBA ?V >OQF@RI>O @>OQFI>DB >Q BFQEBO BKA >KA @LKQ>FKP ?LKB J>OOLT (PELTK FK VBIILT FK QEFP FIIRPQO>QFLK) FK QEB J>OOLT @>SFQV.
Short bones, MP AS@MGB?J @MLCQ, ?PC @MLCQ RF?R ?PC RFC Q?KC UGBRF ?LB JCLERF, EGTGLE RFCK ? AS@CJGIC QF?NC. 'MP CV?KNJC, RFC @MLCQ MD RFC UPGQR (A?PN?JQ) ?LB ?LIJC (R?PQ?JQ) ?PC QFMPR @MLCQ (Figure 38.16). Flat bones ?PC RFGL ?LB PCJ?RGTCJW @PM?B @MLCQ RF?R ?PC DMSLB UFCPC CVRCLQGTC NPMRCARGML MD MPE?LQ GQ PCOSGPCB MP UFCPC @PM?B QSPD?ACQ MD KSQAJC ?RR?AFKCLR ?PC PCOSGPCB. &V?KNJCQ MD DJ?R @MLCQ ?PC RFC QRCPLSK (@PC?QR @MLC), PG@Q, QA?NSJ?C (QFMSJBCP @J?BCQ), ?LB RFC PMMD MD RFC QISJJ (Figure 38.16). Irregular bones ?PC @MLCQ UGRF AMKNJCV QF?NCQ. 5FCQC @MLCQ K?W F?TC QFMPR, DJ?R, LMRAFCB, MP PGBECB QSPD?ACQ. &V?KNJCQ MD GPPCESJ?P @MLCQ ?PC RFC TCPRC@P?C, FGN @MLCQ, ?LB QCTCP?J QISJJ @MLCQ. Sesamoid bones ?PC QK?JJ, DJ?R @MLCQ ?LB ?PC QF?NCB QGKGJ?PJW RM ? QCQ?KC QCCB. 5FC N?RCJJ?C ?PC QCQ?KMGB @MLCQ (Figure 38.18). 4CQ?KMGB @MLCQ BCTCJMN GLQGBC RCLBMLQ ?LB K?W @C DMSLB LC?P HMGLRQ ?R RFC ILCCQ, F?LBQ, ?LB DCCR.
!@>LI< 38.18 4EB M>QBII> LC QEB HKBB FP >K BU>JMIB LC > PBP>JLFA ?LKB.
Sutural bones ?PC QK?JJ, DJ?R, GPPCESJ?PJW QF?NCB @MLCQ. 5FCW K?W @C DMSLB @CRUCCL RFC DJ?R @MLCQ MD RFC QISJJ. 5FCW T?PW GL LSK@CP, QF?NC, QGXC, ?LB NMQGRGML.
1106
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
BFE< /@JJL<
#MLCQ ?PC AMLQGBCPCB MPE?LQ @CA?SQC RFCW AMLR?GL T?PGMSQ RWNCQ MD RGQQSC, QSAF ?Q @JMMB, AMLLCARGTC RGQQSC, LCPTCQ, ?LB @MLC RGQQSC. 0QRCMAWRCQ, RFC JGTGLE ACJJQ MD @MLC RGQQSC, DMPK RFC KGLCP?J K?RPGV MD @MLCQ. 5FCPC ?PC RUM RWNCQ MD @MLC RGQQSC: AMKN?AR ?LB QNMLEW. C425')9 B43+ "/88:+ Compact bone (MP AMPRGA?J @MLC) DMPKQ RFC F?PB CVRCPL?J J?WCP MD ?JJ @MLCQ ?LB QSPPMSLBQ RFC KCBSJJ?PW A?TGRW, MP @MLC K?PPMU. *R NPMTGBCQ NPMRCARGML ?LB QRPCLERF RM @MLCQ. $MKN?AR @MLC RGQQSC AMLQGQRQ MD SLGRQ A?JJCB MQRCMLQ MP )?TCPQG?L QWQRCKQ. Osteons ?PC AWJGLBPGA?J QRPSARSPCQ RF?R AMLR?GL ? KGLCP?J K?RPGV ?LB JGTGLE MQRCMAWRCQ AMLLCARCB @W A?L?JGASJG, UFGAF RP?LQNMPR @JMMB. 5FCW ?PC ?JGELCB N?P?JJCJ RM RFC JMLE ?VGQ MD RFC @MLC. &?AF MQRCML AMLQGQRQ MD lamellae, UFGAF ?PC J?WCPQ MD AMKN?AR K?RPGV RF?R QSPPMSLB ? ACLRP?J A?L?J A?JJCB RFC )?TCPQG?L A?L?J. 5FC Haversian canal (MQRCMLGA A?L?J) AMLR?GLQ RFC @MLCsQ @JMMB TCQQCJQ ?LB LCPTC DG@CPQ (Figure 38.19). 0QRCMLQ GL AMKN?AR @MLC RGQQSC ?PC ?JGELCB GL RFC Q?KC BGPCARGML ?JMLE JGLCQ MD QRPCQQ ?LB FCJN RFC @MLC PCQGQR @CLBGLE MP DP?ARSPGLE. 5FCPCDMPC, AMKN?AR @MLC RGQQSC GQ NPMKGLCLR GL ?PC?Q MD @MLC ?R UFGAF QRPCQQCQ ?PC ?NNJGCB GL MLJW ? DCU BGPCARGMLQ.
!@>LI< 38.19 #LJM>@Q ?LKB QFPPRB @LKPFPQP LC LPQBLKP QE>Q >OB >IFDKBA M>O>IIBI QL QEB ILKD >UFP LC QEB ?LKB, >KA QEB (>SBOPF>K @>K>I QE>Q @LKQ>FKP QEB ?LKBwP ?ILLA SBPPBIP >KA KBOSB CF?BOP. 4EB FKKBO I>VBO LC ?LKBP @LKPFPQP LC PMLKDV ?LKB QFPPRB. 4EB PJ>II A>OH LS>IP FK QEB LPQBLK OBMOBPBKQ QEB IFSFKD LPQBL@VQBP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .#), .)()
7EF@E LC QEB CLIILTFKD PQ>QBJBKQP >?LRQ ?LKB QFPPRB FP C>IPB >. #LJM>@Q ?LKB QFPPRB FP J>AB LC @VIFKAOF@>I LPQBLKP QE>Q >OB >IFDKBA PR@E QE>Q QEBV QO>SBI QEB IBKDQE LC QEB ?LKB. ?. (>SBOPF>K @>K>IP @LKQ>FK ?ILLA SBPPBIP LKIV. @. (>SBOPF>K @>K>IP @LKQ>FK ?ILLA SBPPBIP >KA KBOSB CF?BOP. A. 3MLKDV QFPPRB FP CLRKA LK QEB FKQBOFLO LC QEB ?LKB, >KA @LJM>@Q ?LKB QFPPRB FP CLRKA LK QEB BUQBOFLO. !543-> B43+ "/88:+ 8FCPC?Q AMKN?AR @MLC RGQQSC DMPKQ RFC MSRCP J?WCP MD ?JJ @MLCQ, spongy bone MP A?LACJJMSQ @MLC DMPKQ RFC GLLCP J?WCP MD ?JJ @MLCQ. 4NMLEW @MLC RGQQSC BMCQ LMR AMLR?GL MQRCMLQ RF?R AMLQRGRSRC AMKN?AR @MLC RGQQSC. *LQRC?B, GR AMLQGQRQ MD trabeculae, UFGAF ?PC J?KCJJ?C RF?R ?PC ?PP?LECB ?Q PMBQ MP NJ?RCQ. 3CB @MLC K?PPMU GQ DMSLB @CRUCCL RFC RP?@SASJ?C. #JMMB TCQQCJQ UGRFGL RFGQ RGQQSC BCJGTCP LSRPGCLRQ RM MQRCMAWRCQ ?LB PCKMTC U?QRC. 5FC PCB @MLC K?PPMU MD RFC DCKSP ?LB RFC GLRCPGMP MD MRFCP J?PEC @MLCQ, QSAF ?Q RFC GJCSK, DMPKQ @JMMB ACJJQ. 4NMLEW @MLC PCBSACQ RFC BCLQGRW MD @MLC ?LB ?JJMUQ RFC CLBQ MD JMLE @MLCQ RM AMKNPCQQ ?Q RFC PCQSJR MD QRPCQQCQ ?NNJGCB RM RFC @MLC. 4NMLEW @MLC GQ NPMKGLCLR GL ?PC?Q MD @MLCQ RF?R ?PC LMR FC?TGJW QRPCQQCB MP UFCPC QRPCQQCQ ?PPGTC DPMK K?LW BGPCARGMLQ. 5FC CNGNFWQCQ MD @MLCQ, QSAF ?Q RFC LCAI MD RFC DCKSP, ?PC QS@HCAR RM QRPCQQ DPMK K?LW BGPCARGMLQ. *K?EGLC J?WGLE ? FC?TW DP?KCB NGARSPC DJ?R ML RFC DJMMP. :MS
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1107
AMSJB FMJB SN MLC QGBC MD RFC NGARSPC UGRF ? RMMRFNGAI GD RFC RMMRFNGAI U?Q NCPNCLBGASJ?P RM RFC DJMMP ?LB RFC NGARSPC. /MU BPGJJ ? FMJC ?LB QRGAI RFC RMMRFNGAI GLRM RFC U?JJ RM F?LE SN RFC NGARSPC. *L RFGQ A?QC, RFC DSLARGML MD RFC RMMRFNGAI GQ RM RP?LQKGR RFC BMULU?PB NPCQQSPC MD RFC NGARSPC RM RFC U?JJ. 5FC DMPAC ML RFC NGARSPC GQ QRP?GEFR BMUL RM RFC DJMMP, @SR RFC DMPAC ML RFC RMMRFNGAI GQ @MRF RFC NGARSPC UGPC NSJJGLE BMUL ?LB RFC @MRRMK MD RFC FMJC GL RFC U?JJ NSQFGLE SN. 5FC RMMRFNGAI UGJJ @PC?I MDD PGEFR ?R RFC U?JJ. 5FC LCAI MD RFC DCKSP GQ FMPGXMLR?J JGIC RFC RMMRFNGAI GL RFC U?JJ. 5FC UCGEFR MD RFC @MBW NSQFCQ GR BMUL LC?P RFC HMGLR, @SR RFC TCPRGA?J BG?NFWQGQ MD RFC DCKSP NSQFCQ GR SN ?R RFC MRFCP CLB. 5FC LCAI MD RFC DCKSP KSQR @C QRPMLE CLMSEF RM RP?LQDCP RFC BMULU?PB DMPAC MD RFC @MBW UCGEFR FMPGXMLR?JJW RM RFC TCPRGA?J QF?DR MD RFC DCKSP (Figure 38.20).
!@>LI< 38.20 4O>?B@RI>B FK PMLKDV ?LKB >OB >OO>KDBA PR@E QE>Q LKB PFAB LC QEB ?LKB ?B>OP QBKPFLK >KA QEB LQEBO TFQEPQ>KAP @LJMOBPPFLK.
7GCU micrographs (http://openstaxcollege.org/l/muscle_tissue) MD KSQASJMQICJCR?J RGQQSCQ ?Q WMS PCTGCU RFC ?L?RMKW.
C<CC /PG<J @E BFE<J

#MLC AMLQGQRQ MD DMSP RWNCQ MD ACJJQ: MQRCM@J?QRQ, MQRCMAJ?QRQ, MQRCMAWRCQ, ?LB MQRCMNPMECLGRMP ACJJQ. Osteoblasts ?PC @MLC ACJJQ RF?R ?PC PCQNMLQG@JC DMP @MLC DMPK?RGML. 0QRCM@J?QRQ QWLRFCQGXC ?LB QCAPCRC RFC MPE?LGA N?PR ?LB GLMPE?LGA N?PR MD RFC CVRP?ACJJSJ?P K?RPGV MD @MLC RGQQSC, ?LB AMJJ?ECL DG@CPQ. 0QRCM@J?QRQ @CAMKC RP?NNCB GL RFCQC QCAPCRGMLQ ?LB BGDDCPCLRG?RC GLRM JCQQ ?ARGTC MQRCMAWRCQ. Osteoclasts ?PC J?PEC @MLC ACJJQ UGRF SN RM 50 LSAJCG. 5FCW PCKMTC @MLC QRPSARSPC @W PCJC?QGLE JWQMQMK?J CLXWKCQ ?LB ?AGBQ RF?R BGQQMJTC RFC @MLW K?RPGV. 5FCQC KGLCP?JQ, PCJC?QCB DPMK @MLCQ GLRM RFC @JMMB, FCJN PCESJ?RC A?JAGSK AMLACLRP?RGMLQ GL @MBW DJSGBQ. #MLC K?W ?JQM @C PCQMP@CB DMP PCKMBCJGLE, GD RFC ?NNJGCB QRPCQQCQ F?TC AF?LECB. Osteocytes ?PC K?RSPC @MLC ACJJQ ?LB ?PC RFC K?GL ACJJQ GL @MLW AMLLCARGTC RGQQSC; RFCQC ACJJQ A?LLMR BGTGBC. 0QRCMAWRCQ K?GLR?GL LMPK?J @MLC QRPSARSPC @W PCAWAJGLE RFC KGLCP?J Q?JRQ GL RFC @MLW K?RPGV. Osteoprogenitor cells ?PC QOS?KMSQ QRCK ACJJQ RF?R BGTGBC RM NPMBSAC B?SEFRCP ACJJQ RF?R BGDDCPCLRG?RC GLRM MQRCM@J?QRQ. 0QRCMNPMECLGRMP ACJJQ ?PC GKNMPR?LR GL RFC PCN?GP MD DP?ARSPCQ.
D<M<CFGD<EK F= BFE<
Ossification, MP MQRCMECLCQGQ, GQ RFC NPMACQQ MD @MLC DMPK?RGML @W MQRCM@J?QRQ. 0QQGDGA?RGML GQ BGQRGLAR DPMK RFC NPMACQQ MD A?JAGDGA?RGML; UFCPC?Q A?JAGDGA?RGML R?ICQ NJ?AC BSPGLE RFC MQQGDGA?RGML MD @MLCQ, GR A?L ?JQM MAASP GL MRFCP RGQQSCQ. 0QQGDGA?RGML @CEGLQ ?NNPMVGK?RCJW QGV UCCIQ ?DRCP DCPRGJGX?RGML GL ?L CK@PWM. #CDMPC RFGQ RGKC, RFC CK@PWMLGA QICJCRML AMLQGQRQ CLRGPCJW MD DG@PMSQ KCK@P?LCQ ?LB FW?JGLC A?PRGJ?EC.
1108
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
5FC BCTCJMNKCLR MD @MLC DPMK DG@PMSQ KCK@P?LCQ GQ A?JJCB GLRP?KCK@P?LMSQ MQQGDGA?RGML; BCTCJMNKCLR DPMK FW?JGLC A?PRGJ?EC GQ A?JJCB CLBMAFMLBP?J MQQGDGA?RGML. #MLC EPMURF AMLRGLSCQ SLRGJ ?NNPMVGK?RCJW ?EC 25. #MLCQ A?L EPMU GL RFGAILCQQ RFPMSEFMSR JGDC, @SR ?DRCP ?EC 25, MQQGDGA?RGML DSLARGMLQ NPGK?PGJW GL @MLC PCKMBCJGLE ?LB PCN?GP.
$EKI8D<D9I8EFLJ *JJ@=@:8K@FE
Intramembranous ossification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ndochondral ossification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epiphyseal plate (EPMURF NJ?RC), UFGAF GQ RFC PCEGML @CRUCCL RFC BG?NFWQGQ ?LB CNGNFWQGQ RF?R GQ PCQNMLQG@JC DMP RFC JCLERFUGQC EPMURF MD JMLE @MLCQ (Figure 38.21).
!@>LI< 38.21 %KAL@ELKAO>I LPPFCF@>QFLK FP QEB MOL@BPP LC ?LKB ABSBILMJBKQ COLJ EV>IFKB @>OQFI>DB. 4EB MBOFLPQBRJ FP QEB @LKKB@QFSB QFPPRB LK QEB LRQPFAB LC ?LKB QE>Q >@QP >P QEB FKQBOC>@B ?BQTBBK ?LKB, ?ILLA SBPPBIP, QBKALKP, >KA IFD>JBKQP.
"IFNK? F= BFE<
-MLE @MLCQ AMLRGLSC RM JCLERFCL, NMRCLRG?JJW SLRGJ ?BMJCQACLAC, RFPMSEF RFC ?BBGRGML MD @MLC RGQQSC ?R RFC CNGNFWQC?J NJ?RC. 5FCW ?JQM GLAPC?QC GL UGBRF RFPMSEF ?NNMQGRGML?J EPMURF. L+3-9.+3/3- 4, L43- B43+8 $FMLBPMAWRCQ ML RFC CNGNFWQC?J QGBC MD RFC CNGNFWQC?J NJ?RC BGTGBC; MLC ACJJ PCK?GLQ SLBGDDCPCLRG?RCB LC?P RFC CNGNFWQGQ, ?LB MLC ACJJ KMTCQ RMU?PB RFC BG?NFWQGQ. 5FC ACJJQ, UFGAF ?PC NSQFCB DPMK RFC CNGNFWQGQ, K?RSPC ?LB ?PC BCQRPMWCB @W A?JAGDGA?RGML. 5FGQ NPMACQQ PCNJ?ACQ A?PRGJ?EC UGRF @MLC ML RFC BG?NFWQC?J QGBC MD RFC NJ?RC, PCQSJRGLE GL ? JCLERFCLGLE MD RFC @MLC. -MLE @MLCQ QRMN EPMUGLE ?R ?PMSLB RFC ?EC MD 18 GL DCK?JCQ ?LB RFC ?EC MD 21 GL K?JCQ GL ? NPMACQQ A?JJCB CNGNFWQC?J NJ?RC AJMQSPC. %SPGLE RFGQ NPMACQQ, A?PRGJ?EC ACJJQ QRMN BGTGBGLE ?LB ?JJ MD RFC A?PRGJ?EC GQ
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1109
PCNJ?ACB @W @MLC. 5FC CNGNFWQC?J NJ?RC D?BCQ, JC?TGLE ? QRPSARSPC A?JJCB RFC CNGNFWQC?J JGLC MP CNGNFWQC?J PCKL?LR, ?LB RFC CNGNFWQGQ ?LB BG?NFWQGQ DSQC. "./)0+3/3- 4, L43- B43+8 Appositional growth GQ RFC GLAPC?QC GL RFC BG?KCRCP MD @MLCQ @W RFC ?BBGRGML MD @MLW RGQQSC ?R RFC QSPD?AC MD @MLCQ. 0QRCM@J?QRQ ?R RFC @MLC QSPD?AC QCAPCRC @MLC K?RPGV, ?LB MQRCMAJ?QRQ ML RFC GLLCP QSPD?AC @PC?I BMUL @MLC. 5FC MQRCM@J?QRQ BGDDCPCLRG?RC GLRM MQRCMAWRCQ. " @?J?LAC @CRUCCL RFCQC RUM NPMACQQCQ ?JJMUQ RFC @MLC RM RFGAICL UGRFMSR @CAMKGLE RMM FC?TW.
BFE< -<DF;<C@E> 8E; -<G8@I

#MLC PCLCU?J AMLRGLSCQ ?DRCP @GPRF GLRM ?BSJRFMMB. Bone remodeling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igure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
!@>LI< 38.22 !CQBO QEFP ?LKB FP PBQ, > @>IIRP TFII HKFQ QEB QTL BKAP QLDBQEBO. (@OBAFQ: "FII 2ELABP)
1110
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
D<:8C:@=@:8K@FE F= BFE<J
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
38.3 R %F@EKJ 8E; .B<C<K8C (FM<D<EK

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z $J?QQGDW RFC BGDDCPCLR RWNCQ MD HMGLRQ ML RFC @?QGQ MD QRPSARSPC z &VNJ?GL RFC PMJC MD HMGLRQ GL QICJCR?J KMTCKCLR 5FC NMGLR ?R UFGAF RUM MP KMPC @MLCQ KCCR GQ A?JJCB ? joint, MP articulation. +MGLRQ ?PC PCQNMLQG@JC DMP KMTCKCLR, QSAF ?Q RFC KMTCKCLR MD JGK@Q, ?LB QR?@GJGRW, QSAF ?Q RFC QR?@GJGRW DMSLB GL RFC @MLCQ MD RFC QISJJ.
CC8JJ@=@:8K@FE F= %F@EKJ FE K?< B8J@J F= .KIL:KLI<

5FCPC ?PC RUM U?WQ RM AJ?QQGDW HMGLRQ: ML RFC @?QGQ MD RFCGP QRPSARSPC MP ML RFC @?QGQ MD RFCGP DSLARGML. 5FC QRPSARSP?J AJ?QQGDGA?RGML BGTGBCQ HMGLRQ GLRM @MLW, DG@PMSQ, A?PRGJ?EGLMSQ, ?LB QWLMTG?J HMGLRQ BCNCLBGLE ML RFC K?RCPG?J AMKNMQGLE RFC HMGLR ?LB RFC NPCQCLAC MP ?@QCLAC MD ? A?TGRW GL RFC HMGLR. F/(74:8 J4/398 5FC @MLCQ MD fibrous joints ?PC FCJB RMECRFCP @W DG@PMSQ AMLLCARGTC RGQQSC. 5FCPC GQ LM A?TGRW, MP QN?AC, NPCQCLR @CRUCCL RFC @MLCQ ?LB QM KMQR DG@PMSQ HMGLRQ BM LMR KMTC ?R ?JJ, MP ?PC MLJW A?N?@JC MD KGLMP KMTCKCLRQ. 5FCPC ?PC RFPCC RWNCQ MD DG@PMSQ HMGLRQ: QSRSPCQ, QWLBCQKMQCQ, ?LB EMKNFMQCQ. Sutures ?PC DMSLB MLJW GL RFC QISJJ ?LB NMQQCQQ QFMPR DG@CPQ MD AMLLCARGTC RGQQSC RF?R FMJB RFC QISJJ @MLCQ RGEFRJW GL NJ?AC (Figure 38.23).
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1111
!@>LI< 38.23 3RQROBP >OB CF?OLRP GLFKQP CLRKA LKIV FK QEB PHRII.
Syndesmoses ?PC HMGLRQ GL UFGAF RFC @MLCQ ?PC AMLLCARCB @W ? @?LB MD AMLLCARGTC RGQQSC, ?JJMUGLE DMP KMPC KMTCKCLR RF?L GL ? QSRSPC. "L CV?KNJC MD ? QWLBCQKMQGQ GQ RFC HMGLR MD RFC RG@G? ?LB DG@SJ? GL RFC ?LIJC. 5FC ?KMSLR MD KMTCKCLR GL RFCQC RWNCQ MD HMGLRQ GQ BCRCPKGLCB @W RFC JCLERF MD RFC AMLLCARGTC RGQQSC DG@CPQ. Gomphoses MAASP @CRUCCL RCCRF ?LB RFCGP QMAICRQ; RFC RCPK PCDCPQ RM RFC U?W RFC RMMRF DGRQ GLRM RFC QMAICR JGIC ? NCE (Figure 38.24). 5FC RMMRF GQ AMLLCARCB RM RFC QMAICR @W ? AMLLCARGTC RGQQSC PCDCPPCB RM ?Q RFC NCPGMBMLR?J JGE?KCLR.
!@>LI< 38.24 'LJMELPBP >OB CF?OLRP GLFKQP ?BQTBBK QEB QBBQE >KA QEBFO PL@HBQP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V 'O>V'P !K>QLJV)
C'79/1'-/34:8 J4/398 Cartilaginous joints ?PC HMGLRQ GL UFGAF RFC @MLCQ ?PC AMLLCARCB @W A?PRGJ?EC. 5FCPC ?PC RUM RWNCQ MD A?PRGJ?EGLMSQ HMGLRQ: QWLAFMLBPMQCQ ?LB QWKNFWQCQ. *L ? synchondrosis, RFC @MLCQ ?PC HMGLCB @W FW?JGLC A?PRGJ?EC. 4WLAFMLBPMQCQ ?PC DMSLB GL RFC CNGNFWQC?J NJ?RCQ MD EPMUGLE @MLCQ GL AFGJBPCL. *L symphyses, FW?JGLC A?PRGJ?EC AMTCPQ RFC CLB MD RFC @MLC @SR RFC AMLLCARGML @CRUCCL @MLCQ MAASPQ RFPMSEF DG@PMA?PRGJ?EC. 4WKNFWQCQ ?PC DMSLB ?R RFC HMGLRQ @CRUCCL TCPRC@P?C. &GRFCP RWNC MD A?PRGJ?EGLMSQ HMGLR ?JJMUQ DMP TCPW JGRRJC KMTCKCLR. !>34;/'1 J4/398 Synovial joints ?PC RFC MLJW HMGLRQ RF?R F?TC ? QN?AC @CRUCCL RFC ?BHMGLGLE @MLCQ (Figure 38.25). 5FGQ QN?AC GQ PCDCPPCB RM ?Q RFC QWLMTG?J (MP HMGLR) A?TGRW ?LB GQ DGJJCB UGRF QWLMTG?J DJSGB. 4WLMTG?J DJSGB JS@PGA?RCQ RFC HMGLR, PCBSAGLE DPGARGML @CRUCCL RFC @MLCQ ?LB ?JJMUGLE DMP EPC?RCP KMTCKCLR. 5FC CLBQ MD RFC @MLCQ ?PC AMTCPCB UGRF ?PRGASJ?P A?PRGJ?EC, ? FW?JGLC A?PRGJ?EC, ?LB RFC CLRGPC HMGLR GQ QSPPMSLBCB @W ?L ?PRGASJ?P A?NQSJC AMKNMQCB MD AMLLCARGTC RGQQSC RF?R ?JJMUQ KMTCKCLR MD RFC HMGLR UFGJC PCQGQRGLE
1112
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
BGQJMA?RGML. "PRGASJ?P A?NQSJCQ K?W ?JQM NMQQCQQ JGE?KCLRQ RF?R FMJB RFC @MLCQ RMECRFCP. 4WLMTG?J HMGLRQ ?PC A?N?@JC MD RFC EPC?RCQR KMTCKCLR MD RFC RFPCC QRPSARSP?J HMGLR RWNCQ; FMUCTCP, RFC KMPC KM@GJC ? HMGLR, RFC UC?ICP RFC HMGLR. ,LCCQ, CJ@MUQ, ?LB QFMSJBCPQ ?PC CV?KNJCQ MD QWLMTG?J HMGLRQ.
!@>LI< 38.25 3VKLSF>I GLFKQP >OB QEB LKIV GLFKQP QE>Q E>SB > PM>@B LO xPVKLSF>I @>SFQVy FK QEB GLFKQ.
CC8JJ@=@:8K@FE F= %F@EKJ FE K?< B8J@J F= !LE:K@FE

5FC DSLARGML?J AJ?QQGDGA?RGML BGTGBCQ HMGLRQ GLRM RFPCC A?RCEMPGCQ: QWL?PRFPMQCQ, ?KNFG?PRFPMQCQ, ?LB BG?PRFPMQCQ. " synarthrosis GQ ? HMGLR RF?R GQ GKKMT?@JC. 5FGQ GLAJSBCQ QSRSPCQ, EMKNFMQCQ, ?LB QWLAFMLBPMQCQ. Amphiarthroses ?PC HMGLRQ RF?R ?JJMU QJGEFR KMTCKCLR, GLAJSBGLE QWLBCQKMQCQ ?LB QWKNFWQCQ. Diarthroses ?PC HMGLRQ RF?R ?JJMU DMP DPCC KMTCKCLR MD RFC HMGLR, ?Q GL QWLMTG?J HMGLRQ.
(FM<D<EK 8K .PEFM@8C %F@EKJ

5FC UGBC P?LEC MD KMTCKCLR ?JJMUCB @W QWLMTG?J HMGLRQ NPMBSACQ BGDDCPCLR RWNCQ MD KMTCKCLRQ. 5FC KMTCKCLR MD QWLMTG?J HMGLRQ A?L @C AJ?QQGDGCB ?Q MLC MD DMSP BGDDCPCLR RWNCQ: EJGBGLE, ?LESJ?P, PMR?RGML?J, MP QNCAG?J KMTCKCLR. G1/*/3- M4;+2+39 Gliding movements MAASP ?Q PCJ?RGTCJW DJ?R @MLC QSPD?ACQ KMTC N?QR C?AF MRFCP. (JGBGLE KMTCKCLRQ NPMBSAC TCPW JGRRJC PMR?RGML MP ?LESJ?P KMTCKCLR MD RFC @MLCQ. 5FC HMGLRQ MD RFC A?PN?J ?LB R?PQ?J @MLCQ ?PC CV?KNJCQ MD HMGLRQ RF?R NPMBSAC EJGBGLE KMTCKCLRQ. A3-:1'7 M4;+2+39 Angular movements ?PC NPMBSACB UFCL RFC ?LEJC @CRUCCL RFC @MLCQ MD ? HMGLR AF?LECQ. 5FCPC ?PC QCTCP?J BGDDCPCLR RWNCQ MD ?LESJ?P KMTCKCLRQ, GLAJSBGLE DJCVGML, CVRCLQGML, FWNCPCVRCLQGML, ?@BSARGML, ?BBSARGML, ?LB AGPASKBSARGML. Flexion, MP @CLBGLE, MAASPQ UFCL RFC ?LEJC @CRUCCL RFC @MLCQ BCAPC?QCQ. .MTGLE RFC DMPC?PK SNU?PB ?R RFC CJ@MU MP KMTGLE RFC UPGQR RM KMTC RFC F?LB RMU?PB RFC DMPC?PK ?PC CV?KNJCQ MD DJCVGML. Extension GQ RFC MNNMQGRC MD DJCVGML GL RF?R RFC ?LEJC @CRUCCL RFC @MLCQ MD ? HMGLR GLAPC?QCQ. 4RP?GEFRCLGLE ? JGK@ ?DRCP DJCVGML GQ ?L CV?KNJC MD CVRCLQGML. &VRCLQGML N?QR RFC PCESJ?P ?L?RMKGA?J NMQGRGML GQ PCDCPPCB RM ?Q hyperextension. 5FGQ GLAJSBCQ KMTGLE RFC LCAI @?AI RM JMMI SNU?PB, MP @CLBGLE RFC UPGQR QM RF?R RFC F?LB KMTCQ ?U?W DPMK RFC DMPC?PK. Abduction MAASPQ UFCL ? @MLC KMTCQ ?U?W DPMK RFC KGBJGLC MD RFC @MBW. &V?KNJCQ MD ?@BSARGML ?PC KMTGLE RFC ?PKQ MP JCEQ J?RCP?JJW RM JGDR RFCK QRP?GEFR MSR RM RFC QGBC. Adduction GQ RFC KMTCKCLR MD ? @MLC RMU?PB RFC KGBJGLC MD RFC @MBW. .MTCKCLR MD RFC JGK@Q GLU?PB ?DRCP ?@BSARGML GQ ?L CV?KNJC MD ?BBSARGML. Circumduction GQ RFC KMTCKCLR MD ? JGK@ GL ? AGPASJ?P KMRGML, ?Q GL KMTGLE RFC ?PK GL ? AGPASJ?P KMRGML.
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1113
49'9/43'1 M4;+2+39 Rotational movement GQ RFC KMTCKCLR MD ? @MLC ?Q GR PMR?RCQ ?PMSLB GRQ JMLEGRSBGL?J ?VGQ. 3MR?RGML A?L @C RMU?PB RFC KGBJGLC MD RFC @MBW, UFGAF GQ PCDCPPCB RM ?Q medial rotation, MP ?U?W DPMK RFC KGBJGLC MD RFC @MBW, UFGAF GQ PCDCPPCB RM ?Q lateral rotation. .MTCKCLR MD RFC FC?B DPMK QGBC RM QGBC GQ ?L CV?KNJC MD PMR?RGML. !5+)/'1 M4;+2+398 4MKC KMTCKCLRQ RF?R A?LLMR @C AJ?QQGDGCB ?Q EJGBGLE, ?LESJ?P, MP PMR?RGML?J ?PC A?JJCB QNCAG?J KMTCKCLRQ. Inversion GLTMJTCQ RFC QMJCQ MD RFC DCCR KMTGLE GLU?PB, RMU?PB RFC KGBJGLC MD RFC @MBW. Eversion GQ RFC MNNMQGRC MD GLTCPQGML, KMTCKCLR MD RFC QMJC MD RFC DMMR MSRU?PB, ?U?W DPMK RFC KGBJGLC MD RFC @MBW. Protraction GQ RFC ?LRCPGMP KMTCKCLR MD ? @MLC GL RFC FMPGXMLR?J NJ?LC. Retraction MAASPQ ?Q ? HMGLR KMTCQ @?AI GLRM NMQGRGML ?DRCP NPMRP?ARGML. 1PMRP?ARGML ?LB PCRP?ARGML A?L @C QCCL GL RFC KMTCKCLR MD RFC K?LBG@JC ?Q RFC H?U GQ RFPSQR MSRU?PBQ ?LB RFCL @?AI GLU?PBQ. Elevation GQ RFC KMTCKCLR MD ? @MLC SNU?PB, QSAF ?Q UFCL RFC QFMSJBCPQ ?PC QFPSEECB, JGDRGLE RFC QA?NSJ?C. Depression GQ RFC MNNMQGRC MD CJCT?RGMLqKMTCKCLR BMULU?PB MD ? @MLC, QSAF ?Q ?DRCP RFC QFMSJBCPQ ?PC QFPSEECB ?LB RFC QA?NSJ?C PCRSPL RM RFCGP LMPK?J NMQGRGML DPMK ?L CJCT?RCB NMQGRGML. Dorsiflexion GQ ? @CLBGLE ?R RFC ?LIJC QSAF RF?R RFC RMCQ ?PC JGDRCB RMU?PB RFC ILCC. Plantar flexion GQ ? @CLBGLE ?R RFC ?LIJC UFCL RFC FCCJ GQ JGDRCB, QSAF ?Q UFCL QR?LBGLE ML RFC RMCQ. Supination GQ RFC KMTCKCLR MD RFC P?BGSQ ?LB SJL? @MLCQ MD RFC DMPC?PK QM RF?R RFC N?JK D?ACQ DMPU?PB. Pronation GQ RFC MNNMQGRC KMTCKCLR, GL UFGAF RFC N?JK D?ACQ @?AIU?PB. Opposition GQ RFC KMTCKCLR MD RFC RFSK@ RMU?PB RFC DGLECPQ MD RFC Q?KC F?LB, K?IGLE GR NMQQG@JC RM EP?QN ?LB FMJB M@HCARQ.
/PG<J F= .PEFM@8C %F@EKJ

4WLMTG?J HMGLRQ ?PC DSPRFCP AJ?QQGDGCB GLRM QGV BGDDCPCLR A?RCEMPGCQ ML RFC @?QGQ MD RFC QF?NC ?LB QRPSARSPC MD RFC HMGLR. 5FC QF?NC MD RFC HMGLR ?DDCARQ RFC RWNC MD KMTCKCLR NCPKGRRCB @W RFC HMGLR (Figure 38.26). 5FCQC HMGLRQ A?L @C BCQAPG@CB ?Q NJ?L?P, FGLEC, NGTMR, AMLBWJMGB, Q?BBJC, MP @?JJ-?LB-QMAICR HMGLRQ.
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#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
!@>LI< 38.26 $FCCBOBKQ QVMBP LC GLFKQP >IILT AFCCBOBKQ QVMBP LC JLSBJBKQ. 0I>K>O, EFKDB, MFSLQ, @LKAVILFA, P>AAIB, >KA ?>II->KA-PL@HBQ >OB >II QVMBP LC PVKLSF>I GLFKQP.
P1'3'7 J4/398 Planar joints F?TC @MLCQ UGRF ?PRGASJ?RGLE QSPD?ACQ RF?R ?PC DJ?R MP QJGEFRJW ASPTCB D?ACQ. 5FCQC HMGLRQ ?JJMU DMP EJGBGLE KMTCKCLRQ, ?LB QM RFC HMGLRQ ?PC QMKCRGKCQ PCDCPPCB RM ?Q EJGBGLE HMGLRQ. 5FC P?LEC MD KMRGML GQ JGKGRCB GL RFCQC HMGLRQ ?LB BMCQ LMR GLTMJTC PMR?RGML. 1J?L?P HMGLRQ ?PC DMSLB GL RFC A?PN?J @MLCQ GL RFC F?LB ?LB RFC R?PQ?J @MLCQ MD RFC DMMR, ?Q UCJJ ?Q @CRUCCL TCPRC@P?C (Figure 38.27).
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1115
!@>LI< 38.27 4EB GLFKQP LC QEB @>OM>I ?LKBP FK QEB TOFPQ >OB BU>JMIBP LC MI>K>O GLFKQP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V "OF>K #. 'LPP)
H/3-+ J4/398 *L hinge joints, RFC QJGEFRJW PMSLBCB CLB MD MLC @MLC DGRQ GLRM RFC QJGEFRJW FMJJMU CLB MD RFC MRFCP @MLC. *L RFGQ U?W, MLC @MLC KMTCQ UFGJC RFC MRFCP PCK?GLQ QR?RGML?PW, JGIC RFC FGLEC MD ? BMMP. 5FC CJ@MU GQ ?L CV?KNJC MD ? FGLEC HMGLR. 5FC ILCC GQ QMKCRGKCQ AJ?QQGDGCB ?Q ? KMBGDGCB FGLEC HMGLR ( Figure 38.28).
!@>LI< 38.28 4EB BI?LT GLFKQ, TEBOB QEB O>AFRP >OQF@RI>QBP TFQE QEB ERJBORP, FP >K BU>JMIB LC > EFKDB GLFKQ. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V "OF>K #. 'LPP)
P/;49 J4/398 Pivot joints AMLQGQR MD RFC PMSLBCB CLB MD MLC @MLC DGRRGLE GLRM ? PGLE DMPKCB @W RFC MRFCP @MLC. 5FGQ QRPSARSPC ?JJMUQ PMR?RGML?J KMTCKCLR, ?Q RFC PMSLBCB @MLC KMTCQ ?PMSLB GRQ MUL ?VGQ. "L CV?KNJC MD ? NGTMR HMGLR GQ RFC HMGLR MD RFC DGPQR ?LB QCAMLB TCPRC@P?C MD RFC LCAI RF?R ?JJMUQ RFC FC?B RM KMTC @?AI ?LB DMPRF (Figure 38.29). 5FC HMGLR MD RFC UPGQR RF?R ?JJMUQ RFC N?JK MD RFC F?LB RM @C RSPLCB SN ?LB BMUL GQ ?JQM ? NGTMR HMGLR.
!@>LI< 38.29 4EB GLFKQ FK QEB KB@H QE>Q >IILTP QEB EB>A QL JLSB ?>@H >KA CLOQE FP >K BU>JMIB LC > MFSLQ GLFKQ.
1116
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
C43*>14/* J4/398 Condyloid joints AMLQGQR MD ?L MT?J-QF?NCB CLB MD MLC @MLC DGRRGLE GLRM ? QGKGJ?PJW MT?J-QF?NCB FMJJMU MD ?LMRFCP @MLC (Figure 38.30). 5FGQ GQ ?JQM QMKCRGKCQ A?JJCB ?L CJJGNQMGB?J HMGLR. 5FGQ RWNC MD HMGLR ?JJMUQ ?LESJ?P KMTCKCLR ?JMLE RUM ?VCQ, ?Q QCCL GL RFC HMGLRQ MD RFC UPGQR ?LB DGLECPQ, UFGAF A?L KMTC @MRF QGBC RM QGBC ?LB SN ?LB BMUL.
!@>LI< 38.30 4EB JBQ>@>OMLME>I>KDB>I GLFKQP FK QEB CFKDBO >OB BU>JMIBP LC @LKAVILFA GLFKQP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V 'O>V'P !K>QLJV)
!'**1+ J4/398 Saddle joints ?PC QM L?KCB @CA?SQC RFC CLBQ MD C?AF @MLC PCQCK@JC ? Q?BBJC, UGRF AMLA?TC ?LB AMLTCV NMPRGMLQ RF?R DGR RMECRFCP. 4?BBJC HMGLRQ ?JJMU ?LESJ?P KMTCKCLRQ QGKGJ?P RM AMLBWJMGB HMGLRQ @SR UGRF ? EPC?RCP P?LEC MD KMRGML. "L CV?KNJC MD ? Q?BBJC HMGLR GQ RFC RFSK@ HMGLR, UFGAF A?L KMTC @?AI ?LB DMPRF ?LB SN ?LB BMUL, @SR KMPC DPCCJW RF?L RFC UPGQR MP DGLECPQ (Figure 38.31).
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1117
!@>LI< 38.31 4EB @>OMLJBQ>@>OM>I GLFKQP FK QEB QERJ? >OB BU>JMIBP LC P>AAIB GLFKQP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V "OF>K #. 'LPP)
B'11-'3*-!4)0+9 J4/398 Ball-and-socket joints NMQQCQQ ? PMSLBCB, @?JJ-JGIC CLB MD MLC @MLC DGRRGLE GLRM ? ASNJGIC QMAICR MD ?LMRFCP @MLC. 5FGQ MPE?LGX?RGML ?JJMUQ RFC EPC?RCQR P?LEC MD KMRGML, ?Q ?JJ KMTCKCLR RWNCQ ?PC NMQQG@JC GL ?JJ BGPCARGMLQ. &V?KNJCQ MD @?JJ-?LB-QMAICR HMGLRQ ?PC RFC QFMSJBCP ?LB FGN HMGLRQ ( Figure 38.32).
!@>LI< 38.32 4EB PELRIABO GLFKQ FP >K BU>JMIB LC > ?>II->KA-PL@HBQ GLFKQ.
1118
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
8?RAF RFGQ animation (http://openstaxcollege.org/l/synovial_joints) QFMUGLE RFC QGV RWNCQ MD QWLMTG?J HMGLRQ.
-?<LD8KFCF>@JK
2EBRJ>QLILDFPQP >OB JBAF@>I AL@QLOP TEL PMB@F>IFWB FK QEB AF>DKLPFP >KA QOB>QJBKQ LC AFPLOABOP LC QEB GLFKQP, JRP@IBP, >KA ?LKBP. 4EBV AF>DKLPB >KA QOB>Q AFPB>PBP PR@E >P >OQEOFQFP, JRP@RILPHBIBQ>I AFPLOABOP, LPQBLMLOLPFP, >KA >RQLFJJRKB AFPB>PBP PR@E >P >KHVILPFKD PMLKAVIFQFP >KA OEBRJ>QLFA >OQEOFQFP. 2EBRJ>QLFA >OQEOFQFP (2!) FP >K FKCI>JJ>QLOV AFPLOABO QE>Q MOFJ>OFIV >CCB@QP QEB PVKLSF>I GLFKQP LC QEB E>KAP, CBBQ, >KA @BOSF@>I PMFKB. !CCB@QBA GLFKQP ?B@LJB PTLIIBK, PQFCC, >KA M>FKCRI. !IQELRDE FQ FP HKLTK QE>Q 2! FP >K >RQLFJJRKB AFPB>PB FK TEF@E QEB ?LAVwP FJJRKB PVPQBJ JFPQ>HBKIV >QQ>@HP EB>IQEV QFPPRB, QEB @>RPB LC 2! OBJ>FKP RKHKLTK. )JJRKB @BIIP COLJ QEB ?ILLA BKQBO GLFKQP >KA QEB PVKLSFRJ @>RPFKD @>OQFI>DB ?OB>HALTK, PTBIIFKD, >KA FKCI>JJ>QFLK LC QEB GLFKQ IFKFKD. "OB>HALTK LC @>OQFI>DB @>RPBP ?LKBP QL OR? >D>FKPQ B>@E LQEBO @>RPFKD M>FK. 2! FP JLOB @LJJLK FK TLJBK QE>K JBK >KA QEB >DB LC LKPBQ FP RPR>IIV 40t50 VB>OP LC >DB. 2EBRJ>QLILDFPQP @>K AF>DKLPB 2! LK QEB ?>PFP LC PVJMQLJP PR@E >P GLFKQ FKCI>JJ>QFLK >KA M>FK, 8-O>V >KA -2) FJ>DFKD, >KA ?ILLA QBPQP. !OQEOLDO>MEV FP > QVMB LC JBAF@>I FJ>DFKD LC GLFKQP QE>Q RPBP > @LKQO>PQ >DBKQ, PR@E >P > AVB, QE>Q FP LM>NRB QL 8-O>VP. 4EFP >IILTP QEB PLCQ QFPPRB PQOR@QROBP LC GLFKQPuPR@E >P @>OQFI>DB, QBKALKP, >KA IFD>JBKQPu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
38.4 R (LJ:C< CFEKI8:K@FE 8E; 'F:FDFK@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z $J?QQGDW RFC BGDDCPCLR RWNCQ MD KSQAJC RGQQSC z &VNJ?GL RFC PMJC MD KSQAJCQ GL JMAMKMRGML .SQAJC ACJJQ ?PC QNCAG?JGXCB DMP AMLRP?ARGML. .SQAJCQ ?JJMU DMP KMRGMLQ QSAF ?Q U?JIGLE, ?LB RFCW ?JQM D?AGJGR?RC @MBGJW NPMACQQCQ QSAF ?Q PCQNGP?RGML ?LB BGECQRGML. 5FC @MBW AMLR?GLQ RFPCC RWNCQ MD KSQAJC RGQQSC: QICJCR?J KSQAJC, A?PBG?A KSQAJC, ?LB QKMMRF KSQAJC (Figure 38.33).
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1119
!@>LI< 38.33 4EB ?LAV @LKQ>FKP QEOBB QVMBP LC JRP@IB QFPPRB: PHBIBQ>I JRP@IB, PJLLQE JRP@IB, >KA @>OAF>@ JRP@IB, SFPR>IFWBA EBOB RPFKD IFDEQ JF@OLP@LMV. 3JLLQE JRP@IB @BIIP >OB PELOQ, Q>MBOBA >Q B>@E BKA, >KA E>SB LKIV LKB MIRJM KR@IBRP FK B>@E. #>OAF>@ JRP@IB @BIIP >OB ?O>K@EBA >KA PQOF>QBA, ?RQ PELOQ. 4EB @VQLMI>PJ J>V ?O>K@E, >KA QEBV E>SB LKB KR@IBRP FK QEB @BKQBO LC QEB @BII. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .#), .)(; P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
Skeletal muscle tissue DMPKQ QICJCR?J KSQAJCQ, UFGAF ?RR?AF RM @MLCQ MP QIGL ?LB AMLRPMJ JMAMKMRGML ?LB ?LW KMTCKCLR RF?R A?L @C AMLQAGMSQJW AMLRPMJJCB. #CA?SQC GR A?L @C AMLRPMJJCB @W RFMSEFR, QICJCR?J KSQAJC GQ ?JQM A?JJCB TMJSLR?PW KSQAJC. 4ICJCR?J KSQAJCQ ?PC JMLE ?LB AWJGLBPGA?J GL ?NNC?P?LAC; UFCL TGCUCB SLBCP ? KGAPMQAMNC, QICJCR?J KSQAJC RGQQSC F?Q ? QRPGNCB MP QRPG?RCB ?NNC?P?LAC. 5FC QRPG?RGMLQ ?PC A?SQCB @W RFC PCESJ?P ?PP?LECKCLR MD AMLRP?ARGJC NPMRCGLQ (?ARGL ?LB KWMQGL). Actin GQ ? EJM@SJ?P AMLRP?ARGJC NPMRCGL RF?R GLRCP?ARQ UGRF myosin DMP KSQAJC AMLRP?ARGML. 4ICJCR?J KSQAJC ?JQM F?Q KSJRGNJC LSAJCG NPCQCLR GL ? QGLEJC ACJJ. Smooth muscle tissue MAASPQ GL RFC U?JJQ MD FMJJMU MPE?LQ QSAF ?Q RFC GLRCQRGLCQ, QRMK?AF, ?LB SPGL?PW @J?BBCP, ?LB ?PMSLB N?QQ?ECQ QSAF ?Q RFC PCQNGP?RMPW RP?AR ?LB @JMMB TCQQCJQ. 4KMMRF KSQAJC F?Q LM QRPG?RGMLQ, GQ LMR SLBCP TMJSLR?PW AMLRPMJ, F?Q MLJW MLC LSAJCSQ NCP ACJJ, GQ R?NCPCB ?R @MRF CLBQ, ?LB GQ A?JJCB GLTMJSLR?PW KSQAJC. Cardiac muscle tissue GQ MLJW DMSLB GL RFC FC?PR, ?LB A?PBG?A AMLRP?ARGMLQ NSKN @JMMB RFPMSEFMSR RFC @MBW ?LB K?GLR?GL @JMMB NPCQQSPC. -GIC QICJCR?J KSQAJC, A?PBG?A KSQAJC GQ QRPG?RCB, @SR SLJGIC QICJCR?J KSQAJC, A?PBG?A KSQAJC A?LLMR @C AMLQAGMSQJW AMLRPMJJCB ?LB GQ A?JJCB GLTMJSLR?PW KSQAJC. *R F?Q MLC LSAJCSQ NCP ACJJ, GQ @P?LAFCB, ?LB GQ BGQRGLESGQFCB @W RFC NPCQCLAC MD GLRCPA?J?RCB BGQIQ.
.B<C<K8C (LJ:C< !@9<I .KIL:KLI<

&?AF QICJCR?J KSQAJC DG@CP GQ ? QICJCR?J KSQAJC ACJJ. 5FCQC ACJJQ ?PC GLAPCBG@JW J?PEC, UGRF BG?KCRCPQ MD SN RM 100 ]K ?LB JCLERFQ MD SN RM 30 AK. 5FC NJ?QK? KCK@P?LC MD ? QICJCR?J KSQAJC DG@CP GQ A?JJCB RFC sarcolemma. 5FC Q?PAMJCKK? GQ RFC QGRC MD ?ARGML NMRCLRG?J AMLBSARGML, UFGAF RPGEECPQ KSQAJC AMLRP?ARGML. 8GRFGL C?AF KSQAJC DG@CP ?PC myofibrilsqJMLE AWJGLBPGA?J QRPSARSPCQ RF?R JGC N?P?JJCJ RM RFC KSQAJC DG@CP. .WMDG@PGJQ PSL RFC CLRGPC JCLERF MD RFC KSQAJC DG@CP, ?LB @CA?SQC RFCW ?PC MLJW ?NNPMVGK?RCJW 1.2 ]K GL BG?KCRCP, FSLBPCBQ RM RFMSQ?LBQ A?L @C DMSLB GLQGBC MLC KSQAJC DG@CP. 5FCW ?RR?AF RM RFC Q?PAMJCKK? ?R RFCGP CLBQ, QM RF?R ?Q KWMDG@PGJQ QFMPRCL, RFC CLRGPC KSQAJC ACJJ AMLRP?ARQ (Figure 38.34).
!@>LI< 38.34 ! PHBIBQ>I JRP@IB @BII FP PROOLRKABA ?V > MI>PJ> JBJ?O>KB @>IIBA QEB P>O@LIBJJ> TFQE > @VQLMI>PJ @>IIBA QEB P>O@LMI>PJ. ! JRP@IB CF?BO FP @LJMLPBA LC J>KV CF?OFIP, M>@H>DBA FKQL LOABOIV RKFQP.
5FC QRPG?RCB ?NNC?P?LAC MD QICJCR?J KSQAJC RGQQSC GQ ? PCQSJR MD PCNC?RGLE @?LBQ MD RFC NPMRCGLQ ?ARGL ?LB KWMQGL RF?R ?PC NPCQCLR ?JMLE RFC JCLERF MD KWMDG@PGJQ. %?PI " @?LBQ ?LB JGEFR * @?LBQ PCNC?R ?JMLE KWMDG@PGJQ, ?LB RFC ?JGELKCLR MD KWMDG@PGJQ GL RFC ACJJ A?SQCQ RFC CLRGPC ACJJ RM ?NNC?P QRPG?RCB MP @?LBCB. &?AF * @?LB F?Q ? BCLQC JGLC PSLLGLE TCPRGA?JJW RFPMSEF RFC KGBBJC A?JJCB ? ; BGQA MP ; JGLC. 5FC ; BGQAQ K?PI RFC @MPBCP MD SLGRQ A?JJCB sarcomeres, UFGAF ?PC RFC DSLARGML?J SLGRQ MD QICJCR?J KSQAJC. 0LC
1120
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Q?PAMKCPC GQ RFC QN?AC @CRUCCL RUM AMLQCASRGTC ; BGQAQ ?LB AMLR?GLQ MLC CLRGPC " @?LB ?LB RUM F?JTCQ MD ?L * @?LB, MLC ML CGRFCP QGBC MD RFC " @?LB. " KWMDG@PGJ GQ AMKNMQCB MD K?LW Q?PAMKCPCQ PSLLGLE ?JMLE GRQ JCLERF, ?LB ?Q RFC Q?PAMKCPCQ GLBGTGBS?JJW AMLRP?AR, RFC KWMDG@PGJQ ?LB KSQAJC ACJJQ QFMPRCL (Figure 38.35).
!@>LI< 38.35 ! P>O@LJBOB FP QEB OBDFLK COLJ LKB : IFKB QL QEB KBUQ : IFKB. ->KV P>O@LJBOBP >OB MOBPBKQ FK > JVLCF?OFI, OBPRIQFKD FK QEB PQOF>QFLK M>QQBOK @E>O>@QBOFPQF@ LC PHBIBQ>I JRP@IB.
.WMDG@PGJQ ?PC AMKNMQCB MD QK?JJCP QRPSARSPCQ A?JJCB myofilaments. 5FCPC ?PC RUM K?GL RWNCQ MD DGJ?KCLRQ: RFGAI DGJ?KCLRQ ?LB RFGL DGJ?KCLRQ; C?AF F?Q BGDDCPCLR AMKNMQGRGMLQ ?LB JMA?RGMLQ. Thick filaments MAASP MLJW GL RFC " @?LB MD ? KWMDG@PGJ. Thin filaments ?RR?AF RM ? NPMRCGL GL RFC ; BGQA A?JJCB ?JNF?-?ARGLGL ?LB MAASP ?APMQQ RFC CLRGPC JCLERF MD RFC * @?LB ?LB N?PRU?W GLRM RFC " @?LB. 5FC PCEGML ?R UFGAF RFGAI ?LB RFGL DGJ?KCLRQ MTCPJ?N F?Q ? BCLQC ?NNC?P?LAC, ?Q RFCPC GQ JGRRJC QN?AC @CRUCCL RFC DGJ?KCLRQ. 5FGL DGJ?KCLRQ BM LMR CVRCLB ?JJ RFC U?W GLRM RFC " @?LBQ, JC?TGLE ? ACLRP?J PCEGML MD RFC " @?LB RF?R MLJW AMLR?GLQ RFGAI DGJ?KCLRQ. 5FGQ ACLRP?J PCEGML MD RFC " @?LB JMMIQ QJGEFRJW JGEFRCP RF?L RFC PCQR MD RFC " @?LB ?LB GQ A?JJCB RFC ) XMLC. 5FC KGBBJC MD RFC ) XMLC F?Q ? TCPRGA?J JGLC A?JJCB RFC . JGLC, ?R UFGAF ?AACQQMPW NPMRCGLQ FMJB RMECRFCP RFGAI DGJ?KCLRQ. #MRF RFC ; BGQA ?LB RFC . JGLC FMJB KWMDGJ?KCLRQ GL NJ?AC RM K?GLR?GL RFC QRPSARSP?J ?PP?LECKCLR ?LB J?WCPGLE MD RFC KWMDG@PGJ. .WMDG@PGJQ ?PC AMLLCARCB RM C?AF MRFCP @W GLRCPKCBG?RC, MP BCQKGL, DGJ?KCLRQ RF?R ?RR?AF RM RFC ; BGQA. 5FGAI ?LB RFGL DGJ?KCLRQ ?PC RFCKQCJTCQ AMKNMQCB MD NPMRCGLQ. 5FGAI DGJ?KCLRQ ?PC AMKNMQCB MD RFC NPMRCGL KWMQGL. 5FC R?GJ MD ? KWMQGL KMJCASJC AMLLCARQ UGRF MRFCP KWMQGL KMJCASJCQ RM DMPK RFC ACLRP?J PCEGML MD ? RFGAI DGJ?KCLR LC?P RFC . JGLC, UFCPC?Q RFC FC?BQ ?JGEL ML CGRFCP QGBC MD RFC RFGAI DGJ?KCLR UFCPC RFC RFGL DGJ?KCLRQ MTCPJ?N. 5FC NPGK?PW AMKNMLCLR MD RFGL DGJ?KCLRQ GQ RFC ?ARGL NPMRCGL. 5UM MRFCP AMKNMLCLRQ MD RFC RFGL DGJ?KCLR ?PC RPMNMKWMQGL ?LB RPMNMLGL. "ARGL F?Q @GLBGLE QGRCQ DMP KWMQGL ?RR?AFKCLR. 4RP?LBQ MD RPMNMKWMQGL @JMAI RFC @GLBGLE QGRCQ ?LB NPCTCLR ?ARGLpKWMQGL GLRCP?ARGMLQ UFCL RFC KSQAJCQ ?PC ?R PCQR. 5PMNMLGL AMLQGQRQ MD RFPCC EJM@SJ?P QS@SLGRQ. 0LC QS@SLGR @GLBQ RM RPMNMKWMQGL, MLC QS@SLGR @GLBQ RM ?ARGL, ?LB MLC QS@SLGR @GLBQ $?2+ GMLQ.
7GCU RFGQ animation (http://openstaxcollege.org/l/skeletal_muscle) QFMUGLE RFC MPE?LGX?RGML MD KSQAJC DG@CPQ.
.C@;@E> !@C8D<EK (F;<C F= CFEKI8:K@FE

'MP ? KSQAJC ACJJ RM AMLRP?AR, RFC Q?PAMKCPC KSQR QFMPRCL. )MUCTCP, RFGAI ?LB RFGL DGJ?KCLRQqRFC AMKNMLCLRQ MD Q?PAMKCPCQqBM LMR QFMPRCL. *LQRC?B, RFCW QJGBC @W MLC ?LMRFCP, A?SQGLE RFC Q?PAMKCPC RM QFMPRCL UFGJC RFC DGJ?KCLRQ PCK?GL RFC Q?KC JCLERF. 5FC QJGBGLE DGJ?KCLR RFCMPW MD KSQAJC AMLRP?ARGML U?Q BCTCJMNCB RM DGR RFC BGDDCPCLACQ M@QCPTCB GL RFC L?KCB @?LBQ ML RFC Q?PAMKCPC ?R BGDDCPCLR BCEPCCQ MD KSQAJC AMLRP?ARGML ?LB PCJ?V?RGML. 5FC KCAF?LGQK MD AMLRP?ARGML GQ RFC @GLBGLE MD KWMQGL RM ?ARGL, DMPKGLE APMQQ-@PGBECQ RF?R ECLCP?RC DGJ?KCLR KMTCKCLR (Figure 38.36).
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
1121
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8FCL ? Q?PAMKCPC QFMPRCLQ, QMKC PCEGMLQ QFMPRCL UFCPC?Q MRFCPQ QR?W RFC Q?KC JCLERF. " Q?PAMKCPC GQ BCDGLCB ?Q RFC BGQR?LAC @CRUCCL RUM AMLQCASRGTC ; BGQAQ MP ; JGLCQ; UFCL ? KSQAJC AMLRP?ARQ, RFC BGQR?LAC @CRUCCL RFC ; BGQAQ GQ PCBSACB. 5FC ) XMLCqRFC ACLRP?J PCEGML MD RFC " XMLCqAMLR?GLQ MLJW RFGAI DGJ?KCLRQ ?LB GQ QFMPRCLCB BSPGLE AMLRP?ARGML. 5FC * @?LB AMLR?GLQ MLJW RFGL DGJ?KCLRQ ?LB ?JQM QFMPRCLQ. 5FC " @?LB BMCQ LMR QFMPRCLqGR PCK?GLQ RFC Q?KC JCLERFq@SR " @?LBQ MD BGDDCPCLR Q?PAMKCPCQ KMTC AJMQCP RMECRFCP BSPGLE AMLRP?ARGML, CTCLRS?JJW BGQ?NNC?PGLE. 5FGL DGJ?KCLRQ ?PC NSJJCB @W RFC RFGAI DGJ?KCLRQ RMU?PB RFC ACLRCP MD RFC Q?PAMKCPC SLRGJ RFC ; BGQAQ ?NNPM?AF RFC RFGAI DGJ?KCLRQ. 5FC XMLC MD MTCPJ?N, GL UFGAF RFGL DGJ?KCLRQ ?LB RFGAI DGJ?KCLRQ MAASNW RFC Q?KC ?PC?, GLAPC?QCQ ?Q RFC RFGL DGJ?KCLRQ KMTC GLU?PB.
A/+ 8E; (LJ:C< CFEKI8:K@FE

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tAMAICBu NMQGRGML. 5FC KWMQGL FC?B GQ RFCL GL ? NMQGRGML DMP DSPRFCP KMTCKCLR, NMQQCQQGLE NMRCLRG?J CLCPEW, @SR "%1 ?LB 1G ?PC QRGJJ ?RR?AFCB. *D ?ARGL @GLBGLE QGRCQ ?PC AMTCPCB ?LB SL?T?GJ?@JC, RFC KWMQGL UGJJ PCK?GL GL RFC FGEF CLCPEW AMLDGESP?RGML UGRF "51 FWBPMJWXCB, @SR QRGJJ ?RR?AFCB. *D RFC ?ARGL @GLBGLE QGRCQ ?PC SLAMTCPCB, ? APMQQ-@PGBEC UGJJ DMPK; RF?R GQ, RFC KWMQGL FC?B QN?LQ RFC BGQR?LAC @CRUCCL RFC ?ARGL ?LB KWMQGL KMJCASJCQ. 1G GQ RFCL PCJC?QCB, ?JJMUGLE KWMQGL RM CVNCLB RFC QRMPCB CLCPEW ?Q ? AMLDMPK?RGML?J AF?LEC. 5FC KWMQGL FC?B KMTCQ RMU?PB RFC . JGLC, NSJJGLE RFC ?ARGL ?JMLE UGRF GR. "Q RFC ?ARGL GQ NSJJCB, RFC DGJ?KCLRQ KMTC ?NNPMVGK?RCJW 10 LK RMU?PB RFC . JGLC. 5FGQ KMTCKCLR GQ A?JJCB RFC NMUCP QRPMIC, ?Q GR GQ RFC QRCN ?R UFGAF DMPAC GQ NPMBSACB. "Q RFC ?ARGL GQ NSJJCB RMU?PB RFC . JGLC, RFC Q?PAMKCPC QFMPRCLQ ?LB RFC KSQAJC AMLRP?ARQ.
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8FCL RFC KWMQGL FC?B GQ tAMAICB,u GR AMLR?GLQ CLCPEW ?LB GQ GL ? FGEF-CLCPEW AMLDGESP?RGML. 5FGQ CLCPEW GQ CVNCLBCB ?Q RFC KWMQGL FC?B KMTCQ RFPMSEF RFC NMUCP QRPMIC; ?R RFC CLB MD RFC NMUCP QRPMIC, RFC KWMQGL FC?B GQ GL ? JMU-CLCPEW NMQGRGML. "DRCP RFC NMUCP QRPMIC, "%1 GQ PCJC?QCB; FMUCTCP, RFC APMQQ@PGBEC DMPKCB GQ QRGJJ GL NJ?AC, ?LB ?ARGL ?LB KWMQGL ?PC @MSLB RMECRFCP. "51 A?L RFCL ?RR?AF RM KWMQGL, UFGAF ?JJMUQ RFC APMQQ-@PGBEC AWAJC RM QR?PR ?E?GL ?LB DSPRFCP KSQAJC AMLRP?ARGML A?L MAASP (Figure 38.37).
8?RAF RFGQ video (http://openstaxcollege.org/l/contract_muscle) CVNJ?GLGLE FMU ? KSQAJC AMLRP?ARGML GQ QGEL?JCB.
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!@>LI< 38.37 4EB @OLPP-?OFADB JRP@IB @LKQO>@QFLK @V@IB, TEF@E FP QOFDDBOBA ?V #>2+ ?FKAFKD QL QEB >@QFK >@QFSB PFQB, FP PELTK. 7FQE B>@E @LKQO>@QFLK @V@IB, >@QFK JLSBP OBI>QFSB QL JVLPFK.
7EF@E LC QEB CLIILTFKD PQ>QBJBKQP >?LRQ JRP@IB @LKQO>@QFLK FP QORB >. 4EB MLTBO PQOLHB L@@ROP TEBK !40 FP EVAOLIVWBA QL !$0 >KA MELPME>QB. ?. 4EB MLTBO PQOLHB L@@ROP TEBK !$0 >KA MELPME>QB AFPPL@F>QB COLJ QEB JVLPFK EB>A. @. 4EB MLTBO PQOLHB L@@ROP TEBK !$0 >KA MELPME>QB AFPPL@F>QB COLJ QEB >@QFK >@QFSB PFQB. A. 4EB MLTBO PQOLHB L@@ROP TEBK #>2+ ?FKAP QEB @>I@FRJ EB>A.
7GCU RFGQ animation (http://openstaxcollege.org/l/muscle_contract) MD RFC APMQQ-@PGBEC KSQAJC AMLRP?ARGML.
-<>LC8KFIP +IFK<@EJ
8FCL ? KSQAJC GQ GL ? PCQRGLE QR?RC, ?ARGL ?LB KWMQGL ?PC QCN?P?RCB. 5M ICCN ?ARGL DPMK @GLBGLE RM RFC ?ARGTC QGRC ML KWMQGL, PCESJ?RMPW NPMRCGLQ @JMAI RFC KMJCASJ?P @GLBGLE QGRCQ. Tropomyosin @JMAIQ
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KWMQGL @GLBGLE QGRCQ ML ?ARGL KMJCASJCQ, NPCTCLRGLE APMQQ-@PGBEC DMPK?RGML ?LB NPCTCLRGLE AMLRP?ARGML GL ? KSQAJC UGRFMSR LCPTMSQ GLNSR. Troponin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
O:@K8K@FEVCFEKI8:K@FE CFLGC@E>
&VAGR?RGMLpAMLRP?ARGML AMSNJGLE GQ RFC JGLI (RP?LQBSARGML) @CRUCCL RFC ?ARGML NMRCLRG?J ECLCP?RCB GL RFC Q?PAMJCKK? ?LB RFC QR?PR MD ? KSQAJC AMLRP?ARGML. 5FC RPGEECP DMP A?JAGSK PCJC?QC DPMK RFC Q?PAMNJ?QKGA PCRGASJSK GLRM RFC Q?PAMNJ?QK GQ ? LCSP?J QGEL?J. &?AF QICJCR?J KSQAJC DG@CP GQ AMLRPMJJCB @W ? KMRMP LCSPML, UFGAF AMLBSARQ QGEL?JQ DPMK RFC @P?GL MP QNGL?J AMPB RM RFC KSQAJC. 5FC ?PC? MD RFC Q?PAMJCKK? ML RFC KSQAJC DG@CP RF?R GLRCP?ARQ UGRF RFC LCSPML GQ A?JJCB RFC motor end plate. 5FC CLB MD RFC LCSPMLsQ ?VML GQ A?JJCB RFC QWL?NRGA RCPKGL?J, ?LB GR BMCQ LMR ?ARS?JJW AMLR?AR RFC KMRMP CLB NJ?RC. " QK?JJ QN?AC A?JJCB RFC QWL?NRGA AJCDR QCN?P?RCQ RFC QWL?NRGA RCPKGL?J DPMK RFC KMRMP CLB NJ?RC. &JCARPGA?J QGEL?JQ RP?TCJ ?JMLE RFC LCSPMLs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pNMR?QQGSK "51?QC SQCQ ACJJSJ?P CLCPEW RM KMTC ,+ GMLQ GLQGBC RFC ACJJ ?LB /?+ GMLQ MSRQGBC. 5FGQ ?JMLC ?AASKSJ?RCQ ? QK?JJ CJCARPGA?J AF?PEC, @SR ? @GE AMLACLRP?RGML EP?BGCLR. 5FCPC GQ JMRQ MD ,+ GL RFC ACJJ ?LB JMRQ MD /?+ MSRQGBC RFC ACJJ. 1MR?QQGSK GQ ?@JC RM JC?TC RFC ACJJ RFPMSEF ,+ AF?LLCJQ RF?R ?PC MNCL 90% MD RFC RGKC, ?LB GR BMCQ. )MUCTCP, /?+ AF?LLCJQ ?PC P?PCJW MNCL, QM /?+ PCK?GLQ MSRQGBC RFC ACJJ. 8FCL ,+ JC?TCQ RFC ACJJ, M@CWGLE GRQ AMLACLRP?RGML EP?BGCLR, RF?R CDDCARGTCJW JC?TCQ ? LCE?RGTC AF?PEC @CFGLB. 4M ?R PCQR, RFCPC GQ ? J?PEC AMLACLRP?RGML EP?BGCLR DMP /?+ RM CLRCP RFC ACJJ, ?LB RFCPC GQ ?L ?AASKSJ?RGML MD LCE?RGTC AF?PECQ JCDR @CFGLB GL RFC ACJJ. 5FGQ GQ RFC PCQRGLE KCK@P?LC NMRCLRG?J. 1MRCLRG?J GL RFGQ AMLRCVR KC?LQ ? QCN?P?RGML MD CJCARPGA?J AF?PEC RF?R GQ A?N?@JC MD BMGLE UMPI. *R GQ KC?QSPCB GL TMJRQ, HSQR JGIC ? @?RRCPW. )MUCTCP, RFC RP?LQKCK@P?LC NMRCLRG?J GQ AMLQGBCP?@JW QK?JJCP (0.07 7); RFCPCDMPC, RFC QK?JJ T?JSC GQ CVNPCQQCB ?Q KGJJGTMJRQ (K7) MP 70 K7. #CA?SQC RFC GLQGBC MD ? ACJJ GQ LCE?RGTC AMKN?PCB UGRF RFC MSRQGBC, ? KGLSQ QGEL QGELGDGCQ RFC CVACQQ MD LCE?RGTC AF?PECQ GLQGBC RFC ACJJ, y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qDMJBQ GL RFC Q?PAMJCKK? RF?R APC?RC ? J?PEC QSPD?AC ?PC? DMP RFC LCSPMRP?LQKGRRCP RM @GLB RM PCACNRMPQ. 5FC PCACNRMPQ ?PC ?ARS?JJW QMBGSK AF?LLCJQ RF?R MNCL RM ?JJMU RFC N?QQ?EC MD /?+ GLRM RFC ACJJ UFCL RFCW PCACGTC LCSPMRP?LQKGRRCP QGEL?J. "ACRWJAFMJGLC ("$F) GQ ? LCSPMRP?LQKGRRCP PCJC?QCB @W KMRMP LCSPMLQ RF?R @GLBQ RM PCACNRMPQ GL RFC KMRMP CLB NJ?RC. /CSPMRP?LQKGRRCP PCJC?QC MAASPQ UFCL ?L ?ARGML NMRCLRG?J RP?TCJQ BMUL RFC KMRMP LCSPMLs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"$F GQ @PMICL BMUL @W RFC CLXWKC acetylcholinesterase ("$F&) GLRM ?ACRWJ ?LB AFMJGLC. "$F& PCQGBCQ GL RFC QWL?NRGA AJCDR, @PC?IGLE BMUL "$F QM RF?R GR BMCQ LMR PCK?GL @MSLB RM "$F PCACNRMPQ, UFGAF UMSJB A?SQC SLU?LRCB CVRCLBCB KSQAJC AMLRP?ARGML (Figure 38.38).
!@>LI< 38.38 4EFP AF>DO>J PELTP BU@FQ>QFLK-@LKQO>@QFLK @LRMIFKD FK > PHBIBQ>I JRP@IB @LKQO>@QFLK. 4EB P>O@LMI>PJF@ OBQF@RIRJ FP > PMB@F>IFWBA BKALMI>PJF@ OBQF@RIRJ CLRKA FK JRP@IB @BIIP.
4EB AB>AIV KBOSB D>P 3>OFK FOOBSBOPF?IV FKEF?FQP >@BQV@ELIFKBPQBO>PB. 7E>Q BCCB@Q TLRIA 3>OFK E>SB LK JRP@IB @LKQO>@QFLK "DRCP BCNMJ?PGX?RGML, RFC KCK@P?LC PCRSPLQ RM GRQ PCQRGLE QR?RC. 5FGQ GQ A?JJCB PCNMJ?PGX?RGML, BSPGLE UFGAF TMJR?EC-E?RCB QMBGSK AF?LLCJQ AJMQC. 1MR?QQGSK AF?LLCJQ AMLRGLSC ?R 90% AMLBSAR?LAC. #CA?SQC RFC NJ?QK? KCK@P?LC QMBGSKpNMR?QQGSK "51?QC ?JU?WQ RP?LQNMPRQ GMLQ, RFC PCQRGLE QR?RC (LCE?RGTCJW AF?PECB GLQGBC PCJ?RGTC RM RFC MSRQGBC) GQ PCQRMPCB. 5FC NCPGMB GKKCBG?RCJW DMJJMUGLE RFC RP?LQKGQQGML MD ?L GKNSJQC GL ? LCPTC MP KSQAJC, GL UFGAF ? LCSPML MP KSQAJC ACJJ PCE?GLQ GRQ ?@GJGRW RM RP?LQKGR ?LMRFCP GKNSJQC, GQ A?JJCB RFC PCDP?ARMPW NCPGMB. %SPGLE RFC PCDP?ARMPW NCPGMB, RFC KCK@P?LC A?LLMR ECLCP?RC ?LMRFCP ?ARGML NMRCLRG?J. . 5FC PCDP?ARMPW NCPGMB ?JJMUQ RFC TMJR?EC-QCLQGRGTC GML AF?LLCJQ RM PCRSPL RM RFCGP PCQRGLE AMLDGESP?RGMLQ. 5FC QMBGSK NMR?QQGSK "51?QC AMLRGLS?JJW KMTCQ /?+ @?AI MSR MD RFC ACJJ ?LB ,+ @?AI GLRM RFC ACJJ, ?LB RFC ,+ JC?IQ MSR JC?TGLE LCE?RGTC AF?PEC @CFGLB. 7CPW OSGAIJW, RFC KCK@P?LC PCNMJ?PGXCQ, QM RF?R GR A?L ?E?GL @C BCNMJ?PGXCB.
CFEKIFC F= (LJ:C< /<EJ@FE

/CSP?J AMLRPMJ GLGRG?RCQ RFC DMPK?RGML MD ?ARGLpKWMQGL APMQQ-@PGBECQ, JC?BGLE RM RFC Q?PAMKCPC QFMPRCLGLE GLTMJTCB GL KSQAJC AMLRP?ARGML. 5FCQC AMLRP?ARGMLQ CVRCLB DPMK RFC KSQAJC DG@CP RFPMSEF AMLLCARGTC
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RGQQSC RM NSJJ ML @MLCQ, A?SQGLE QICJCR?J KMTCKCLR. 5FC NSJJ CVCPRCB @W ? KSQAJC GQ A?JJCB RCLQGML, ?LB RFC ?KMSLR MD DMPAC APC?RCB @W RFGQ RCLQGML A?L T?PW. 5FGQ CL?@JCQ RFC Q?KC KSQAJCQ RM KMTC TCPW JGEFR M@HCARQ ?LB TCPW FC?TW M@HCARQ. *L GLBGTGBS?J KSQAJC DG@CPQ, RFC ?KMSLR MD RCLQGML NPMBSACB BCNCLBQ ML RFC APMQQ-QCARGML?J ?PC? MD RFC KSQAJC DG@CP ?LB RFC DPCOSCLAW MD LCSP?J QRGKSJ?RGML. 5FC LSK@CP MD APMQQ-@PGBECQ DMPKCB @CRUCCL ?ARGL ?LB KWMQGL BCRCPKGLC RFC ?KMSLR MD RCLQGML RF?R ? KSQAJC DG@CP A?L NPMBSAC. $PMQQ-@PGBECQ A?L MLJW DMPK UFCPC RFGAI ?LB RFGL DGJ?KCLRQ MTCPJ?N, ?JJMUGLE KWMQGL RM @GLB RM ?ARGL. *D KMPC APMQQ-@PGBECQ ?PC DMPKCB, KMPC KWMQGL UGJJ NSJJ ML ?ARGL, ?LB KMPC RCLQGML UGJJ @C NPMBSACB. 5FC GBC?J JCLERF MD ? Q?PAMKCPC BSPGLE NPMBSARGML MD K?VGK?J RCLQGML MAASPQ UFCL RFGAI ?LB RFGL DGJ?KCLRQ MTCPJ?N RM RFC EPC?RCQR BCEPCC. *D ? Q?PAMKCPC ?R PCQR GQ QRPCRAFCB N?QR ?L GBC?J PCQRGLE JCLERF, RFGAI ?LB RFGL DGJ?KCLRQ BM LMR MTCPJ?N RM RFC EPC?RCQR BCEPCC, ?LB DCUCP APMQQ-@PGBECQ A?L DMPK. 5FGQ PCQSJRQ GL DCUCP KWMQGL FC?BQ NSJJGLE ML ?ARGL, ?LB JCQQ RCLQGML GQ NPMBSACB. "Q ? Q?PAMKCPC GQ QFMPRCLCB, RFC XMLC MD MTCPJ?N GQ PCBSACB ?Q RFC RFGL DGJ?KCLRQ PC?AF RFC ) XMLC, UFGAF GQ AMKNMQCB MD KWMQGL R?GJQ. #CA?SQC GR GQ KWMQGL FC?BQ RF?R DMPK APMQQ-@PGBECQ, ?ARGL UGJJ LMR @GLB RM KWMQGL GL RFGQ XMLC, PCBSAGLE RFC RCLQGML NPMBSACB @W RFGQ KWMDG@CP. *D RFC Q?PAMKCPC GQ QFMPRCLCB CTCL KMPC, RFGL DGJ?KCLRQ @CEGL RM MTCPJ?N UGRF C?AF MRFCPqPCBSAGLE APMQQ-@PGBEC DMPK?RGML CTCL DSPRFCP, ?LB NPMBSAGLE CTCL JCQQ RCLQGML. $MLTCPQCJW, GD RFC Q?PAMKCPC GQ QRPCRAFCB RM RFC NMGLR ?R UFGAF RFGAI ?LB RFGL DGJ?KCLRQ BM LMR MTCPJ?N ?R ?JJ, LM APMQQ-@PGBECQ ?PC DMPKCB ?LB LM RCLQGML GQ NPMBSACB. 5FGQ ?KMSLR MD QRPCRAFGLE BMCQ LMR SQS?JJW MAASP @CA?SQC ?AACQQMPW NPMRCGLQ, GLRCPL?J QCLQMPW LCPTCQ, ?LB AMLLCARGTC RGQQSC MNNMQC CVRPCKC QRPCRAFGLE. 5FC NPGK?PW T?PG?@JC BCRCPKGLGLE DMPAC NPMBSARGML GQ RFC LSK@CP MD KWMDG@CPQ UGRFGL RFC KSQAJC RF?R PCACGTC ?L ?ARGML NMRCLRG?J DPMK RFC LCSPML RF?R AMLRPMJQ RF?R DG@CP. 8FCL SQGLE RFC @GACNQ RM NGAI SN ? NCLAGJ, RFC KMRMP AMPRCV MD RFC @P?GL MLJW QGEL?JQ ? DCU LCSPMLQ MD RFC @GACNQ, ?LB MLJW ? DCU KWMDG@CPQ PCQNMLB. *L TCPRC@P?RCQ, C?AF KWMDG@CP PCQNMLBQ DSJJW GD QRGKSJ?RCB. 8FCL NGAIGLE SN ? NG?LM, RFC KMRMP AMPRCV QGEL?JQ ?JJ MD RFC LCSPMLQ GL RFC @GACNQ ?LB CTCPW KWMDG@CP N?PRGAGN?RCQ. 5FGQ GQ AJMQC RM RFC K?VGKSK DMPAC RFC KSQAJC A?L NPMBSAC. "Q KCLRGMLCB ?@MTC, GLAPC?QGLE RFC DPCOSCLAW MD ?ARGML NMRCLRG?JQ (RFC LSK@CP MD QGEL?JQ NCP QCAMLB) A?L GLAPC?QC RFC DMPAC ? @GR KMPC, @CA?SQC RFC RPMNMKWMQGL GQ DJMMBCB UGRF A?JAGSK.
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& 4 / -(.
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s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C8D<CC8 J?WCP MD AMKN?AR RGQQSC RF?R QSPPMSLBQ ? ACLRP?J A?L?J A?JJCB RFC )?TCPQG?L A?L?J C8K<I8C IFK8K@FE PMR?RGML ?U?W DPMK RFC KGBJGLC MD RFC @MBW CFE> 9FE< @MLC RF?R GQ JMLECP RF?L UGBC, ?LB F?Q ? QF?DR ?LB RUM CLBQ CFN<I C@D9 AMLQGQRQ MD RFC RFGEF, RFC JCE, ?LB RFC DMMR D<;@8C IFK8K@FE PMR?RGML RMU?PB RFC KGBJGLC MD RFC @MBW D<K8:8IGLJ DGTC @MLCQ RF?R AMKNPGQC RFC N?JK D<K8K8IJ8C MLC MD RFC DGTC @MLCQ MD RFC DMMR DFKFI <E; GC8K< Q?PAMJCKK? MD RFC KSQAJC DG@CP RF?R GLRCP?ARQ UGRF RFC LCSPML DPF=@9I@C JMLE AWJGLBPGA?J QRPSARSPCQ RF?R JGC N?P?JJCJ RM RFC KSQAJC DG@CP DPF=@C8D<EK QK?JJ QRPSARSPCQ RF?R K?IC SN KWMDG@PGJQ DPFJ@E AMLRP?ARGJC NPMRCGL RF?R GLRCP?ARQ UGRF ?ARGL DMP KSQAJC AMLRP?ARGML FGGFJ@K@FE KMTCKCLR MD RFC RFSK@ RMU?PB RFC DGLECPQ MD RFC Q?KC F?LB, K?IGLE GR NMQQG@JC RM EP?QN ?LB FMJB M@HCARQ FJJ<FLJ K@JJL< AMLLCARGTC RGQQSC RF?R AMLQRGRSRCQ RFC CLBMQICJCRML FJJ@=@:8K@FE (?JQM, MQRCMECLCQGQ) NPMACQQ MD @MLC DMPK?RGML @W MQRCM@J?QRQ FJK<F9C8JK @MLC ACJJ PCQNMLQG@JC DMP @MLC DMPK?RGML FJK<F:C8JK J?PEC @MLC ACJJQ UGRF SN RM 50 LSAJCG, PCQNMLQG@JC DMP @MLC PCKMBCJGLE FJK<F:PK< K?RSPC @MLC ACJJQ ?LB RFC K?GL ACJJ GL @MLC RGQQSC FJK<FE AWJGLBPGA?J QRPSARSPC ?JGELCB N?P?JJCJ RM RFC JMLE ?VGQ MD RFC @MLC G8K<CC8 (?JQM, ILCCA?N) RPG?LESJ?P @MLC RF?R JGCQ ?LRCPGMP RM RFC ILCC HMGLR G<:KFI8C >@I;C< @MLCQ RF?R RP?LQKGR RFC DMPAC ECLCP?RCB @W RFC SNNCP JGK@Q RM RFC ?VG?J QICJCRML G<CM@: >@I;C< @MLCQ RF?R RP?LQKGR RFC DMPAC ECLCP?RCB @W RFC JMUCP JGK@Q RM RFC ?VG?J QICJCRML G?8C8E>< MLC MD RFC @MLCQ MD RFC DGLECPQ MP RMCQ G@MFK AF@EK HMGLR UGRF RFC PMSLBCB CLB MD MLC @MLC DGRRGLE GLRM ? PGLE DMPKCB @W RFC MRFCP @MLC GC8E8I AF@EK HMGLR UGRF @MLCQ UFMQC ?PRGASJ?RGLE QSPD?ACQ ?PC DJ?R GC8EK8I =C<O@FE @CLBGLE ?R RFC ?LIJC QSAF RF?R RFC FCCJ GQ JGDRCB, QSAF ?Q UFCL QR?LBGLE ML RFC RMCQ GIFE8K@FE KMTCKCLR GL UFGAF RFC N?JK D?ACQ @?AIU?PB GIFKI8:K@FE ?LRCPGMP KMTCKCLR MD ? @MLC GL RFC FMPGXMLR?J NJ?LC I8;@LJ @MLC JMA?RCB ?JMLE RFC J?RCP?J (RFSK@) QGBC MD RFC DMPC?PK; ?PRGASJ?RCQ UGRF RFC FSKCPSQ ?R RFC CJ@MU I<JFIGK@FE NPMACQQ @W UFGAF MQRCMAJ?QRQ PCJC?QC KGLCP?JQ QRMPCB GL @MLCQ I<KI8:K@FE KMTCKCLR GL UFGAF ? HMGLR KMTCQ @?AI GLRM NMQGRGML ?DRCP NPMRP?ARGML I@9 MLC MD 12 N?GPQ MD JMLE, ASPTCB @MLCQ RF?R ?RR?AF RM RFC RFMP?AGA TCPRC@P?C ?LB ASPTC RMU?PB RFC DPMLR MD RFC @MBW RM DMPK RFC PG@A?EC
IFK8K@FE8C DFM<D<EK KMTCKCLR MD ? @MLC ?Q GR PMR?RCQ ?PMSLB GRQ MUL JMLEGRSBGL?J ?VGQ
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M<IK<9I8C :FCLDE (?JQM, QNGLC) QSPPMSLBQ ?LB NPMRCARQ RFC QNGL?J AMPB, QSNNMPRQ RFC FC?B, ?LB ?ARQ ?Q ?L ?RR?AFKCLR NMGLR DMP PG@Q ?LB KSQAJCQ MD RFC @?AI ?LB LCAI
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38.1 /PG<J F= .B<C<K8C .PJK<DJ
5FC RFPCC RWNCQ MD QICJCRML BCQGELQ ?PC FWBPMQR?RGA QICJCRMLQ, CVMQICJCRMLQ, ?LB CLBMQICJCRMLQ. " FWBPMQR?RGA QICJCRML GQ DMPKCB @W ? DJSGB-DGJJCB AMKN?PRKCLR FCJB SLBCP FWBPMQR?RGA NPCQQSPC; KMTCKCLR GQ APC?RCB @W RFC KSQAJCQ NPMBSAGLE NPCQQSPC ML RFC DJSGB. "L CVMQICJCRML GQ ? F?PB CVRCPL?J QICJCRML RF?R NPMRCARQ RFC MSRCP QSPD?AC MD ?L MPE?LGQK ?LB CL?@JCQ KMTCKCLR RFPMSEF KSQAJCQ ?RR?AFCB ML RFC GLQGBC. "L CLBMQICJCRML GQ ?L GLRCPL?J QICJCRML AMKNMQCB MD F?PB, KGLCP?JGXCB RGQQSC RF?R ?JQM CL?@JCQ KMTCKCLR @W ?RR?AFKCLR RM KSQAJCQ. 5FC FSK?L QICJCRML GQ ?L CLBMQICJCRML RF?R GQ AMKNMQCB MD RFC ?VG?J ?LB ?NNCLBGASJ?P QICJCRML. 5FC ?VG?J QICJCRML GQ AMKNMQCB MD RFC @MLCQ MD RFC QISJJ, MQQGAJCQ MD RFC C?P, FWMGB @MLC, TCPRC@P?J AMJSKL, ?LB PG@A?EC. 5FC QISJJ AMLQGQRQ MD CGEFR AP?LG?J @MLCQ ?LB 14 D?AG?J @MLCQ. 4GV @MLCQ K?IC SN RFC MQQGAJCQ MD RFC KGBBJC C?P, UFGJC RFC FWMGB @MLC GQ JMA?RCB GL RFC LCAI SLBCP RFC K?LBG@JC. 5FC TCPRC@P?J AMJSKL AMLR?GLQ 26 @MLCQ, ?LB GR QSPPMSLBQ ?LB NPMRCARQ RFC QNGL?J AMPB. 5FC RFMP?AGA A?EC AMLQGQRQ MD RFC QRCPLSK, PG@Q, RFMP?AGA TCPRC@P?C, ?LB AMQR?J A?PRGJ?ECQ. 5FC ?NNCLBGASJ?P QICJCRML GQ K?BC SN MD RFC JGK@Q MD RFC SNNCP ?LB JMUCP JGK@Q. 5FC NCARMP?J EGPBJC GQ AMKNMQCB MD RFC AJ?TGAJCQ ?LB RFC QA?NSJ?C. 5FC SNNCP JGK@ AMLR?GLQ 30 @MLCQ GL RFC ?PK, RFC DMPC?PK, ?LB RFC F?LB. 5FC NCJTGA EGPBJC ?RR?AFCQ RFC JMUCP JGK@Q RM RFC ?VG?J QICJCRML. 5FC JMUCP JGK@ GLAJSBCQ RFC @MLCQ MD RFC RFGEF, RFC JCE, ?LB RFC DMMR.
38.2 BFE<
#MLC, MP MQQCMSQ RGQQSC, GQ AMLLCARGTC RGQQSC RF?R GLAJSBCQ QNCAG?JGXCB ACJJQ, KGLCP?J Q?JRQ, ?LB AMJJ?ECL DG@CPQ. 5FC FSK?L QICJCRML A?L @C BGTGBCB GLRM JMLE @MLCQ, QFMPR @MLCQ, DJ?R @MLCQ, ?LB GPPCESJ?P @MLCQ. $MKN?AR @MLC RGQQSC GQ AMKNMQCB MD MQRCMLQ ?LB DMPKQ RFC CVRCPL?J J?WCP MD ?JJ @MLCQ. 4NMLEW @MLC RGQQSC GQ AMKNMQCB MD RP?@CASJ?C ?LB DMPKQ RFC GLLCP N?PR MD ?JJ @MLCQ. 'MSP RWNCQ MD ACJJQ AMKNMQC @MLW RGQQSC: MQRCMAWRCQ, MQRCMAJ?QRQ, MQRCMNPMECLGRMP ACJJQ, ?LB MQRCM@J?QRQ. 0QQGDGA?RGML GQ RFC NPMACQQ MD @MLC DMPK?RGML @W MQRCM@J?QRQ. *LRP?KCK@P?LMSQ MQQGDGA?RGML GQ RFC NPMACQQ MD @MLC BCTCJMNKCLR DPMK DG@PMSQ KCK@P?LCQ. &LBMAFMLBP?J MQQGDGA?RGML GQ RFC NPMACQQ MD @MLC BCTCJMNKCLR DPMK FW?JGLC A?PRGJ?EC. -MLE @MLCQ JCLERFCL ?Q AFMLBPMAWRCQ BGTGBC ?LB QCAPCRC FW?JGLC A?PRGJ?EC. 0QRCM@J?QRQ PCNJ?AC A?PRGJ?EC UGRF @MLC. "NNMQGRGML?J EPMURF GQ RFC GLAPC?QC GL RFC BG?KCRCP MD @MLCQ @W RFC ?BBGRGML MD @MLC RGQQSC ?R RFC QSPD?AC MD @MLCQ. #MLC PCKMBCJGLE GLTMJTCQ RFC NPMACQQCQ MD @MLC BCNMQGRGML @W MQRCM@J?QRQ ?LB @MLC PCQMPNRGML @W MQRCMAJ?QRQ. #MLC PCN?GP MAASPQ GL DMSP QR?ECQ ?LB A?L R?IC QCTCP?J KMLRFQ.
38.3 %F@EKJ 8E; .B<C<K8C (FM<D<EK

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
38.4 (LJ:C< CFEKI8:K@FE 8E; 'F:FDFK@FE

5FC @MBW AMLR?GLQ RFPCC RWNCQ MD KSQAJC RGQQSC: QICJCR?J KSQAJC, A?PBG?A KSQAJC, ?LB QKMMRF KSQAJC. 4ICJCRML KSQAJC RGQQSC GQ AMKNMQCB MD Q?PAMKCPCQ, RFC DSLARGML?J SLGRQ MD KSQAJC RGQQSC. .SQAJC AMLRP?ARGML MAASPQ UFCL Q?PAMKCPCQ QFMPRCL, ?Q RFGAI ?LB RFGL DGJ?KCLRQ QJGBC N?QR C?AF MRFCP, UFGAF GQ A?JJCB RFC QJGBGLE DGJ?KCLR KMBCJ MD KSQAJC AMLRP?ARGML. "51 NPMTGBCQ RFC CLCPEW DMP APMQQ-@PGBEC DMPK?RGML ?LB DGJ?KCLR QJGBGLE. 3CESJ?RMPW NPMRCGLQ, QSAF ?Q RPMNMLGL ?LB RPMNMKWMQGL, AMLRPMJ APMQQ-
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@PGBEC DMPK?RGML. &VAGR?RGMLpAMLRP?ARGML AMSNJGLE RP?LQBSACQ RFC CJCARPGA?J QGEL?J MD RFC LCSPML, TG? ?ACRWJAFMJGLC, RM ?L CJCARPGA?J QGEL?J ML RFC KSQAJC KCK@P?LC, UFGAF GLGRG?RCQ DMPAC NPMBSARGML. 5FC LSK@CP MD KSQAJC DG@CPQ AMLRP?ARGLE BCRCPKGLCQ FMU KSAF DMPAC RFC UFMJC KSQAJC NPMBSACQ.
A-/ C*)) C/$*) ,0 ./$*).

1. Figure 38.198FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR @MLC RGQQSC GQ D?JQC >. $MKN?AR @MLC RGQQSC GQ K?BC MD AWJGLBPGA?J MQRCMLQ RF?R ?PC ?JGELCB QSAF RF?R RFCW RP?TCJ RFC JCLERF MD RFC @MLC. ?. )?TCPQG?L A?L?JQ AMLR?GL @JMMB TCQQCJQ MLJW. @. )?TCPQG?L A?L?JQ AMLR?GL @JMMB TCQQCJQ ?LB LCPTC DG@CPQ. A. 4NMLEW RGQQSC GQ DMSLB ML RFC GLRCPGMP MD RFC @MLC, ?LB AMKN?AR @MLC RGQQSC GQ DMSLB ML RFC CVRCPGMP. 2. Figure 38.37 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR KSQAJC AMLRP?ARGML GQ RPSC >. 5FC NMUCP QRPMIC MAASPQ UFCL "51 GQ FWBPMJWXCB RM "%1 ?LB NFMQNF?RC. ?. 5FC NMUCP QRPMIC MAASPQ UFCL "%1 ?LB NFMQNF?RC BGQQMAG?RC DPMK RFC KWMQGL FC?B. @. 5FC NMUCP QRPMIC MAASPQ UFCL "%1 ?LB NFMQNF?RC BGQQMAG?RC DPMK RFC ?ARGL ?ARGTC QGRC. A. 5FC NMUCP QRPMIC MAASPQ UFCL $?2+ @GLBQ RFC A?JAGSK FC?B. 3. Figure 38.38 5FC BC?BJW LCPTC E?Q 4?PGL GPPCTCPQG@JW GLFG@GRQ ?ACRWAFMJGLCQRCP?QC. 8F?R CDDCAR UMSJB 4?PGL F?TC ML KSQAJC AMLRP?ARGML
- 1$ 2 ,0 ./$*).
4. 5FC DMPC?PK AMLQGQRQ MD RFC: >. P?BGSQ ?LB SJL? ?. P?BGSQ ?LB FSKCPSQ @. SJL? ?LB FSKCPSQ A. FSKCPSQ ?LB A?PNSQ 5. 5FC NCARMP?J EGPBJC AMLQGQRQ MD RFC: >. AJ?TGAJC ?LB QRCPLSK ?. QRCPLSK ?LB QA?NSJ? @. AJ?TGAJC ?LB QA?NSJ? A. AJ?TGAJC ?LB AMAAWV 6. "JJ MD RFC DMJJMUGLE ?PC EPMSNQ MD TCPRC@P?C CVACNR >>>>>>>>, UFGAF GQ ? ASPT?RSPC. >. RFMP?AGA ?. ACPTGA?J @. JSK@?P A. NCJTGA 7. 8FGAF MD RFCQC GQ ? D?AG?J @MLC >. DPMLR?J ?. MAAGNGR?J @. J?APGK?J A. RCKNMP?J 8. 5FC )?TCPQG?L A?L?J: >. GQ ?PP?LECB ?Q PMBQ MP NJ?RCQ ?. AMLR?GLQ RFC @MLCsQ @JMMB TCQQCJQ ?LB LCPTC DG@CPQ @. GQ PCQNMLQG@JC DMP RFC JCLERFUGQC EPMURF MD JMLE @MLCQ A. QWLRFCQGXCQ ?LB QCAPCRCQ K?RPGV 9. 5FC CNGNFWQC?J NJ?RC: >. GQ ?PP?LECB ?Q PMBQ MP NJ?RCQ ?. AMLR?GLQ RFC @MLCsQ @JMMB TCQQCJQ ?LB LCPTC DG@CPQ @. GQ PCQNMLQG@JC DMP RFC JCLERFUGQC EPMURF MD JMLE @MLCQ A. QWLRFCQGXCQ ?LB QCAPCRCQ @MLC K?RPGV 10. 5FC ACJJQ PCQNMLQG@JC DMP @MLC PCQMPNRGML ?PC >>>>>>>>. >. MQRCMAJ?QRQ ?. MQRCM@J?QRQ @. DG@PM@J?QRQ A. MQRCMAWRCQ 11. $MKN?AR @MLC GQ AMKNMQCB MD >>>>>>>>. >. ?. @. A. RP?@CASJ?C AMKN?ARCB AMJJ?ECL MQRCMLQ A?JAGSK NFMQNF?RC MLJW
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
#(!04%2 38 X 4(% -53#5,/3+%,%4!, 3934%-
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16. *L PCJ?VCB KSQAJC, RFC KWMQGL-@GLBGLE QGRC ML ?ARGL GQ @JMAICB @W >>>>>>>>. >. RGRGL ?. RPMNMLGL @. KWMEJM@GL A. RPMNMKWMQGL 17. 5FC ACJJ KCK@P?LC MD ? KSQAJC DG@CP GQ A?JJCB ? >>>>>>>>. >. KWMDG@PGJ ?. Q?PAMJCKK? @. Q?PAMNJ?QK A. KWMDGJ?KCLR 18. 5FC KSQAJC PCJ?VCQ GD LM LCU LCPTC QGEL?J ?PPGTCQ. )MUCTCP RFC LCSPMRP?LQKGRRCP DPMK RFC
NPCTGMSQ QRGKSJ?RGML GQ QRGJJ NPCQCLR GL RFC QWL?NQC. 5FC ?ARGTGRW MD >>>>>>>> FCJNQ RM PCKMTC RFGQ LCSPMRP?LQKGRRCP. >. KWMQGL ?. ?ARGML NMRCLRG?J @. RPMNMKWMQGL A. ?ACRWJAFMJGLCQRCP?QC 19. 5FC ?@GJGRW MD ? KSQAJC RM ECLCP?RC RCLQGML GKKCBG?RCJW ?DRCP QRGKSJ?RGML GQ BCNCLBCLR ML: >. ?. @. A. KWMQGL GLRCP?ARGML UGRF RFC . JGLC MTCPJ?N MD KWMQGL ?LB ?ARGL ?ARGL ?RR?AFKCLRQ RM RFC ; JGLC LMLC MD RFC ?@MTC
C-$/$CA' /#$)&$)" ,0 ./$*).

20. 8F?R ?PC RFC K?HMP BGDDCPCLACQ @CRUCCL RFC K?JC NCJTGQ ?LB DCK?JC NCJTGQ RF?R NCPKGR AFGJB@GPRF GL DCK?JCQ 21. 8F?R ?PC RFC K?HMP BGDDCPCLACQ @CRUCCL RFC NCJTGA EGPBJC ?LB RFC NCARMP?J EGPBJC RF?R ?JJMU RFC NCJTGA EGPBJC RM @C?P RFC UCGEFR MD RFC @MBW 22. 8F?R ?PC RFC K?HMP BGDDCPCLACQ @CRUCCL QNMLEW @MLC ?LB AMKN?AR @MLC 23. 8F?R ?PC RFC PMJCQ MD MQRCM@J?QRQ, MQRCMAWRCQ, ?LB MQRCMAJ?QRQ 24. 8F?R KMTCKCLRQ MAASP ?R RFC FGN HMGLR ?LB ILCCQ ?Q WMS @CLB BMUL RM RMSAF WMSP RMCQ 25. 8F?R KMTCKCLR(Q) MAASP(Q) ?R RFC QA?NSJ?C UFCL WMS QFPSE WMSP QFMSJBCPQ 26. )MU UMSJB KSQAJC AMLRP?ARGMLQ @C ?DDCARCB GD "51 U?Q AMKNJCRCJW BCNJCRCB GL ? KSQAJC DG@CP 27. 8F?R D?ARMPQ AMLRPG@SRC RM RFC ?KMSLR MD RCLQGML NPMBSACB GL ?L GLBGTGBS?J KSQAJC DG@CP 28. 8F?R CDDCAR UGJJ JMU @JMMB A?JAGSK F?TC ML LCSPMLQ 8F?R CDDCAR UGJJ JMU @JMMB A?JAGSK F?TC ML QICJCR?J KSQAJCQ
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#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
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39 R /# - .+$-A/*-4 .4./ (
!@>LI< 39.1 ,RKDP, TEF@E >MMB>O >P KB>OIV QO>KPM>OBKQ QFPPRB PROOLRKAFKD QEB EB>OQ FK QEFP 8-O>V LC > ALD (IBCQ), >OB QEB @BKQO>I LOD>KP LC QEB OBPMFO>QLOV PVPQBJ. 4EB IBCQ IRKD FP PJ>IIBO QE>K QEB OFDEQ IRKD QL >@@LJJLA>QB PM>@B CLO QEB EB>OQ. ! ALDwP KLPB (OFDEQ) E>P > PIFQ LK QEB PFAB LC B>@E KLPQOFI. 7EBK QO>@HFKD > P@BKQ, QEB PIFQP LMBK, ?IL@HFKD QEB COLKQ LC QEB KLPQOFIP. 4EFP >IILTP QEB ALD QL BUE>IB QELRDE QEB KLT-LMBK >OB> LK QEB PFAB LC QEB KLPQOFIP TFQELRQ ILPFKD QEB P@BKQ QE>Q FP ?BFKD CLIILTBA. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V 'BLCC 3QB>OKP; @OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V #LOV :>KHBO)
C?8GK< 39.2: "8J O:?8E>< 8:IFJJ -<JG@I8KFIP .LI=8:<J 39.3: BI<8K?@E> 39.4: /I8EJGFIK F= "8J<J @E #LD8E BF;@CP !CL@;J
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#PC?RFGLE GQ ?L GLTMJSLR?PW CTCLR. )MU MDRCL ? @PC?RF GQ R?ICL ?LB FMU KSAF ?GP GQ GLF?JCB MP CVF?JCB ?PC RGEFRJW PCESJ?RCB @W RFC PCQNGP?RMPW ACLRCP GL RFC @P?GL. )SK?LQ, UFCL RFCW ?PCLsR CVCPRGLE RFCKQCJTCQ, @PC?RFC ?NNPMVGK?RCJW 15 RGKCQ NCP KGLSRC ML ?TCP?EC. $?LGLCQ, JGIC RFC BME GL Figure 39.1, F?TC ? PCQNGP?RMPW P?RC MD ?@MSR 15p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39.1 R .PJK<DJ F= "8J O:?8E><

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC N?QQ?EC MD ?GP DPMK RFC MSRQGBC CLTGPMLKCLR RM RFC JSLEQ z &VNJ?GL FMU RFC JSLEQ ?PC NPMRCARCB DPMK N?PRGASJ?RC K?RRCP 5FC NPGK?PW DSLARGML MD RFC PCQNGP?RMPW QWQRCK GQ RM BCJGTCP MVWECL RM RFC ACJJQ MD RFC @MBWsQ RGQQSCQ ?LB PCKMTC A?P@ML BGMVGBC, ? ACJJ U?QRC NPMBSAR. 5FC K?GL QRPSARSPCQ MD RFC FSK?L PCQNGP?RMPW QWQRCK ?PC RFC L?Q?J A?TGRW, RFC RP?AFC?, ?LB JSLEQ. "JJ ?CPM@GA MPE?LGQKQ PCOSGPC MVWECL RM A?PPW MSR RFCGP KCR?@MJGA DSLARGMLQ. "JMLE RFC CTMJSRGML?PW RPCC, BGDDCPCLR MPE?LGQKQ F?TC BCTGQCB BGDDCPCLR KC?LQ MD M@R?GLGLE MVWECL DPMK RFC QSPPMSLBGLE ?RKMQNFCPC. 5FC CLTGPMLKCLR GL UFGAF RFC ?LGK?J JGTCQ EPC?RJW BCRCPKGLCQ FMU ?L ?LGK?J PCQNGPCQ. 5FC AMKNJCVGRW MD RFC PCQNGP?RMPW QWQRCK GQ AMPPCJ?RCB UGRF RFC QGXC MD RFC MPE?LGQK. "Q ?LGK?J QGXC GLAPC?QCQ, BGDDSQGML BGQR?LACQ GLAPC?QC ?LB RFC P?RGM MD QSPD?AC ?PC? RM TMJSKC BPMNQ. *L SLGACJJSJ?P MPE?LGQKQ, BGDDSQGML ?APMQQ RFC ACJJ KCK@P?LC GQ QSDDGAGCLR DMP QSNNJWGLE MVWECL RM RFC ACJJ (Figure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
!@>LI< 39.2 4EB @BII LC QEB RKF@BIIRI>O >ID>B #,5;90*(90( =,5;90*6:( FP LKB LC QEB I>ODBPQ HKLTK, OB>@EFKD LKB QL CFSB @BKQFJBQBOP FK AF>JBQBO. ,FHB >II PFKDIB-@BIIBA LOD>KFPJP, #. =,5;90*6:( BU@E>KDBP D>PBP >@OLPP QEB @BII JBJ?O>KB.
D@I<:K D@==LJ@FE
'MP QK?JJ KSJRGACJJSJ?P MPE?LGQKQ, BGDDSQGML ?APMQQ RFC MSRCP KCK@P?LC GQ QSDDGAGCLR RM KCCR RFCGP MVWECL LCCBQ. (?Q CVAF?LEC @W BGPCAR BGDDSQGML ?APMQQ QSPD?AC KCK@P?LCQ GQ CDDGAGCLR DMP MPE?LGQKQ JCQQ RF?L 1 KK GL BG?KCRCP. *L QGKNJC MPE?LGQKQ, QSAF ?Q ALGB?PG?LQ ?LB DJ?RUMPKQ, CTCPW ACJJ GL RFC @MBW GQ AJMQC RM RFC CVRCPL?J CLTGPMLKCLR. 5FCGP ACJJQ ?PC ICNR KMGQR ?LB E?QCQ BGDDSQC OSGAIJW TG? BGPCAR BGDDSQGML. 'J?RUMPKQ ?PC QK?JJ, JGRCP?JJW DJ?R UMPKQ, UFGAF r@PC?RFCs RFPMSEF BGDDSQGML ?APMQQ RFC MSRCP KCK@P?LC (Figure 39.3). 5FC DJ?R QF?NC MD RFCQC MPE?LGQKQ GLAPC?QCQ RFC QSPD?AC ?PC? DMP BGDDSQGML, CLQSPGLE RF?R C?AF ACJJ UGRFGL RFC @MBW GQ AJMQC RM RFC MSRCP KCK@P?LC QSPD?AC ?LB F?Q ?AACQQ RM MVWECL. *D RFC DJ?RUMPK F?B ? AWJGLBPGA?J @MBW, RFCL RFC ACJJQ GL RFC ACLRCP UMSJB LMR @C ?@JC RM ECR MVWECL.
#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
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!@>LI< 39.3 4EFP CI>QTLOJwP MOL@BPP LC OBPMFO>QFLK TLOHP ?V AFCCRPFLK >@OLPP QEB LRQBO JBJ?O>KB. (@OBAFQ: 3QBMEBK #EFIAP)
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igure 39.4). (GJJQ ?PC RFGL RGQQSC DGJ?KCLRQ RF?R ?PC FGEFJW @P?LAFCB ?LB DMJBCB. 8FCL U?RCP N?QQCQ MTCP RFC EGJJQ, RFC BGQQMJTCB MVWECL GL U?RCP P?NGBJW BGDDSQCQ ?APMQQ RFC EGJJQ GLRM RFC @JMMBQRPC?K. 5FC AGPASJ?RMPW QWQRCK A?L RFCL A?PPW RFC MVWECL?RCB @JMMB RM RFC MRFCP N?PRQ MD RFC @MBW. *L ?LGK?JQ RF?R AMLR?GL AMCJMKGA DJSGB GLQRC?B MD @JMMB, MVWECL BGDDSQCQ ?APMQQ RFC EGJJ QSPD?ACQ GLRM RFC AMCJMKGA DJSGB. (GJJQ ?PC DMSLB GL KMJJSQIQ, ?LLCJGBQ, ?LB APSQR?AC?LQ.
!@>LI< 39.4 4EFP @LJJLK @>OM, IFHB J>KV LQEBO >NR>QF@ LOD>KFPJP, E>P DFIIP QE>Q >IILT FQ QL L?Q>FK LUVDBK COLJ T>QBO. (@OBAFQ: "'RFQ>OARAB012"/7FHFJBAF> #LJJLKP)
5FC DMJBCB QSPD?ACQ MD RFC EGJJQ NPMTGBC ? J?PEC QSPD?AC ?PC? RM CLQSPC RF?R RFC DGQF ECRQ QSDDGAGCLR MVWECL. %GDDSQGML GQ ? NPMACQQ GL UFGAF K?RCPG?J RP?TCJQ DPMK PCEGMLQ MD FGEF AMLACLRP?RGML RM JMU AMLACLRP?RGML SLRGJ COSGJG@PGSK GQ PC?AFCB. *L RFGQ A?QC, @JMMB UGRF ? JMU AMLACLRP?RGML MD MVWECL KMJCASJCQ AGPASJ?RCQ RFPMSEF RFC EGJJQ. 5FC AMLACLRP?RGML MD MVWECL KMJCASJCQ GL U?RCP GQ FGEFCP RF?L RFC AMLACLRP?RGML MD MVWECL KMJCASJCQ GL EGJJQ. "Q ? PCQSJR, MVWECL KMJCASJCQ BGDDSQC DPMK U?RCP (FGEF AMLACLRP?RGML) RM @JMMB (JMU AMLACLRP?RGML), ?Q QFMUL GL Figure 39.5. 4GKGJ?PJW, A?P@ML BGMVGBC KMJCASJCQ GL RFC @JMMB BGDDSQC DPMK RFC @JMMB (FGEF AMLACLRP?RGML) RM U?RCP (JMU AMLACLRP?RGML).
1138
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!@>LI< 39.5 !P T>QBO CILTP LSBO QEB DFIIP, LUVDBK FP QO>KPCBOOBA QL ?ILLA SF> QEB SBFKP. (@OBAFQ "CFPE": JLAFCF@>QFLK LC TLOH ?V $R>KB 2>SBO, ./!!)
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igure 39.6) ?LB PCESJ?RGLE RFC BGDDSQGML MD $02 ?LB U?RCP T?NMP. "GP CLRCPQ ?LB JC?TCQ RFC RP?AFC?J QWQRCK RFPMSEF RFC QNGP?AJCQ. 4MKC GLQCARQ A?L TCLRGJ?RC RFC RP?AFC?J QWQRCK UGRF @MBW KMTCKCLRQ.
!@>LI< 39.6 )KPB@QP MBOCLOJ OBPMFO>QFLK SF> > QO>@EB>I PVPQBJ.
(8DD8C@8E .PJK<DJ
*L K?KK?JQ, NSJKML?PW TCLRGJ?RGML MAASPQ TG? GLF?J?RGML (@PC?RFGLE). %SPGLE GLF?J?RGML, ?GP CLRCPQ RFC @MBW RFPMSEF RFC nasal cavity JMA?RCB HSQR GLQGBC RFC LMQC (Figure 39.7). "Q ?GP N?QQCQ RFPMSEF RFC L?Q?J A?TGRW, RFC ?GP GQ U?PKCB RM @MBW RCKNCP?RSPC ?LB FSKGBGDGCB. 5FC PCQNGP?RMPW RP?AR GQ AM?RCB UGRF KSASQ RM QC?J RFC RGQQSCQ DPMK BGPCAR AMLR?AR UGRF ?GP. .SASQ GQ FGEF GL U?RCP. "Q ?GP APMQQCQ RFCQC QSPD?ACQ MD RFC KSAMSQ KCK@P?LCQ, GR NGAIQ SN U?RCP. 5FCQC NPMACQQCQ FCJN COSGJG@P?RC RFC ?GP RM RFC @MBW AMLBGRGMLQ, PCBSAGLE ?LW B?K?EC RF?R AMJB, BPW ?GP A?L A?SQC. 1?PRGASJ?RC K?RRCP RF?R GQ DJM?RGLE GL RFC ?GP GQ PCKMTCB GL RFC L?Q?J N?QQ?ECQ TG? KSASQ ?LB AGJG?. 5FC NPMACQQCQ MD U?PKGLE, FSKGBGDWGLE, ?LB PCKMTGLE N?PRGAJCQ ?PC GKNMPR?LR NPMRCARGTC KCAF?LGQKQ RF?R NPCTCLR B?K?EC RM RFC RP?AFC? ?LB JSLEQ. 5FSQ, GLF?J?RGML QCPTCQ QCTCP?J NSPNMQCQ GL ?BBGRGML RM @PGLEGLE MVWECL GLRM RFC PCQNGP?RMPW QWQRCK.
#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
1139
!@>LI< 39.7 !FO BKQBOP QEB OBPMFO>QLOV PVPQBJ QEOLRDE QEB K>P>I @>SFQV >KA ME>OVKU, >KA QEBK M>PPBP QEOLRDE QEB QO>@EB> >KA FKQL QEB ?OLK@EF, TEF@E ?OFKD >FO FKQL QEB IRKDP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .#))
7EF@E LC QEB CLIILTFKD PQ>QBJBKQP >?LRQ QEB J>JJ>IF>K OBPMFO>QLOV PVPQBJ FP C>IPB >. 7EBK TB ?OB>QEB FK, >FO QO>SBIP COLJ QEB ME>OVKU QL QEB QO>@EB>. ?. 4EB ?OLK@EFLIBP ?O>K@E FKQL ?OLK@EF. @. !ISBLI>O AR@QP @LKKB@Q QL >ISBLI>O P>@P. A. '>P BU@E>KDB ?BQTBBK QEB IRKD >KA ?ILLA Q>HBP MI>@B FK QEB >ISBLIRP. 'PMK RFC L?Q?J A?TGRW, ?GP N?QQCQ RFPMSEF RFC pharynx (RFPM?R) ?LB RFC larynx (TMGAC @MV), ?Q GR K?ICQ GRQ U?W RM RFC trachea (Figure 39.7). 5FC K?GL DSLARGML MD RFC RP?AFC? GQ RM DSLLCJ RFC GLF?JCB ?GP RM RFC JSLEQ ?LB RFC CVF?JCB ?GP @?AI MSR MD RFC @MBW. 5FC FSK?L RP?AFC? GQ ? AWJGLBCP ?@MSR 10 RM 12 AK JMLE ?LB 2 AK GL BG?KCRCP RF?R QGRQ GL DPMLR MD RFC CQMNF?ESQ ?LB CVRCLBQ DPMK RFC J?PWLV GLRM RFC AFCQR A?TGRW UFCPC GR BGTGBCQ GLRM RFC RUM NPGK?PW @PMLAFG ?R RFC KGBRFMP?V. *R GQ K?BC MD GLAMKNJCRC PGLEQ MD FW?JGLC A?PRGJ?EC ?LB QKMMRF KSQAJC (Figure 39.8). 5FC RP?AFC? GQ JGLCB UGRF KSASQ-NPMBSAGLE EM@JCR ACJJQ ?LB AGJG?RCB CNGRFCJG?. 5FC AGJG? NPMNCJ DMPCGEL N?PRGAJCQ RP?NNCB GL RFC KSASQ RMU?PB RFC NF?PWLV. 5FC A?PRGJ?EC NPMTGBCQ QRPCLERF ?LB QSNNMPR RM RFC RP?AFC? RM ICCN RFC N?QQ?EC MNCL. 5FC QKMMRF KSQAJC A?L AMLRP?AR, BCAPC?QGLE RFC RP?AFC?sQ BG?KCRCP, UFGAF A?SQCQ CVNGPCB ?GP RM PSQF SNU?PBQ DPMK RFC JSLEQ ?R ? EPC?R DMPAC. 5FC DMPACB CVF?J?RGML FCJNQ CVNCJ KSASQ UFCL UC AMSEF. 4KMMRF KSQAJC A?L AMLRP?AR MP PCJ?V, BCNCLBGLE ML QRGKSJG DPMK RFC CVRCPL?J CLTGPMLKCLR MP RFC @MBWsQ LCPTMSQ QWQRCK.
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!@>LI< 39.8 4EB QO>@EB> >KA ?OLK@EF >OB J>AB LC FK@LJMIBQB OFKDP LC @>OQFI>DB. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V 'O>V'P !K>QLJV)
L:3-8: B743)./ '3* A1;+41/ 5FC CLB MD RFC RP?AFC? @GDSPA?RCQ (BGTGBCQ) RM RFC PGEFR ?LB JCDR JSLEQ. 5FC JSLEQ ?PC LMR GBCLRGA?J. 5FC PGEFR JSLE GQ J?PECP ?LB AMLR?GLQ RFPCC JM@CQ, UFCPC?Q RFC QK?JJCP JCDR JSLE AMLR?GLQ RUM JM@CQ (Figure 39.9). 5FC KSQASJ?P diaphragm, UFGAF D?AGJGR?RCQ @PC?RFGLE, GQ GLDCPGMP (@CJMU) RM RFC JSLEQ ?LB K?PIQ RFC CLB MD RFC RFMP?AGA A?TGRW.
!@>LI< 39.9 4EB QO>@EB> ?FCRO@>QBP FKQL QEB OFDEQ >KA IBCQ ?OLK@EF FK QEB IRKDP. 4EB OFDEQ IRKD FP J>AB LC QEOBB IL?BP >KA FP I>ODBO. 4L >@@LJJLA>QB QEB EB>OQ, QEB IBCQ IRKD FP PJ>IIBO >KA E>P LKIV QTL IL?BP.
*L RFC JSLEQ, ?GP GQ BGTCPRCB GLRM QK?JJCP ?LB QK?JJCP N?QQ?ECQ, MP bronchi. "GP CLRCPQ RFC JSLEQ RFPMSEF RFC RUM primary (main) bronchi (QGLESJ?P: @PMLAFSQ). &?AF @PMLAFSQ BGTGBCQ GLRM QCAMLB?PW @PMLAFG, RFCL GLRM RCPRG?PW @PMLAFG, UFGAF GL RSPL BGTGBC, APC?RGLE QK?JJCP ?LB QK?JJCP BG?KCRCP bronchioles ?Q RFCW QNJGR ?LB QNPC?B RFPMSEF RFC JSLE. -GIC RFC RP?AFC?, RFC @PMLAFG ?PC K?BC MD A?PRGJ?EC ?LB QKMMRF KSQAJC. "R RFC @PMLAFGMJCQ, RFC A?PRGJ?EC GQ PCNJ?ACB UGRF CJ?QRGA DG@CPQ. #PMLAFG ?PC GLLCPT?RCB @W LCPTCQ MD @MRF RFC N?P?QWKN?RFCRGA ?LB QWKN?RFCRGA LCPTMSQ QWQRCKQ RF?R AMLRPMJ KSQAJC AMLRP?ARGML (N?P?QWKN?RFCRGA) MP PCJ?V?RGML (QWKN?RFCRGA) GL RFC @PMLAFG ?LB @PMLAFGMJCQ, BCNCLBGLE ML RFC LCPTMSQ QWQRCKsQ ASCQ. *L FSK?LQ, @PMLAFGMJCQ UGRF ? BG?KCRCP QK?JJCP RF?L 0.5 KK ?PC RFC respiratory bronchioles. 5FCW J?AI A?PRGJ?EC ?LB RFCPCDMPC PCJW ML GLF?JCB ?GP RM QSNNMPR RFCGP QF?NC. "Q RFC N?QQ?ECU?WQ BCAPC?QC GL BG?KCRCP, RFC PCJ?RGTC ?KMSLR MD QKMMRF KSQAJC GLAPC?QCQ. 5FC terminal bronchioles QS@BGTGBC GLRM KGAPMQAMNGA @P?LAFCQ A?JJCB PCQNGP?RMPW @PMLAFGMJCQ. 5FC PCQNGP?RMPW @PMLAFGMJCQ QS@BGTGBC GLRM QCTCP?J ?JTCMJ?P BSARQ. /SKCPMSQ ?JTCMJG ?LB ?JTCMJ?P Q?AQ QSPPMSLB RFC ?JTCMJ?P BSARQ. 5FC ?JTCMJ?P Q?AQ PCQCK@JC @SLAFCQ MD EP?NCQ RCRFCPCB RM RFC CLB MD RFC @PMLAFGMJCQ (Figure 39.10). *L RFC ?AGL?P PCEGML, RFC alveolar ducts ?PC ?RR?AFCB RM RFC CLB MD C?AF @PMLAFGMJC. "R RFC CLB MD C?AF BSAR ?PC ?NNPMVGK?RCJW 100 alveolar sacs, C?AF AMLR?GLGLE 20 RM 30
#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
1141
alveoli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
!@>LI< 39.10 4BOJFK>I ?OLK@EFLIBP >OB @LKKB@QBA ?V OBPMFO>QLOV ?OLK@EFLIBP QL >ISBLI>O AR@QP >KA >ISBLI>O P>@P. %>@E >ISBLI>O P>@ @LKQ>FKP 20 QL 30 PMEBOF@>I >ISBLIF >KA E>P QEB >MMB>O>K@B LC > ?RK@E LC DO>MBP. !FO CILTP FKQL QEB >QOFRJ LC QEB >ISBLI>O P>@, QEBK @FO@RI>QBP FKQL >ISBLIF TEBOB D>P BU@E>KDB L@@ROP TFQE QEB @>MFII>OFBP. -R@LRP DI>KAP PB@OBQB JR@LRP FKQL QEB >FOT>VP, HBBMFKD QEBJ JLFPQ >KA CIBUF?IB. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
8?RAF RFC DMJJMUGLE video (http://openstaxcollege.org/l/lungs_pulmonary) RM PCTGCU RFC PCQNGP?RMPW QWQRCK.
+IFK<:K@M< (<:?8E@JDJ
5FC ?GP RF?R MPE?LGQKQ @PC?RFC AMLR?GLQ particulate matter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mucusq? QRGAIW QS@QR?LAC K?BC
1142
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MD mucin, ? AMKNJCV EJWAMNPMRCGL, ?Q UCJJ ?Q Q?JRQ ?LB U?RCPqRF?R RP?NQ N?PRGASJ?RCQ. 5FC @PMLAFG ?LB @PMLAFGMJCQ AMLR?GL AGJG?, QK?JJ F?GP-JGIC NPMHCARGMLQ RF?R JGLC RFC U?JJQ MD RFC @PMLAFG ?LB @PMLAFGMJCQ (Figure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
!@>LI< 39.11 4EB ?OLK@EF >KA ?OLK@EFLIBP @LKQ>FK @FIF> QE>Q EBIM JLSB JR@RP >KA LQEBO M>OQF@IBP LRQ LC QEB IRKDP. (@OBAFQ: ,LRFP> (LT>OA, JLAFCF@>QFLK LC TLOH ?V $>OQJLRQE %IB@QOLK -F@OLP@LMB &>@FIFQV)
39.2 R "8J O:?8E>< 8:IFJJ -<JG@I8KFIP .LI=8:<J

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z /?KC ?LB BCQAPG@C JSLE TMJSKCQ ?LB A?N?AGRGCQ z 6LBCPQR?LB FMU E?Q NPCQQSPC GLDJSCLACQ FMU E?QCQ KMTC GLRM ?LB MSR MD RFC @MBW 5FC QRPSARSPC MD RFC JSLE K?VGKGXCQ GRQ QSPD?AC ?PC? RM GLAPC?QC E?Q BGDDSQGML. #CA?SQC MD RFC CLMPKMSQ LSK@CP MD ?JTCMJG (?NNPMVGK?RCJW 300 KGJJGML GL C?AF FSK?L JSLE), RFC QSPD?AC ?PC? MD RFC JSLE GQ TCPW J?PEC (75 K2). )?TGLE QSAF ? J?PEC QSPD?AC ?PC? GLAPC?QCQ RFC ?KMSLR MD E?Q RF?R A?L BGDDSQC GLRM ?LB MSR MD RFC JSLEQ.
B8J@: +I@E:@GC<J F= "8J O:?8E><

(?Q CVAF?LEC BSPGLE PCQNGP?RGML MAASPQ NPGK?PGJW RFPMSEF BGDDSQGML. %GDDSQGML GQ ? NPMACQQ GL UFGAF RP?LQNMPR GQ BPGTCL @W ? AMLACLRP?RGML EP?BGCLR. (?Q KMJCASJCQ KMTC DPMK ? PCEGML MD FGEF AMLACLRP?RGML RM ? PCEGML MD JMU AMLACLRP?RGML. #JMMB RF?R GQ JMU GL MVWECL AMLACLRP?RGML ?LB FGEF GL A?P@ML BGMVGBC AMLACLRP?RGML SLBCPEMCQ E?Q CVAF?LEC UGRF ?GP GL RFC JSLEQ. 5FC ?GP GL RFC JSLEQ F?Q ? FGEFCP AMLACLRP?RGML MD MVWECL RF?L RF?R MD MVWECL-BCNJCRCB @JMMB ?LB ? JMUCP AMLACLRP?RGML MD A?P@ML BGMVGBC. 5FGQ AMLACLRP?RGML EP?BGCLR ?JJMUQ DMP E?Q CVAF?LEC BSPGLE PCQNGP?RGML. Partial pressure GQ ? KC?QSPC MD RFC AMLACLRP?RGML MD RFC GLBGTGBS?J AMKNMLCLRQ GL ? KGVRSPC MD E?QCQ. 5FC RMR?J NPCQQSPC CVCPRCB @W RFC KGVRSPC GQ RFC QSK MD RFC N?PRG?J NPCQQSPCQ MD RFC AMKNMLCLRQ GL RFC KGVRSPC. 5FC P?RC MD BGDDSQGML MD ? E?Q GQ NPMNMPRGML?J RM GRQ N?PRG?J NPCQQSPC UGRFGL RFC RMR?J E?Q KGVRSPC. 5FGQ AMLACNR GQ BGQASQQCB DSPRFCP GL BCR?GJ @CJMU.
'LE> 1FCLD<J 8E; C8G8:@K@<J

%GDDCPCLR ?LGK?JQ F?TC BGDDCPCLR JSLE A?N?AGRGCQ @?QCB ML RFCGP ?ARGTGRGCQ. $FCCR?FQ F?TC CTMJTCB ? KSAF FGEFCP JSLE A?N?AGRW RF?L FSK?LQ; GR FCJNQ NPMTGBC MVWECL RM ?JJ RFC KSQAJCQ GL RFC @MBW ?LB ?JJMUQ RFCK RM PSL TCPW D?QR. &JCNF?LRQ ?JQM F?TC ? FGEF JSLE A?N?AGRW. *L RFGQ A?QC, GR GQ LMR @CA?SQC RFCW PSL D?QR @SR @CA?SQC RFCW F?TC ? J?PEC @MBW ?LB KSQR @C ?@JC RM R?IC SN MVWECL GL ?AAMPB?LAC UGRF RFCGP @MBW QGXC.
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1143
)SK?L JSLE QGXC GQ BCRCPKGLCB @W ECLCRGAQ, ECLBCP, ?LB FCGEFR. "R K?VGK?J A?N?AGRW, ?L ?TCP?EC JSLE A?L FMJB ?JKMQR QGV JGRCPQ MD ?GP, @SR JSLEQ BM LMR SQS?JJW MNCP?RC ?R K?VGK?J A?N?AGRW. "GP GL RFC JSLEQ GQ KC?QSPCB GL RCPKQ MD lung volumes ?LB lung capacities (Figure 39.12 ?LB Table 39.1). 7MJSKC KC?QSPCQ RFC ?KMSLR MD ?GP DMP MLC DSLARGML (QSAF ?Q GLF?J?RGML MP CVF?J?RGML). $?N?AGRW GQ ?LW RUM MP KMPC TMJSKCQ (DMP CV?KNJC, FMU KSAF A?L @C GLF?JCB DPMK RFC CLB MD ? K?VGK?J CVF?J?RGML).
!@>LI< 39.12 (RJ>K IRKD SLIRJBP >KA @>M>@FQFBP >OB PELTK. 4EB QLQ>I IRKD @>M>@FQV LC QEB >ARIQ J>IB FP PFU IFQBOP. 4FA>I SLIRJB FP QEB SLIRJB LC >FO FKE>IBA FK > PFKDIB, KLOJ>I ?OB>QE. )KPMFO>QLOV @>M>@FQV FP QEB >JLRKQ LC >FO Q>HBK FK AROFKD > ABBM ?OB>QE, >KA OBPFAR>I SLIRJB FP QEB >JLRKQ LC >FO IBCQ FK QEB IRKDP >CQBO CLO@BCRI OBPMFO>QFLK.
'LE> 1FCLD<J 8E; C8G8:@K@<J (AM> A;LCK (8C<)

1FCLD</ C8G8:@KP
4FA>I SLIRJB (46)
D<=@E@K@FE
!JLRKQ LC >FO FKE>IBA AROFKD > KLOJ>I ?OB>QE
1FCLD< (C@K<IJ)
0.5 1.2 -
HL8K@FEJ
%UMFO>QLOV !JLRKQ LC >FO QE>Q @>K ?B BUE>IBA >CQBO > OBPBOSB SLIRJB KLOJ>I BUE>I>QFLK (%26) )KPMFO>QLOV !JLRKQ LC >FO QE>Q @>K ?B CROQEBO FKE>IBA OBPBOSB SLIRJB >CQBO > KLOJ>I FKE>I>QFLK ()26) 2BPFAR>I SLIRJB (26) 6FQ>I @>M>@FQV (6#) )KPMFO>QLOV @>M>@FQV ()#) !FO IBCQ FK QEB IRKDP >CQBO > CLO@BA BUE>I>QFLK
3.1 1.2
%26+46+)26 46+)26 %26+26 26+%26+46+)26 -
->UFJRJ >JLRKQ LC >FO QE>Q @>K ?B JLSBA FK LO LRQ LC QEB IRKDP FK > PFKDIB OBPMFO>QLOV 4.8 @V@IB 6LIRJB LC >FO QE>Q @>K ?B FKE>IBA FK >AAFQFLK QL > KLOJ>I BUE>I>QFLK 3.6 2.4 6.0 Y4.1 QL 5.5
&RK@QFLK>I 6LIRJB LC >FO OBJ>FKFKD >CQBO > KLOJ>I OBPFAR>I BUE>I>QFLK @>M>@FQV (&2#) 4LQ>I IRKD @>M>@FQV (4,#) 4LQ>I SLIRJB LC >FO FK QEB IRKDP >CQBO > J>UFJ>I FKPMFO>QFLK
&LO@BA (LT JR@E >FO @>K ?B CLO@BA LRQ LC QEB BUMFO>QLOV IRKDP LSBO > PMB@FCF@ QFJB MBOFLA, RPR>IIV SLIRJB (&%61) LKB PB@LKA /89C< 39.1
1144
#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
5FC TMJSKC GL RFC JSLE A?L @C BGTGBCB GLRM DMSP SLGRQ: RGB?J TMJSKC, CVNGP?RMPW PCQCPTC TMJSKC, GLQNGP?RMPW PCQCPTC TMJSKC, ?LB PCQGBS?J TMJSKC. Tidal volume (TV) KC?QSPCQ RFC ?KMSLR MD ?GP RF?R GQ GLQNGPCB ?LB CVNGPCB BSPGLE ? LMPK?J @PC?RF. 0L ?TCP?EC, RFGQ TMJSKC GQ ?PMSLB MLC-F?JD JGRCP, UFGAF GQ ? JGRRJC JCQQ RF?L RFC A?N?AGRW MD ? 20-MSLAC BPGLI @MRRJC. 5FC expiratory reserve volume (ERV) GQ RFC ?BBGRGML?J ?KMSLR MD ?GP RF?R A?L @C CVF?JCB ?DRCP ? LMPK?J CVF?J?RGML. *R GQ RFC PCQCPTC ?KMSLR RF?R A?L @C CVF?JCB @CWMLB UF?R GQ LMPK?J. $MLTCPQCJW, RFC inspiratory reserve volume (IRV) GQ RFC ?BBGRGML?J ?KMSLR MD ?GP RF?R A?L @C GLF?JCB ?DRCP ? LMPK?J GLF?J?RGML. 5FC residual volume (RV) GQ RFC ?KMSLR MD ?GP RF?R GQ JCDR ?DRCP CVNGP?RMPW PCQCPTC TMJSKC GQ CVF?JCB. 5FC JSLEQ ?PC LCTCP AMKNJCRCJW CKNRW: 5FCPC GQ ?JU?WQ QMKC ?GP JCDR GL RFC JSLEQ ?DRCP ? K?VGK?J CVF?J?RGML. *D RFGQ PCQGBS?J TMJSKC BGB LMR CVGQR ?LB RFC JSLEQ CKNRGCB AMKNJCRCJW, RFC JSLE RGQQSCQ UMSJB QRGAI RMECRFCP ?LB RFC CLCPEW LCACQQ?PW RM PC-GLDJ?RC RFC JSLE AMSJB @C RMM EPC?R RM MTCPAMKC. 5FCPCDMPC, RFCPC GQ ?JU?WQ QMKC ?GP PCK?GLGLE GL RFC JSLEQ. 3CQGBS?J TMJSKC GQ ?JQM GKNMPR?LR DMP NPCTCLRGLE J?PEC DJSARS?RGMLQ GL PCQNGP?RMPW E?QCQ (02 ?LB $02). 5FC PCQGBS?J TMJSKC GQ RFC MLJW JSLE TMJSKC RF?R A?LLMR @C KC?QSPCB BGPCARJW @CA?SQC GR GQ GKNMQQG@JC RM AMKNJCRCJW CKNRW RFC JSLE MD ?GP. 5FGQ TMJSKC A?L MLJW @C A?JASJ?RCB P?RFCP RF?L KC?QSPCB. $?N?AGRGCQ ?PC KC?QSPCKCLRQ MD RUM MP KMPC TMJSKCQ. 5FC vital capacity (VC) KC?QSPCQ RFC K?VGKSK ?KMSLR MD ?GP RF?R A?L @C GLF?JCB MP CVF?JCB BSPGLE ? PCQNGP?RMPW AWAJC. *R GQ RFC QSK MD RFC CVNGP?RMPW PCQCPTC TMJSKC, RGB?J TMJSKC, ?LB GLQNGP?RMPW PCQCPTC TMJSKC. 5FC inspiratory capacity (IC) GQ RFC ?KMSLR MD ?GP RF?R A?L @C GLF?JCB ?DRCP RFC CLB MD ? LMPK?J CVNGP?RGML. *R GQ, RFCPCDMPC, RFC QSK MD RFC RGB?J TMJSKC ?LB GLQNGP?RMPW PCQCPTC TMJSKC. 5FC functional residual capacity (FRC) GLAJSBCQ RFC CVNGP?RMPW PCQCPTC TMJSKC ?LB RFC PCQGBS?J TMJSKC. 5FC '3$ KC?QSPCQ RFC ?KMSLR MD ?BBGRGML?J ?GP RF?R A?L @C CVF?JCB ?DRCP ? LMPK?J CVF?J?RGML. -?QRJW, RFC total lung capacity (TLC) GQ ? KC?QSPCKCLR MD RFC RMR?J ?KMSLR MD ?GP RF?R RFC JSLE A?L FMJB. *R GQ RFC QSK MD RFC PCQGBS?J TMJSKC, CVNGP?RMPW PCQCPTC TMJSKC, RGB?J TMJSKC, ?LB GLQNGP?RMPW PCQCPTC TMJSKC. -SLE TMJSKCQ ?PC KC?QSPCB @W ? RCAFLGOSC A?JJCB spirometry. "L GKNMPR?LR KC?QSPCKCLR R?ICL BSPGLE QNGPMKCRPW GQ RFC forced expiratory volume (FEV), UFGAF KC?QSPCQ FMU KSAF ?GP A?L @C DMPACB MSR MD RFC JSLE MTCP ? QNCAGDGA NCPGMB, SQS?JJW MLC QCAMLB ('&71). *L ?BBGRGML, RFC DMPACB TGR?J A?N?AGRW ('7$), UFGAF GQ RFC RMR?J ?KMSLR MD ?GP RF?R A?L @C DMPAG@JW CVF?JCB, GQ KC?QSPCB. 5FC P?RGM MD RFCQC T?JSCQ ( FEV1/FVC ratio) GQ SQCB RM BG?ELMQC JSLE BGQC?QCQ GLAJSBGLE ?QRFK?, CKNFWQCK?, ?LB DG@PMQGQ. *D RFC '&71/'7$ P?RGM GQ FGEF, RFC JSLEQ ?PC LMR AMKNJG?LR (KC?LGLE RFCW ?PC QRGDD ?LB SL?@JC RM @CLB NPMNCPJW), ?LB RFC N?RGCLR KMQR JGICJW F?Q JSLE DG@PMQGQ. 1?RGCLRQ CVF?JC KMQR MD RFC JSLE TMJSKC TCPW OSGAIJW. $MLTCPQCJW, UFCL RFC '&71/'7$ P?RGM GQ JMU, RFCPC GQ PCQGQR?LAC GL RFC JSLE RF?R GQ AF?P?ARCPGQRGA MD ?QRFK?. *L RFGQ GLQR?LAC, GR GQ F?PB DMP RFC N?RGCLR RM ECR RFC ?GP MSR MD FGQ MP FCP JSLEQ, ?LB GR R?ICQ ? JMLE RGKC RM PC?AF RFC K?VGK?J CVF?J?RGML TMJSKC. *L CGRFCP A?QC, @PC?RFGLE GQ BGDDGASJR ?LB AMKNJGA?RGMLQ ?PGQC.
-<JG@I8KFIP /?<I8G@JK
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wP ABDOBB TFQE > OBPMFO>QLOV QEBO>MFPQ PMB@F>IQV. "B@>RPB LC > DOLTFKD >DFKD MLMRI>QFLK, @>OBBO LMMLOQRKFQFBP >P > OBPMFO>QLOV QEBO>MFPQ >OB BUMB@QBA QL OBJ>FK PQOLKD.
"8J +I<JJLI< 8E; -<JG@I8K@FE

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
#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
1145
P = (P atm ) (percent content in mixture).

1?RK, RFC ?RKMQNFCPGA NPCQQSPC, GQ RFC QSK MD ?JJ MD RFC N?PRG?J NPCQQSPCQ MD RFC ?RKMQNFCPGA E?QCQ ?BBCB RMECRFCP,
P atm = P N + P O + P H
2 2
2O
+ P CO = 760 mm Hg
2
` (NCPACLR AMLRCLR GL KGVRSPC). 5FC NPCQQSPC MD RFC ?RKMQNFCPC ?R QC? JCTCJ GQ 760 KK )E. 5FCPCDMPC, RFC N?PRG?J NPCQQSPC MD MVWECL GQ: ?LB DMP A?P@ML BGMVGBC:
P O = (760 mm Hg) (0.21) = 160 mm Hg

2
P CO = (760 mm Hg) (0.0004) = 0.3 mm Hg.

2
"R FGEF ?JRGRSBCQ, 1?RK BCAPC?QCQ @SR AMLACLRP?RGML BMCQ LMR AF?LEC; RFC N?PRG?J NPCQQSPC BCAPC?QC GQ BSC RM RFC PCBSARGML GL 1?RK. 8FCL RFC ?GP KGVRSPC PC?AFCQ RFC JSLE, GR F?Q @CCL FSKGBGDGCB. 5FC NPCQQSPC MD RFC U?RCP T?NMP GL RFC JSLE BMCQ LMR AF?LEC RFC NPCQQSPC MD RFC ?GP, @SR GR KSQR @C GLAJSBCB GL RFC N?PRG?J NPCQQSPC COS?RGML. 'MP RFGQ A?JASJ?RGML, RFC U?RCP NPCQQSPC (47 KK )E) GQ QS@RP?ARCB DPMK RFC ?RKMQNFCPGA NPCQQSPC:
760 mm Hg 47 mm Hg = 713 mm Hg
?LB RFC N?PRG?J NPCQQSPC MD MVWECL GQ:
(760 mm Hg 47 mm Hg) 0.21 = 150 mm Hg.
5FCQC NPCQQSPCQ BCRCPKGLC RFC E?Q CVAF?LEC, MP RFC DJMU MD E?Q, GL RFC QWQRCK. 0VWECL ?LB A?P@ML BGMVGBC UGJJ DJMU ?AAMPBGLE RM RFCGP NPCQQSPC EP?BGCLR DPMK FGEF RM JMU. 5FCPCDMPC, SLBCPQR?LBGLE RFC N?PRG?J NPCQQSPC MD C?AF E?Q UGJJ ?GB GL SLBCPQR?LBGLE FMU E?QCQ KMTC GL RFC PCQNGP?RMPW QWQRCK.
"8J O:?8E>< 8:IFJJ K?< ACM<FC@

*L RFC @MBW, MVWECL GQ SQCB @W ACJJQ MD RFC @MBWsQ RGQQSCQ ?LB A?P@ML BGMVGBC GQ NPMBSACB ?Q ? U?QRC NPMBSAR. 5FC P?RGM MD A?P@ML BGMVGBC NPMBSARGML RM MVWECL AMLQSKNRGML GQ RFC respiratory quotient (RQ). 32 T?PGCQ @CRUCCL 0.7 ?LB 1.0. *D HSQR EJSAMQC UCPC SQCB RM DSCJ RFC @MBW, RFC 32 UMSJB COS?J MLC. 0LC KMJC MD A?P@ML BGMVGBC UMSJB @C NPMBSACB DMP CTCPW KMJC MD MVWECL AMLQSKCB. (JSAMQC, FMUCTCP, GQ LMR RFC MLJW DSCJ DMP RFC @MBW. 1PMRCGL ?LB D?R ?PC ?JQM SQCB ?Q DSCJQ DMP RFC @MBW. #CA?SQC MD RFGQ, JCQQ A?P@ML BGMVGBC GQ NPMBSACB RF?L MVWECL GQ AMLQSKCB ?LB RFC 32 GQ, ML ?TCP?EC, ?@MSR 0.7 DMP D?R ?LB ?@MSR 0.8 DMP NPMRCGL. 5FC 32 GQ SQCB RM A?JASJ?RC RFC N?PRG?J NPCQQSPC MD MVWECL GL RFC ?JTCMJ?P QN?ACQ UGRFGL RFC JSLE, RFC alveolar PO "@MTC, RFC N?PRG?J NPCQQSPC MD MVWECL GL RFC JSLEQ U?Q A?JASJ?RCB RM @C 150 KK
2
)E. )MUCTCP, JSLEQ LCTCP DSJJW BCDJ?RC UGRF ?L CVF?J?RGML; RFCPCDMPC, RFC GLQNGPCB ?GP KGVCQ UGRF RFGQ PCQGBS?J ?GP ?LB JMUCPQ RFC N?PRG?J NPCQQSPC MD MVWECL UGRFGL RFC ?JTCMJG. 5FGQ KC?LQ RF?R RFCPC GQ ? JMUCP AMLACLRP?RGML MD MVWECL GL RFC JSLEQ RF?L GQ DMSLB GL RFC ?GP MSRQGBC RFC @MBW. ,LMUGLE RFC 32, RFC N?PRG?J NPCQQSPC MD MVWECL GL RFC ?JTCMJG A?L @C A?JASJ?RCB:
alveolar P O = inspired P O (
2 2 2
alveolar P O 2) RQ
8GRF ?L 32 MD 0.8 ?LB ? P CO GL RFC ?JTCMJG MD 40 KK )E, RFC ?JTCMJ?P P O GQ COS?J RM:
alveolar P O
2
40 mm Hg = 150 mm Hg ( ) = mm Hg. 0.8

2
/MRGAC RF?R RFGQ NPCQQSPC GQ JCQQ RF?L RFC CVRCPL?J ?GP. 5FCPCDMPC, RFC MVWECL UGJJ DJMU DPMK RFC GLQNGPCB ?GP GL RFC JSLE ( P O = 150 KK )E) GLRM RFC @JMMBQRPC?K ( P O = 100 KK )E) (Figure 39.13). *L RFC JSLEQ, MVWECL BGDDSQCQ MSR MD RFC ?JTCMJG ?LB GLRM RFC A?NGJJ?PGCQ QSPPMSLBGLE RFC ?JTCMJG. 0VWECL (?@MSR 98 NCPACLR) @GLBQ PCTCPQG@JW RM RFC PCQNGP?RMPW NGEKCLR FCKMEJM@GL DMSLB GL PCB @JMMB ACJJQ (3#$Q). 3#$Q A?PPW MVWECL RM RFC RGQQSCQ UFCPC MVWECL BGQQMAG?RCQ DPMK RFC FCKMEJM@GL ?LB BGDDSQCQ GLRM RFC ACJJQ MD RFC RGQQSCQ. .MPC QNCAGDGA?JJW, ?JTCMJ?P P O GQ FGEFCP GL RFC ?JTCMJG ( P ALVO = 100 KK )E) RF?L @JMMB P O
2 2 2
(40 KK )E) GL RFC A?NGJJ?PGCQ. #CA?SQC RFGQ NPCQQSPC EP?BGCLR CVGQRQ,
MVWECL BGDDSQCQ BMUL GRQ NPCQQSPC EP?BGCLR, KMTGLE MSR MD RFC ?JTCMJG ?LB CLRCPGLE RFC @JMMB MD RFC A?NGJJ?PGCQ UFCPC 02 @GLBQ RM FCKMEJM@GL. "R RFC Q?KC RGKC, ?JTCMJ?P P CO GQ JMUCP P ALVO = 40 KK )E RF?L @JMMB P CO
2 2 2
= (45 KK )E). $02 BGDDSQCQ BMUL GRQ NPCQQSPC EP?BGCLR, KMTGLE MSR MD RFC
A?NGJJ?PGCQ ?LB CLRCPGLE RFC ?JTCMJG.
1146
#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
0VWECL ?LB A?P@ML BGMVGBC KMTC GLBCNCLBCLRJW MD C?AF MRFCP; RFCW BGDDSQC BMUL RFCGP MUL NPCQQSPC EP?BGCLRQ. "Q @JMMB JC?TCQ RFC JSLEQ RFPMSEF RFC NSJKML?PW TCGLQ, RFC venous PO = 100 KK )E, UFCPC?Q RFC venous PCO = 40 KK )E. "Q @JMMB CLRCPQ RFC QWQRCKGA A?NGJJ?PGCQ, RFC @JMMB UGJJ JMQC 2 MVWECL ?LB E?GL A?P@ML BGMVGBC @CA?SQC MD RFC NPCQQSPC BGDDCPCLAC MD RFC RGQQSCQ ?LB @JMMB. *L QWQRCKGA A?NGJJ?PGCQ, P O = 100 KK )E, @SR GL RFC RGQQSC ACJJQ, P O = 40 KK )E. 5FGQ NPCQQSPC EP?BGCLR BPGTCQ RFC BGDDSQGML MD MVWECL MSR MD RFC A?NGJJ?PGCQ ?LB GLRM RFC RGQQSC ACJJQ. "R RFC Q?KC RGKC, @JMMB P CO = 40 KK )E ?LB QWQRCKGA RGQQSC P CO = 45 KK )E. 5FC NPCQQSPC EP?BGCLR BPGTCQ $02 MSR MD RGQQSC ACJJQ 2 ?LB GLRM RFC A?NGJJ?PGCQ. 5FC @JMMB PCRSPLGLE RM RFC JSLEQ RFPMSEF RFC NSJKML?PW ?PRCPGCQ F?Q ? TCLMSQ P O = 40 KK )E ?LB ? P CO = 45 KK )E. 5FC @JMMB CLRCPQ RFC JSLE A?NGJJ?PGCQ UFCPC RFC NPMACQQ MD
2 2 2 2 2 2
CVAF?LEGLE E?QCQ @CRUCCL RFC A?NGJJ?PGCQ ?LB ?JTCMJG @CEGLQ ?E?GL (Figure 39.13).
!@>LI< 39.13 4EB M>OQF>I MOBPPROBP LC LUVDBK >KA @>O?LK AFLUFAB @E>KDB >P ?ILLA JLSBP QEOLRDE QEB ?LAV.
7EF@E LC QEB CLIILTFKD PQ>QBJBKQP FP C>IPB >. )K QEB QFPPRBP, P O AOLMP >P ?ILLA M>PPBP COLJ QEB >OQBOFBP QL QEB SBFKP, TEFIB P CO 2 2 FK@OB>PBP. ?. "ILLA QO>SBIP COLJ QEB IRKDP QL QEB EB>OQ QL ?LAV QFPPRBP, QEBK ?>@H QL QEB EB>OQ, QEBK QEB IRKDP. @. "ILLA QO>SBIP COLJ QEB IRKDP QL QEB EB>OQ QL ?LAV QFPPRBP, QEBK ?>@H QL QEB IRKDP, QEBK QEB EB>OQ. A. P O FP EFDEBO FK >FO QE>K FK QEB IRKDP.
2
*L QFMPR, RFC AF?LEC GL N?PRG?J NPCQQSPC DPMK RFC ?JTCMJG RM RFC A?NGJJ?PGCQ BPGTCQ RFC MVWECL GLRM RFC RGQQSCQ ?LB RFC A?P@ML BGMVGBC GLRM RFC @JMMB DPMK RFC RGQQSCQ. 5FC @JMMB GQ RFCL RP?LQNMPRCB RM RFC JSLEQ UFCPC BGDDCPCLACQ GL NPCQQSPC GL RFC ?JTCMJG PCQSJR GL RFC KMTCKCLR MD A?P@ML BGMVGBC MSR MD RFC @JMMB GLRM RFC JSLEQ, ?LB MVWECL GLRM RFC @JMMB.
#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
1147
8?RAF RFGQ video (http://openstaxcollege.org/l/spirometry) RM JC?PL FMU RM A?PPW MSR QNGPMKCRPW.
39.3 R BI<8K?@E>
#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C FMU RFC QRPSARSPCQ MD RFC JSLEQ ?LB RFMP?AGA A?TGRW AMLRPMJ RFC KCAF?LGAQ MD @PC?RFGLE z &VNJ?GL RFC GKNMPR?LAC MD AMKNJG?LAC ?LB PCQGQR?LAC GL RFC JSLEQ z %GQASQQ NPM@JCKQ RF?R K?W ?PGQC BSC RM ? 7/2 KGQK?RAF .?KK?JG?L JSLEQ ?PC JMA?RCB GL RFC RFMP?AGA A?TGRW UFCPC RFCW ?PC QSPPMSLBCB ?LB NPMRCARCB @W RFC PG@ A?EC, GLRCPAMQR?J KSQAJCQ, ?LB @MSLB @W RFC AFCQR U?JJ. 5FC @MRRMK MD RFC JSLEQ GQ AMLR?GLCB @W RFC BG?NFP?EK, ? QICJCR?J KSQAJC RF?R D?AGJGR?RCQ @PC?RFGLE. #PC?RFGLE PCOSGPCQ RFC AMMPBGL?RGML MD RFC JSLEQ, RFC AFCQR U?JJ, ?LB KMQR GKNMPR?LRJW, RFC BG?NFP?EK.
/PG<J F= BI<8K?@E>
"KNFG@G?LQ F?TC CTMJTCB KSJRGNJC U?WQ MD @PC?RFGLE. :MSLE ?KNFG@G?LQ, JGIC R?BNMJCQ, SQC EGJJQ RM @PC?RFC, ?LB RFCW BMLsR JC?TC RFC U?RCP. 4MKC ?KNFG@G?LQ PCR?GL EGJJQ DMP JGDC. "Q RFC R?BNMJC EPMUQ, RFC EGJJQ BGQ?NNC?P ?LB JSLEQ EPMU. 5FCQC JSLEQ ?PC NPGKGRGTC ?LB LMR ?Q CTMJTCB ?Q K?KK?JG?L JSLEQ. "BSJR ?KNFG@G?LQ ?PC J?AIGLE MP F?TC ? PCBSACB BG?NFP?EK, QM @PC?RFGLE TG? JSLEQ GQ DMPACB. 5FC MRFCP KC?LQ MD @PC?RFGLE DMP ?KNFG@G?LQ GQ BGDDSQGML ?APMQQ RFC QIGL. 5M ?GB RFGQ BGDDSQGML, ?KNFG@G?L QIGL KSQR PCK?GL KMGQR. #GPBQ D?AC ? SLGOSC AF?JJCLEC UGRF PCQNCAR RM @PC?RFGLE: 5FCW DJW. 'JWGLE AMLQSKCQ ? EPC?R ?KMSLR MD CLCPEW; RFCPCDMPC, @GPBQ PCOSGPC ? JMR MD MVWECL RM ?GB RFCGP KCR?@MJGA NPMACQQCQ. #GPBQ F?TC CTMJTCB ? PCQNGP?RMPW QWQRCK RF?R QSNNJGCQ RFCK UGRF RFC MVWECL LCCBCB RM CL?@JC DJWGLE. 4GKGJ?P RM K?KK?JQ, @GPBQ F?TC JSLEQ, UFGAF ?PC MPE?LQ QNCAG?JGXCB DMP E?Q CVAF?LEC. 0VWECL?RCB ?GP, R?ICL GL BSPGLE GLF?J?RGML, BGDDSQCQ ?APMQQ RFC QSPD?AC MD RFC JSLEQ GLRM RFC @JMMBQRPC?K, ?LB A?P@ML BGMVGBC BGDDSQCQ DPMK RFC @JMMB GLRM RFC JSLEQ ?LB CVNCJJCB BSPGLE CVF?J?RGML. 5FC BCR?GJQ MD @PC?RFGLE @CRUCCL @GPBQ ?LB K?KK?JQ BGDDCP QS@QR?LRG?JJW. *L ?BBGRGML RM JSLEQ, @GPBQ F?TC ?GP Q?AQ GLQGBC RFCGP @MBW. "GP DJMUQ GL MLC BGPCARGML DPMK RFC NMQRCPGMP ?GP Q?AQ RM RFC JSLEQ ?LB MSR MD RFC ?LRCPGMP ?GP Q?AQ. 5FC DJMU MD ?GP GQ GL RFC MNNMQGRC BGPCARGML DPMK @JMMB DJMU, ?LB E?Q CVAF?LEC R?ICQ NJ?AC KSAF KMPC CDDGAGCLRJW. 5FGQ RWNC MD @PC?RFGLE CL?@JCQ @GPBQ RM M@R?GL RFC PCOSGQGRC MVWECL, CTCL ?R FGEFCP ?JRGRSBCQ UFCPC RFC MVWECL AMLACLRP?RGML GQ JMU. 5FGQ BGPCARGML?JGRW MD ?GPDJMU PCOSGPCQ RUM AWAJCQ MD ?GP GLR?IC ?LB CVF?J?RGML RM AMKNJCRCJW ECR RFC ?GP MSR MD RFC JSLEQ.
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#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
AM@8E -<JG@I8K@FE
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
!@>LI< 39.14 (>) "FOAP E>SB > CILT-QEOLRDE OBPMFO>QLOV PVPQBJ FK TEF@E >FO CILTP RKFAFOB@QFLK>IIV COLJ QEB MLPQBOFLO P>@P FKQL QEB IRKDP, QEBK FKQL QEB >KQBOFLO >FO P>@P. 4EB >FO P>@P @LKKB@Q QL LMBKFKDP FK ELIILT ?LKBP. (?) $FKLP>ROP, COLJ TEF@E ?FOAP ABP@BKABA, E>SB PFJFI>O ELIILT ?LKBP >KA >OB ?BIFBSBA QL E>SB E>A > PFJFI>O OBPMFO>QLOV PVPQBJ. (@OBAFQ ?: JLAFCF@>QFLK LC TLOH ?V :FK> $BOBQPHV, .>QFLK>I 3@FBK@B &LRKA>QFLK)
-LPQ LC RP @LKPFABO QE>Q AFKLP>ROP >OB BUQFK@Q. (LTBSBO, JLABOK ?FOAP >OB ABP@BKA>KQP LC >SF>K AFKLP>ROP. 4EB OBPMFO>QLOV PVPQBJ LC JLABOK ?FOAP E>P ?BBK BSLISFKD CLO ERKAOBAP LC JFIIFLKP LC VB>OP. "JJ K?KK?JQ F?TC JSLEQ RF?R ?PC RFC K?GL MPE?LQ DMP @PC?RFGLE. -SLE A?N?AGRW F?Q CTMJTCB RM QSNNMPR RFC ?LGK?JsQ ?ARGTGRGCQ. %SPGLE GLF?J?RGML, RFC JSLEQ CVN?LB UGRF ?GP, ?LB MVWECL BGDDSQCQ ?APMQQ
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RFC JSLEsQ QSPD?AC ?LB CLRCPQ RFC @JMMBQRPC?K. %SPGLE CVF?J?RGML, RFC JSLEQ CVNCJ ?GP ?LB JSLE TMJSKC BCAPC?QCQ. *L RFC LCVR DCU QCARGMLQ, RFC NPMACQQ MD FSK?L @PC?RFGLE UGJJ @C CVNJ?GLCB.
/?< (<:?8E@:J F= #LD8E BI<8K?@E>

#MWJCsQ -?U GQ RFC E?Q J?U RF?R QR?RCQ RF?R GL ? AJMQCB QN?AC, NPCQQSPC ?LB TMJSKC ?PC GLTCPQCJW PCJ?RCB. "Q TMJSKC BCAPC?QCQ, NPCQQSPC GLAPC?QCQ ?LB TGAC TCPQ? (Figure 39.15). 5FC PCJ?RGMLQFGN @CRUCCL E?Q NPCQQSPC ?LB TMJSKC FCJNQ RM CVNJ?GL RFC KCAF?LGAQ MD @PC?RFGLE.
!@>LI< 39.15 4EFP DO>ME PELTP A>Q> COLJ "LVIBwP LOFDFK>I 1662 BUMBOFJBKQ, TEF@E PELTP QE>Q MOBPPROB >KA SLIRJB >OB FKSBOPBIV OBI>QBA. .L RKFQP >OB DFSBK >P "LVIB RPBA >O?FQO>OV RKFQP FK EFP BUMBOFJBKQP.
5FCPC GQ ?JU?WQ ? QJGEFRJW LCE?RGTC NPCQQSPC UGRFGL RFC RFMP?AGA A?TGRW, UFGAF ?GBQ GL ICCNGLE RFC ?GPU?WQ MD RFC JSLEQ MNCL. %SPGLE GLF?J?RGML, TMJSKC GLAPC?QCQ ?Q ? PCQSJR MD AMLRP?ARGML MD RFC BG?NFP?EK, ?LB NPCQQSPC BCAPC?QCQ (?AAMPBGLE RM #MWJCsQ -?U). 5FGQ BCAPC?QC MD NPCQQSPC GL RFC RFMP?AGA A?TGRW PCJ?RGTC RM RFC CLTGPMLKCLR K?ICQ RFC A?TGRW JCQQ RF?L RFC ?RKMQNFCPC (Figure 39.16a). #CA?SQC MD RFGQ BPMN GL NPCQQSPC, ?GP PSQFCQ GLRM RFC PCQNGP?RMPW N?QQ?ECQ. 5M GLAPC?QC RFC TMJSKC MD RFC JSLEQ, RFC AFCQR U?JJ CVN?LBQ. 5FGQ PCQSJRQ DPMK RFC AMLRP?ARGML MD RFC intercostal muscles, RFC KSQAJCQ RF?R ?PC AMLLCARCB RM RFC PG@ A?EC. -SLE TMJSKC CVN?LBQ @CA?SQC RFC BG?NFP?EK AMLRP?ARQ ?LB RFC GLRCPAMQR?JQ KSQAJCQ AMLRP?AR, RFSQ CVN?LBGLE RFC RFMP?AGA A?TGRW. 5FGQ GLAPC?QC GL RFC TMJSKC MD RFC RFMP?AGA A?TGRW JMUCPQ NPCQQSPC AMKN?PCB RM RFC ?RKMQNFCPC, QM ?GP PSQFCQ GLRM RFC JSLEQ, RFSQ GLAPC?QGLE GRQ TMJSKC. 5FC PCQSJRGLE GLAPC?QC GL TMJSKC GQ J?PECJW ?RRPG@SRCB RM ?L GLAPC?QC GL ?JTCMJ?P QN?AC, @CA?SQC RFC @PMLAFGMJCQ ?LB @PMLAFG ?PC QRGDD QRPSARSPCQ RF?R BM LMR AF?LEC GL QGXC.
!@>LI< 39.16 4EB IRKDP, @EBPQ T>II, >KA AF>MEO>DJ >OB >II FKSLISBA FK OBPMFO>QFLK, ?LQE (>) FKE>I>QFLK >KA (?) BUMFO>QFLK. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
5FC AFCQR U?JJ CVN?LBQ MSR ?LB ?U?W DPMK RFC JSLEQ. 5FC JSLEQ ?PC CJ?QRGA; RFCPCDMPC, UFCL ?GP DGJJQ RFC JSLEQ, RFC elastic recoil UGRFGL RFC RGQQSCQ MD RFC JSLE CVCPRQ NPCQQSPC @?AI RMU?PB RFC GLRCPGMP MD RFC JSLEQ. 5FCQC MSRU?PB ?LB GLU?PB DMPACQ AMKNCRC RM GLDJ?RC ?LB BCDJ?RC RFC JSLE UGRF CTCPW @PC?RF. 6NML CVF?J?RGML, RFC JSLEQ PCAMGJ RM DMPAC RFC ?GP MSR MD RFC JSLEQ, ?LB RFC GLRCPAMQR?J KSQAJCQ PCJ?V, PCRSPLGLE RFC AFCQR U?JJ @?AI RM GRQ MPGEGL?J NMQGRGML (Figure 39.16b). 5FC BG?NFP?EK ?JQM PCJ?VCQ ?LB KMTCQ FGEFCP GLRM RFC RFMP?AGA A?TGRW. 5FGQ GLAPC?QCQ RFC NPCQQSPC UGRFGL RFC RFMP?AGA A?TGRW PCJ?RGTC RM
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RFC CLTGPMLKCLR, ?LB ?GP PSQFCQ MSR MD RFC JSLEQ. 5FC KMTCKCLR MD ?GP MSR MD RFC JSLEQ GQ ? N?QQGTC CTCLR. /M KSQAJCQ ?PC AMLRP?ARGLE RM CVNCJ RFC ?GP. &?AF JSLE GQ QSPPMSLBCB @W ?L GLT?EGL?RCB Q?A. 5FC J?WCP MD RGQQSC RF?R AMTCPQ RFC JSLE ?LB BGNQ GLRM QN?ACQ GQ A?JJCB RFC TGQACP?J pleura. " QCAMLB J?WCP MD N?PGCR?J NJCSP? JGLCQ RFC GLRCPGMP MD RFC RFMP?V (Figure 39.17). 5FC QN?AC @CRUCCL RFCQC J?WCPQ, RFC intrapleural space, AMLR?GLQ ? QK?JJ ?KMSLR MD DJSGB RF?R NPMRCARQ RFC RGQQSC ?LB PCBSACQ RFC DPGARGML ECLCP?RCB DPMK PS@@GLE RFC RGQQSC J?WCPQ RMECRFCP ?Q RFC JSLEQ AMLRP?AR ?LB PCJ?V. Pleurisy PCQSJRQ UFCL RFCQC J?WCPQ MD RGQQSC @CAMKC GLDJ?KCB; GR GQ N?GLDSJ @CA?SQC RFC GLDJ?KK?RGML GLAPC?QCQ RFC NPCQQSPC UGRFGL RFC RFMP?AGA A?TGRW ?LB PCBSACQ RFC TMJSKC MD RFC JSLE.
!@>LI< 39.17 ! QFPPRB I>VBO @>IIBA MIBRO> PROOLRKAP QEB IRKD >KA FKQBOFLO LC QEB QELO>@F@ @>SFQV. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .#))
View (http://openstaxcollege.org/l/boyle_breathing) FMU #MWJCsQ -?U GQ PCJ?RCB RM @PC?RFGLE ?LB U?RAF this video (http://openstaxcollege.org/l/boyles_law) ML #MWJCsQ -?U.
/?< 2FIB F= BI<8K?@E>

5FC LSK@CP MD @PC?RFQ NCP KGLSRC GQ RFC respiratory rate. 0L ?TCP?EC, SLBCP LML-CVCPRGML AMLBGRGMLQ, RFC FSK?L PCQNGP?RMPW P?RC GQ 12p15 @PC?RFQ/KGLSRC. 5FC PCQNGP?RMPW P?RC AMLRPG@SRCQ RM RFC alveolar ventilation, MP FMU KSAF ?GP KMTCQ GLRM ?LB MSR MD RFC ?JTCMJG. "JTCMJ?P TCLRGJ?RGML NPCTCLRQ A?P@ML BGMVGBC @SGJBSN GL RFC ?JTCMJG. 5FCPC ?PC RUM U?WQ RM ICCN RFC ?JTCMJ?P TCLRGJ?RGML AMLQR?LR: GLAPC?QC RFC PCQNGP?RMPW P?RC UFGJC BCAPC?QGLE RFC RGB?J TMJSKC MD ?GP NCP @PC?RF (QF?JJMU @PC?RFGLE), MP BCAPC?QC RFC PCQNGP?RMPW P?RC UFGJC GLAPC?QGLE RFC RGB?J TMJSKC NCP @PC?RF. *L CGRFCP A?QC, RFC TCLRGJ?RGML PCK?GLQ RFC Q?KC, @SR RFC UMPI BMLC ?LB RWNC MD UMPI LCCBCB ?PC OSGRC BGDDCPCLR. #MRF RGB?J TMJSKC ?LB PCQNGP?RMPW P?RC ?PC AJMQCJW PCESJ?RCB UFCL MVWECL BCK?LB GLAPC?QCQ. 5FCPC ?PC RUM RWNCQ MD UMPI AMLBSARCB BSPGLE PCQNGP?RGML, DJMU-PCQGQRGTC ?LB CJ?QRGA UMPI. Flowresistive PCDCPQ RM RFC UMPI MD RFC ?JTCMJG ?LB RGQQSCQ GL RFC JSLE, UFCPC?Q elastic work PCDCPQ RM RFC UMPI MD RFC GLRCPAMQR?J KSQAJCQ, AFCQR U?JJ, ?LB BG?NFP?EK. *LAPC?QGLE RFC PCQNGP?RGML P?RC GLAPC?QCQ RFC DJMU-
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PCQGQRGTC UMPI MD RFC ?GPU?WQ ?LB BCAPC?QCQ RFC CJ?QRGA UMPI MD RFC KSQAJCQ. %CAPC?QGLE RFC PCQNGP?RMPW P?RC PCTCPQCQ RFC RWNC MD UMPI PCOSGPCB. !:7,')9'39 5FC ?GP-RGQQSC/U?RCP GLRCPD?AC MD RFC ?JTCMJG F?Q ? FGEF QSPD?AC RCLQGML. 5FGQ QSPD?AC RCLQGML GQ QGKGJ?P RM RFC QSPD?AC RCLQGML MD U?RCP ?R RFC JGOSGB-?GP GLRCPD?AC MD ? U?RCP BPMNJCR RF?R PCQSJRQ GL RFC @MLBGLE MD RFC U?RCP KMJCASJCQ RMECRFCP. Surfactant GQ ? AMKNJCV KGVRSPC MD NFMQNFMJGNGBQ ?LB JGNMNPMRCGLQ RF?R UMPIQ RM PCBSAC RFC QSPD?AC RCLQGML RF?R CVGQRQ @CRUCCL RFC ?JTCMJG RGQQSC ?LB RFC ?GP DMSLB UGRFGL RFC ?JTCMJG. #W JMUCPGLE RFC QSPD?AC RCLQGML MD RFC ?JTCMJ?P DJSGB, GR PCBSACQ RFC RCLBCLAW MD ?JTCMJG RM AMJJ?NQC. 4SPD?AR?LR UMPIQ JGIC ? BCRCPECLR RM PCBSAC RFC QSPD?AC RCLQGML ?LB ?JJMUQ DMP C?QGCP GLDJ?RGML MD RFC ?GPU?WQ. 8FCL ? @?JJMML GQ DGPQR GLDJ?RCB, GR R?ICQ ? J?PEC ?KMSLR MD CDDMPR RM QRPCRAF RFC NJ?QRGA ?LB QR?PR RM GLDJ?RC RFC @?JJMML. *D ? JGRRJC @GR MD BCRCPECLR U?Q ?NNJGCB RM RFC GLRCPGMP MD RFC @?JJMML, RFCL RFC ?KMSLR MD CDDMPR MP UMPI LCCBCB RM @CEGL RM GLDJ?RC RFC @?JJMML UMSJB BCAPC?QC, ?LB GR UMSJB @CAMKC KSAF C?QGCP RM QR?PR @JMUGLE SN RFC @?JJMML. 5FGQ Q?KC NPGLAGNJC ?NNJGCQ RM RFC ?GPU?WQ. " QK?JJ ?KMSLR MD QSPD?AR?LR RM RFC ?GPU?W RGQQSCQ PCBSACQ RFC CDDMPR MP UMPI LCCBCB RM GLDJ?RC RFMQC ?GPU?WQ. #?@GCQ @MPL NPCK?RSPCJW QMKCRGKCQ BM LMR NPMBSAC CLMSEF QSPD?AR?LR. "Q ? PCQSJR, RFCW QSDDCP DPMK respiratory distress syndrome, @CA?SQC GR PCOSGPCQ KMPC CDDMPR RM GLDJ?RC RFCGP JSLEQ. 4SPD?AR?LR GQ ?JQM GKNMPR?LR DMP NPCTCLRGLE AMJJ?NQC MD QK?JJ ?JTCMJG PCJ?RGTC RM J?PEC ?JTCMJG. L:3- +8/89'3)+ '3* C4251/'3)+ 1SJKML?PW BGQC?QCQ PCBSAC RFC P?RC MD E?Q CVAF?LEC GLRM ?LB MSR MD RFC JSLEQ. 5UM K?GL A?SQCQ MD BCAPC?QCB E?Q CVAF?LEC ?PC compliance (FMU CJ?QRGA RFC JSLE GQ) ?LB resistance (FMU KSAF M@QRPSARGML CVGQRQ GL RFC ?GPU?WQ). " AF?LEC GL CGRFCP A?L BP?K?RGA?JJW ?JRCP @PC?RFGLE ?LB RFC ?@GJGRW RM R?IC GL MVWECL ?LB PCJC?QC A?P@ML BGMVGBC. &V?KNJCQ MD restrictive diseases ?PC PCQNGP?RMPW BGQRPCQQ QWLBPMKC ?LB NSJKML?PW DG@PMQGQ. *L @MRF BGQC?QCQ, RFC ?GPU?WQ ?PC JCQQ AMKNJG?LR ?LB RFCW ?PC QRGDD MP DG@PMRGA. 5FCPC GQ ? BCAPC?QC GL AMKNJG?LAC @CA?SQC RFC JSLE RGQQSC A?LLMR @CLB ?LB KMTC. *L RFCQC RWNCQ MD PCQRPGARGTC BGQC?QCQ, RFC GLRP?NJCSP?J NPCQQSPC GQ KMPC NMQGRGTC ?LB RFC ?GPU?WQ AMJJ?NQC SNML CVF?J?RGML, UFGAF RP?NQ ?GP GL RFC JSLEQ. 'MPACB MP functional vital capacity (FVC), UFGAF GQ RFC ?KMSLR MD ?GP RF?R A?L @C DMPAG@JW CVF?JCB ?DRCP R?IGLE RFC BCCNCQR @PC?RF NMQQG@JC, GQ KSAF JMUCP RF?L GL LMPK?J N?RGCLRQ, ?LB RFC RGKC GR R?ICQ RM CVF?JC KMQR MD RFC ?GP GQ EPC?RJW NPMJMLECB (Figure 39.18). " N?RGCLR QSDDCPGLE DPMK RFCQC BGQC?QCQ A?LLMR CVF?JC RFC LMPK?J ?KMSLR MD ?GP. Obstructive diseases ?LB AMLBGRGMLQ GLAJSBC CKNFWQCK?, ?QRFK?, ?LB NSJKML?PW CBCK?. *L CKNFWQCK?, UFGAF KMQRJW ?PGQCQ DPMK QKMIGLE RM@?AAM, RFC U?JJQ MD RFC ?JTCMJG ?PC BCQRPMWCB, BCAPC?QGLE RFC QSPD?AC ?PC? DMP E?Q CVAF?LEC. 5FC MTCP?JJ AMKNJG?LAC MD RFC JSLEQ GQ GLAPC?QCB, @CA?SQC ?Q RFC ?JTCMJ?P U?JJQ ?PC B?K?ECB, JSLE CJ?QRGA PCAMGJ BCAPC?QCQ BSC RM ? JMQQ MD CJ?QRGA DG@CPQ, ?LB KMPC ?GP GQ RP?NNCB GL RFC JSLEQ ?R RFC CLB MD CVF?J?RGML. "QRFK? GQ ? BGQC?QC GL UFGAF GLDJ?KK?RGML GQ RPGEECPCB @W CLTGPMLKCLR?J D?ARMPQ. *LDJ?KK?RGML M@QRPSARQ RFC ?GPU?WQ. 5FC M@QRPSARGML K?W @C BSC RM CBCK? (DJSGB ?AASKSJ?RGML), QKMMRF KSQAJC QN?QKQ GL RFC U?JJQ MD RFC @PMLAFGMJCQ, GLAPC?QCB KSASQ QCAPCRGML, B?K?EC RM RFC CNGRFCJG? MD RFC ?GPU?WQ, MP ? AMK@GL?RGML MD RFCQC CTCLRQ. 5FMQC UGRF ?QRFK? MP CBCK? CVNCPGCLAC GLAPC?QCB MAAJSQGML DPMK GLAPC?QCB GLDJ?KK?RGML MD RFC ?GPU?WQ. 5FGQ RCLBQ RM @JMAI RFC ?GPU?WQ, NPCTCLRGLE RFC NPMNCP KMTCKCLR MD E?QCQ (Figure 39.18). 5FMQC UGRF M@QRPSARGTC BGQC?QCQ F?TC J?PEC TMJSKCQ MD ?GP RP?NNCB ?DRCP CVF?J?RGML ?LB @PC?RFC ?R ? TCPW FGEF JSLE TMJSKC RM AMKNCLQ?RC DMP RFC J?AI MD ?GPU?W PCAPSGRKCLR.
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!@>LI< 39.18 4EB O>QFL LC &%61 (QEB >JLRKQ LC >FO QE>Q @>K ?B CLO@F?IV BUE>IBA FK LKB PB@LKA >CQBO Q>HFKD > ABBM ?OB>QE) QL &6# (QEB QLQ>I >JLRKQ LC >FO QE>Q @>K ?B CLO@F?IV BUE>IBA) @>K ?B RPBA QL AF>DKLPB TEBQEBO > MBOPLK E>P OBPQOF@QFSB LO L?PQOR@QFSB IRKD AFPB>PB. )K OBPQOF@QFSB IRKD AFPB>PB, &6# FP OBAR@BA ?RQ >FOT>VP >OB KLQ L?PQOR@QBA, PL QEB MBOPLK FP >?IB QL BUMBI >FO OB>PLK>?IV C>PQ. )K L?PQOR@QFSB IRKD AFPB>PB, >FOT>V L?PQOR@QFLK OBPRIQP FK PILT BUE>I>QFLK >P TBII >P OBAR@BA &6#. 4ERP, QEB &%61/&6# O>QFL FP ILTBO FK MBOPLKP TFQE L?PQOR@QFSB IRKD AFPB>PB (IBPP QE>K 69 MBO@BKQ) QE>K FK MBOPLKP TFQE OBPQOF@QFSB AFPB>PB (88 QL 90 MBO@BKQ).
D<8; .G8:<: 1/, (@JD8K:?

1SJKML?PW AGPASJ?RGML NPCQQSPC GQ TCPW JMU AMKN?PCB RM RF?R MD RFC QWQRCKGA AGPASJ?RGML. *R GQ ?JQM GLBCNCLBCLR MD A?PBG?A MSRNSR. 5FGQ GQ @CA?SQC MD ? NFCLMKCLML A?JJCB recruitment, UFGAF GQ RFC NPMACQQ MD MNCLGLE ?GPU?WQ RF?R LMPK?JJW PCK?GL AJMQCB UFCL A?PBG?A MSRNSR GLAPC?QCQ. "Q A?PBG?A MSRNSR GLAPC?QCQ, RFC LSK@CP MD A?NGJJ?PGCQ ?LB ?PRCPGCQ RF?R ?PC NCPDSQCB (DGJJCB UGRF @JMMB) GLAPC?QCQ. 5FCQC A?NGJJ?PGCQ ?LB ?PRCPGCQ ?PC LMR ?JU?WQ GL SQC @SR ?PC PC?BW GD LCCBCB. "R RGKCQ, FMUCTCP, RFCPC GQ ? KGQK?RAF @CRUCCL RFC ?KMSLR MD ?GP (TCLRGJ?RGML, 7) ?LB RFC ?KMSLR MD @JMMB (NCPDSQGML, 2) GL RFC JSLEQ. 5FGQ GQ PCDCPPCB RM ?Q ventilation/perfusion (V/Q) mismatch. 5FCPC ?PC RUM RWNCQ MD 7/2 KGQK?RAF. #MRF NPMBSAC dead space, PCEGMLQ MD @PMICL BMUL MP @JMAICB JSLE RGQQSC. %C?B QN?ACQ A?L QCTCPCJW GKN?AR @PC?RFGLE, @CA?SQC RFCW PCBSAC RFC QSPD?AC ?PC? ?T?GJ?@JC DMP E?Q BGDDSQGML. "Q ? PCQSJR, RFC ?KMSLR MD MVWECL GL RFC @JMMB BCAPC?QCQ, UFCPC?Q RFC A?P@ML BGMVGBC JCTCJ GLAPC?QCQ. %C?B QN?AC GQ APC?RCB UFCL LM TCLRGJ?RGML ?LB/MP NCPDSQGML R?ICQ NJ?AC. Anatomical dead space MP ?L?RMKGA?J QFSLR, ?PGQCQ DPMK ?L ?L?RMKGA?J D?GJSPC, UFGJC physiological dead space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
#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
1153
7GQGR RFGQ site (http://openstaxcollege.org/l/breathing) RM TGCU RFC KCAF?LGAQ MD @PC?RFGLE.
39.4 R /I8EJGFIK F= "8J<J @E #LD8E BF;@CP !CL@;J

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C FMU MVWECL GQ @MSLB RM FCKMEJM@GL ?LB RP?LQNMPRCB RM @MBW RGQQSCQ z &VNJ?GL FMU A?P@ML BGMVGBC GQ RP?LQNMPRCB DPMK @MBW RGQQSCQ RM RFC JSLEQ 0LAC RFC MVWECL BGDDSQCQ ?APMQQ RFC ?JTCMJG, GR CLRCPQ RFC @JMMBQRPC?K ?LB GQ RP?LQNMPRCB RM RFC RGQQSCQ UFCPC GR GQ SLJM?BCB, ?LB A?P@ML BGMVGBC BGDDSQCQ MSR MD RFC @JMMB ?LB GLRM RFC ?JTCMJG RM @C CVNCJJCB DPMK RFC @MBW. "JRFMSEF E?Q CVAF?LEC GQ ? AMLRGLSMSQ NPMACQQ, RFC MVWECL ?LB A?P@ML BGMVGBC ?PC RP?LQNMPRCB @W BGDDCPCLR KCAF?LGQKQ.
/I8EJGFIK F= *OP><E @E K?< BCFF;

"JRFMSEF MVWECL BGQQMJTCQ GL @JMMB, MLJW ? QK?JJ ?KMSLR MD MVWECL GQ RP?LQNMPRCB RFGQ U?W. 0LJW 1.5 NCPACLR MD MVWECL GL RFC @JMMB GQ BGQQMJTCB BGPCARJW GLRM RFC @JMMB GRQCJD. .MQR MVWECLq98.5 NCPACLRqGQ @MSLB RM ? NPMRCGL A?JJCB FCKMEJM@GL ?LB A?PPGCB RM RFC RGQQSCQ. H+24-14(/3 Hemoglobin, MP )@, GQ ? NPMRCGL KMJCASJC DMSLB GL PCB @JMMB ACJJQ (CPWRFPMAWRCQ) K?BC MD DMSP QS@SLGRQ: RUM ?JNF? QS@SLGRQ ?LB RUM @CR? QS@SLGRQ (Figure 39.19). &?AF QS@SLGR QSPPMSLBQ ? ACLRP?J heme group RF?R AMLR?GLQ GPML ?LB @GLBQ MLC MVWECL KMJCASJC, ?JJMUGLE C?AF FCKMEJM@GL KMJCASJC RM @GLB DMSP MVWECL KMJCASJCQ. .MJCASJCQ UGRF KMPC MVWECL @MSLB RM RFC FCKC EPMSNQ ?PC @PGEFRCP PCB. "Q ? PCQSJR, MVWECL?RCB ?PRCPG?J @JMMB UFCPC RFC )@ GQ A?PPWGLE DMSP MVWECL KMJCASJCQ GQ @PGEFR PCB, UFGJC TCLMSQ @JMMB RF?R GQ BCMVWECL?RCB GQ B?PICP PCB.
!@>LI< 39.19 4EB MOLQBFK FKPFAB (>) OBA ?ILLA @BIIP QE>Q @>OOFBP LUVDBK QL @BIIP >KA @>O?LK AFLUFAB QL QEB IRKDP FP (?) EBJLDIL?FK. (BJLDIL?FK FP J>AB RM LC CLRO PVJJBQOF@>I PR?RKFQP >KA CLRO EBJB DOLRMP. )OLK >PPL@F>QBA TFQE QEB EBJB ?FKAP LUVDBK. )Q FP QEB FOLK FK EBJLDIL?FK QE>Q DFSBP ?ILLA FQP OBA @LILO.
*R GQ C?QGCP RM @GLB ? QCAMLB ?LB RFGPB MVWECL KMJCASJC RM )@ RF?L RFC DGPQR KMJCASJC. 5FGQ GQ @CA?SQC RFC FCKMEJM@GL KMJCASJC AF?LECQ GRQ QF?NC, MP AMLDMPK?RGML, ?Q MVWECL @GLBQ. 5FC DMSPRF MVWECL GQ RFCL KMPC BGDDGASJR RM @GLB. 5FC @GLBGLE MD MVWECL RM FCKMEJM@GL A?L @C NJMRRCB ?Q ? DSLARGML MD RFC N?PRG?J NPCQQSPC MD MVWECL GL RFC @JMMB (V-?VGQ) TCPQSQ RFC PCJ?RGTC )@-MVWECL Q?RSP?RGML (W-?VGQ). 5FC PCQSJRGLE
1154
#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
EP?NFq?L oxygen dissociation curveqGQ QGEKMGB?J, MP 4-QF?NCB (Figure 39.20). "Q RFC N?PRG?J NPCQQSPC MD MVWECL GLAPC?QCQ, RFC FCKMEJM@GL @CAMKCQ GLAPC?QGLEJW Q?RSP?RCB UGRF MVWECL.
!@>LI< 39.20 4EB LUVDBK AFPPL@F>QFLK @ROSB ABJLKPQO>QBP QE>Q, >P QEB M>OQF>I MOBPPROB LC LUVDBK FK@OB>PBP, JLOB LUVDBK ?FKAP EBJLDIL?FK. (LTBSBO, QEB >CCFKFQV LC EBJLDIL?FK CLO LUVDBK J>V PEFCQ QL QEB IBCQ LO QEB OFDEQ ABMBKAFKD LK BKSFOLKJBKQ>I @LKAFQFLKP.
4EB HFAKBVP >OB OBPMLKPF?IB CLO OBJLSFKD BU@BPP (+ FLKP COLJ QEB ?ILLA. )C QEB HFAKBVP C>FI, TE>Q TLRIA E>MMBK QL ?ILLA M( >KA QL EBJLDIL?FK >CCFKFQV CLO LUVDBK F')9478 ".'9 A,,+)9 O=>-+3 B/3*/35FC oxygen-carrying capacity MD FCKMEJM@GL BCRCPKGLCQ FMU KSAF MVWECL GQ A?PPGCB GL RFC @JMMB. *L ?BBGRGML RM P O , MRFCP CLTGPMLKCLR?J D?ARMPQ ?LB BGQC?QCQ A?L ?DDCAR MVWECL A?PPWGLE A?N?AGRW ?LB
2
BCJGTCPW. $?P@ML BGMVGBC JCTCJQ, @JMMB N), ?LB @MBW RCKNCP?RSPC ?DDCAR MVWECL-A?PPWGLE A?N?AGRW (Figure 39.20). 8FCL A?P@ML BGMVGBC GQ GL RFC @JMMB, GR PC?ARQ UGRF U?RCP RM DMPK @GA?P@ML?RC (HCO 3 ) ?LB FWBPMECL GMLQ ()+). "Q RFC JCTCJ MD A?P@ML BGMVGBC GL RFC @JMMB GLAPC?QCQ, KMPC )+ GQ NPMBSACB ?LB RFC N) BCAPC?QCQ. 5FGQ GLAPC?QC GL A?P@ML BGMVGBC ?LB QS@QCOSCLR BCAPC?QC GL N) PCBSAC RFC ?DDGLGRW MD FCKMEJM@GL DMP MVWECL. 5FC MVWECL BGQQMAG?RCQ DPMK RFC )@ KMJCASJC, QFGDRGLE RFC MVWECL BGQQMAG?RGML ASPTC RM RFC PGEFR. 5FCPCDMPC, KMPC MVWECL GQ LCCBCB RM PC?AF RFC Q?KC FCKMEJM@GL Q?RSP?RGML JCTCJ ?Q UFCL RFC N) U?Q FGEFCP. " QGKGJ?P QFGDR GL RFC ASPTC ?JQM PCQSJRQ DPMK ?L GLAPC?QC GL @MBW RCKNCP?RSPC. *LAPC?QCB RCKNCP?RSPC, QSAF ?Q DPMK GLAPC?QCB ?ARGTGRW MD QICJCR?J KSQAJC, A?SQCQ RFC ?DDGLGRW MD FCKMEJM@GL DMP MVWECL RM @C PCBSACB. %GQC?QCQ JGIC QGAIJC ACJJ ?LCKG? ?LB RF?J?QQCKG? BCAPC?QC RFC @JMMBsQ ?@GJGRW RM BCJGTCP MVWECL RM RGQQSCQ ?LB GRQ MVWECL-A?PPWGLE A?N?AGRW. *L sickle cell anemia, RFC QF?NC MD RFC PCB @JMMB ACJJ GQ APCQACLRQF?NCB, CJMLE?RCB, ?LB QRGDDCLCB, PCBSAGLE GRQ ?@GJGRW RM BCJGTCP MVWECL (Figure 39.21). *L RFGQ DMPK, PCB @JMMB ACJJQ A?LLMR N?QQ RFPMSEF RFC A?NGJJ?PGCQ. 5FGQ GQ N?GLDSJ UFCL GR MAASPQ. Thalassemia GQ ? P?PC ECLCRGA BGQC?QC A?SQCB @W ? BCDCAR GL CGRFCP RFC ?JNF? MP RFC @CR? QS@SLGR MD )@. 1?RGCLRQ UGRF RF?J?QQCKG? NPMBSAC ? FGEF LSK@CP MD PCB @JMMB ACJJQ, @SR RFCQC ACJJQ F?TC JMUCP-RF?L-LMPK?J JCTCJQ MD FCKMEJM@GL. 5FCPCDMPC, RFC MVWECL-A?PPWGLE A?N?AGRW GQ BGKGLGQFCB.
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/I8EJGFIK F= C8I9FE D@FO@;< @E K?< BCFF;

$?P@ML BGMVGBC KMJCASJCQ ?PC RP?LQNMPRCB GL RFC @JMMB DPMK @MBW RGQQSCQ RM RFC JSLEQ @W MLC MD RFPCC KCRFMBQ: BGQQMJSRGML BGPCARJW GLRM RFC @JMMB, @GLBGLE RM FCKMEJM@GL, MP A?PPGCB ?Q ? @GA?P@ML?RC GML. 4CTCP?J NPMNCPRGCQ MD A?P@ML BGMVGBC GL RFC @JMMB ?DDCAR GRQ RP?LQNMPR. 'GPQR, A?P@ML BGMVGBC GQ KMPC QMJS@JC GL @JMMB RF?L MVWECL. "@MSR 5 RM 7 NCPACLR MD ?JJ A?P@ML BGMVGBC GQ BGQQMJTCB GL RFC NJ?QK?. 4CAMLB, A?P@ML BGMVGBC A?L @GLB RM NJ?QK? NPMRCGLQ MP A?L CLRCP PCB @JMMB ACJJQ ?LB @GLB RM FCKMEJM@GL. 5FGQ DMPK RP?LQNMPRQ ?@MSR 10 NCPACLR MD RFC A?P@ML BGMVGBC. 8FCL A?P@ML BGMVGBC @GLBQ RM FCKMEJM@GL, ? KMJCASJC A?JJCB carbaminohemoglobin GQ DMPKCB. #GLBGLE MD A?P@ML BGMVGBC RM FCKMEJM@GL GQ PCTCPQG@JC. 5FCPCDMPC, UFCL GR PC?AFCQ RFC JSLEQ, RFC A?P@ML BGMVGBC A?L DPCCJW BGQQMAG?RC DPMK RFC FCKMEJM@GL ?LB @C CVNCJJCB DPMK RFC @MBW. 5FGPB, RFC K?HMPGRW MD A?P@ML BGMVGBC KMJCASJCQ (85 NCPACLR) ?PC A?PPGCB ?Q N?PR MD RFC bicarbonate buffer system. *L RFGQ QWQRCK, A?P@ML BGMVGBC BGDDSQCQ GLRM RFC PCB @JMMB ACJJQ. Carbonic anhydrase (CA) UGRFGL RFC PCB @JMMB ACJJQ OSGAIJW AMLTCPRQ RFC A?P@ML BGMVGBC GLRM A?P@MLGA ?AGB ()2$03). $?P@MLGA ?AGB GQ ?L SLQR?@JC GLRCPKCBG?RC KMJCASJC RF?R GKKCBG?RCJW BGQQMAG?RCQ GLRM bicarbonate ions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chloride shift. 8FCL RFC @JMMB PC?AFCQ RFC JSLEQ, RFC @GA?P@ML?RC GML GQ RP?LQNMPRCB @?AI GLRM RFC PCB @JMMB ACJJ GL CVAF?LEC DMP RFC AFJMPGBC GML. 5FC )+ GML BGQQMAG?RCQ DPMK RFC FCKMEJM@GL ?LB @GLBQ RM RFC @GA?P@ML?RC GML. 5FGQ NPMBSACQ RFC A?P@MLGA ?AGB GLRCPKCBG?RC, UFGAF GQ AMLTCPRCB @?AI GLRM A?P@ML BGMVGBC RFPMSEF RFC CLXWK?RGA ?ARGML MD $". 5FC A?P@ML BGMVGBC NPMBSACB GQ CVNCJJCB RFPMSEF RFC JSLEQ BSPGLE CVF?J?RGML.
CO 2 + H 2 O
H 2 CO 3 HCO 3 + H+ (carbonic acid) (bicarbonate)
5FC @CLCDGR MD RFC @GA?P@ML?RC @SDDCP QWQRCK GQ RF?R A?P@ML BGMVGBC GQ tQM?ICB SNu GLRM RFC @JMMB UGRF JGRRJC AF?LEC RM RFC N) MD RFC QWQRCK. 5FGQ GQ GKNMPR?LR @CA?SQC GR R?ICQ MLJW ? QK?JJ AF?LEC GL RFC MTCP?JJ N) MD RFC @MBW DMP QCTCPC GLHSPW MP BC?RF RM PCQSJR. 5FC NPCQCLAC MD RFGQ @GA?P@ML?RC @SDDCP QWQRCK ?JQM ?JJMUQ DMP NCMNJC RM RP?TCJ ?LB JGTC ?R FGEF ?JRGRSBCQ: 8FCL RFC N?PRG?J NPCQQSPC MD MVWECL ?LB A?P@ML BGMVGBC AF?LEC ?R FGEF ?JRGRSBCQ, RFC @GA?P@ML?RC @SDDCP QWQRCK ?BHSQRQ RM PCESJ?RC A?P@ML BGMVGBC UFGJC K?GLR?GLGLE RFC AMPPCAR N) GL RFC @MBW. C'7(43 M434=/*+ P4/843/38FGJC A?P@ML BGMVGBC A?L PC?BGJW ?QQMAG?RC ?LB BGQQMAG?RC DPMK FCKMEJM@GL, MRFCP KMJCASJCQ QSAF ?Q A?P@ML KMLMVGBC ($0) A?LLMR. $?P@ML KMLMVGBC F?Q ? EPC?RCP ?DDGLGRW DMP FCKMEJM@GL RF?L MVWECL. 5FCPCDMPC, UFCL A?P@ML KMLMVGBC GQ NPCQCLR, GR @GLBQ RM FCKMEJM@GL NPCDCPCLRG?JJW MTCP MVWECL. "Q ? PCQSJR, MVWECL A?LLMR @GLB RM FCKMEJM@GL, QM TCPW JGRRJC MVWECL GQ RP?LQNMPRCB RFPMSEF RFC @MBW (Figure 39.22). $?P@ML KMLMVGBC GQ ? AMJMPJCQQ, MBMPJCQQ E?Q ?LB GQ RFCPCDMPC BGDDGASJR RM BCRCAR. *R GQ NPMBSACB @W E?Q-NMUCPCB TCFGAJCQ ?LB RMMJQ. $?P@ML KMLMVGBC A?L A?SQC FC?B?AFCQ, AMLDSQGML, ?LB L?SQC?; JMLE-
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#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
RCPK CVNMQSPC A?L A?SQC @P?GL B?K?EC MP BC?RF. "BKGLGQRCPGLE 100 NCPACLR (NSPC) MVWECL GQ RFC SQS?J RPC?RKCLR DMP A?P@ML KMLMVGBC NMGQMLGLE. "BKGLGQRP?RGML MD NSPC MVWECL QNCCBQ SN RFC QCN?P?RGML MD A?P@ML KMLMVGBC DPMK FCKMEJM@GL.
!@>LI< 39.22 !P MBO@BKQ #/ FK@OB>PBP, QEB LUVDBK P>QRO>QFLK LC EBJLDIL?FK AB@OB>PBP.
#(!04%2 39 X 4(% 2%30)2!4/29 3934%-
1157
& 4 / -(.
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
+ 9@:8I9FE8K< (HCO 3 ) @FE GML APC?RCB UFCL A?P@MLGA ?AGB BGQQMAG?RCQ GLRM ) ?LB (HCO 3 )
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
1158
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
C#A+/ - .0((A-4
39.1 .PJK<DJ F= "8J O:?8E><
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
39.2 "8J O:?8E>< 8:IFJJ -<JG@I8KFIP .LI=8:<J

5FC JSLEQ A?L FMJB ? J?PEC TMJSKC MD ?GP, @SR RFCW ?PC LMR SQS?JJW DGJJCB RM K?VGK?J A?N?AGRW. -SLE TMJSKC KC?QSPCKCLRQ GLAJSBC RGB?J TMJSKC, CVNGP?RMPW PCQCPTC TMJSKC, GLQNGP?RMPW PCQCPTC TMJSKC, ?LB PCQGBS?J TMJSKC. 5FC QSK MD RFCQC COS?JQ RFC RMR?J JSLE A?N?AGRW. (?Q KMTCKCLR GLRM MP MSR MD RFC JSLEQ GQ BCNCLBCLR ML RFC NPCQQSPC MD RFC E?Q. "GP GQ ? KGVRSPC MD E?QCQ; RFCPCDMPC, RFC N?PRG?J NPCQQSPC MD C?AF E?Q A?L @C A?JASJ?RCB RM BCRCPKGLC FMU RFC E?Q UGJJ DJMU GL RFC JSLE. 5FC BGDDCPCLAC @CRUCCL RFC N?PRG?J NPCQQSPC MD RFC E?Q GL RFC ?GP BPGTCQ MVWECL GLRM RFC RGQQSCQ ?LB A?P@ML BGMVGBC MSR MD RFC @MBW.
39.3 BI<8K?@E>
5FC QRPSARSPC MD RFC JSLEQ ?LB RFMP?AGA A?TGRW AMLRPMJ RFC KCAF?LGAQ MD @PC?RFGLE. 6NML GLQNGP?RGML, RFC BG?NFP?EK AMLRP?ARQ ?LB JMUCPQ. 5FC GLRCPAMQR?J KSQAJCQ AMLRP?AR ?LB CVN?LB RFC AFCQR U?JJ MSRU?PB. 5FC GLRP?NJCSP?J NPCQQSPC BPMNQ, RFC JSLEQ CVN?LB, ?LB ?GP GQ BP?UL GLRM RFC ?GPU?WQ. 8FCL CVF?JGLE, RFC GLRCPAMQR?J KSQAJCQ ?LB BG?NFP?EK PCJ?V, PCRSPLGLE RFC GLRP?NJCSP?J NPCQQSPC @?AI RM RFC PCQRGLE QR?RC. 5FC JSLEQ PCAMGJ ?LB ?GPU?WQ AJMQC. 5FC ?GP N?QQGTCJW CVGRQ RFC JSLE. 5FCPC GQ FGEF QSPD?AC RCLQGML ?R RFC ?GP-?GPU?W GLRCPD?AC GL RFC JSLE. 4SPD?AR?LR, ? KGVRSPC MD NFMQNFMJGNGBQ ?LB JGNMNPMRCGLQ, ?ARQ JGIC ? BCRCPECLR GL RFC ?GPU?WQ RM PCBSAC QSPD?AC RCLQGML ?LB ?JJMU DMP MNCLGLE MD RFC ?JTCMJG. #PC?RFGLE ?LB E?Q CVAF?LEC ?PC @MRF ?JRCPCB @W AF?LECQ GL RFC AMKNJG?LAC ?LB PCQGQR?LAC MD RFC JSLE. *D RFC AMKNJG?LAC MD RFC JSLE BCAPC?QCQ, ?Q MAASPQ GL PCQRPGARGTC BGQC?QCQ JGIC DG@PMQGQ, RFC ?GPU?WQ QRGDDCL ?LB AMJJ?NQC SNML CVF?J?RGML. "GP @CAMKCQ RP?NNCB GL RFC JSLEQ, K?IGLE @PC?RFGLE KMPC BGDDGASJR. *D PCQGQR?LAC GLAPC?QCQ, ?Q F?NNCLQ UGRF ?QRFK? MP CKNFWQCK?, RFC ?GPU?WQ @CAMKC M@QRPSARCB, RP?NNGLE ?GP GL RFC JSLEQ ?LB A?SQGLE @PC?RFGLE RM @CAMKC BGDDGASJR. "JRCP?RGMLQ GL RFC TCLRGJ?RGML MD RFC
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?GPU?WQ MP NCPDSQGML MD RFC ?PRCPGCQ A?L ?DDCAR E?Q CVAF?LEC. 5FCQC AF?LECQ GL TCLRGJ?RGML ?LB NCPDSQGML, A?JJCB 7/2 KGQK?RAF, A?L ?PGQC DPMK ?L?RMKGA?J MP NFWQGMJMEGA?J AF?LECQ.
39.4 /I8EJGFIK F= "8J<J @E #LD8E BF;@CP !CL@;J

)CKMEJM@GL GQ ? NPMRCGL DMSLB GL PCB @JMMB ACJJQ RF?R GQ AMKNPGQCB MD RUM ?JNF? ?LB RUM @CR? QS@SLGRQ RF?R QSPPMSLB ?L GPML-AMLR?GLGLE FCKC EPMSN. 0VWECL PC?BGJW @GLBQ RFGQ FCKC EPMSN. 5FC ?@GJGRW MD MVWECL RM @GLB GLAPC?QCQ ?Q KMPC MVWECL KMJCASJCQ ?PC @MSLB RM FCKC. %GQC?QC QR?RCQ ?LB ?JRCPCB AMLBGRGMLQ GL RFC @MBW A?L ?DDCAR RFC @GLBGLE ?@GJGRW MD MVWECL, ?LB GLAPC?QC MP BCAPC?QC GRQ ?@GJGRW RM BGQQMAG?RC DPMK FCKMEJM@GL. $?P@ML BGMVGBC A?L @C RP?LQNMPRCB RFPMSEF RFC @JMMB TG? RFPCC KCRFMBQ. *R GQ BGQQMJTCB BGPCARJW GL RFC @JMMB, @MSLB RM NJ?QK? NPMRCGLQ MP FCKMEJM@GL, MP AMLTCPRCB GLRM @GA?P@ML?RC. 5FC K?HMPGRW MD A?P@ML BGMVGBC GQ RP?LQNMPRCB ?Q N?PR MD RFC @GA?P@ML?RC QWQRCK. $?P@ML BGMVGBC BGDDSQCQ GLRM PCB @JMMB ACJJQ. *LQGBC, A?P@MLGA ?LFWBP?QC AMLTCPRQ A?P@ML BGMVGBC GLRM A?P@MLGA ?AGB ()2$03), UFGAF GQ @JMMB ACJJQ, ?LB @GA?P@ML?RC GQ RP?LQNMPRCB MSR MD RFC PCB @JMMB ACJJQ GL CVAF?LEC DMP ? AFJMPGBC GML. 5FGQ GQ A?JJCB RFC AFJMPGBC QFGDR. #GA?P@ML?RC JC?TCQ RFC PCB @JMMB ACJJQ ?LB CLRCPQ RFC @JMMB NJ?QK?. *L RFC JSLEQ, @GA?P@ML?RC GQ RP?LQNMPRCB @?AI GLRM RFC PCB @JMMB ACJJQ GL CVAF?LEC DMP AFJMPGBC. 5FC ) + BGQQMAG?RCQ DPMK FCKMEJM@GL ?LB AMK@GLCQ UGRF @GA?P@ML?RC RM DMPK A?P@MLGA ?AGB UGRF RFC FCJN MD A?P@MLGA ?LFWBP?QC, UFGAF DSPRFCP A?R?JWXCQ RFC PC?ARGML RM AMLTCPR A?P@MLGA ?AGB @?AI GLRM A?P@ML BGMVGBC ?LB U?RCP. 5FC A?P@ML BGMVGBC GQ RFCL CVNCJJCB DPMK RFC JSLEQ.
+ + QS@QCOSCLRJW FWBPMJWXCB GLRM @GA?P@ML?RC (HCO 3 ) ?LB ) . 5FC ) GML @GLBQ RM FCKMEJM@GL GL PCB
A-/ C*)) C/$*) ,0 ./$*).

1. Figure 39.7 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR RFC K?KK?JG?L PCQNGP?RMPW QWQRCK GQ D?JQC >. 8FCL UC @PC?RFC GL, ?GP RP?TCJQ DPMK RFC NF?PWLV RM RFC RP?AFC?. ?. 5FC @PMLAFGMJCQ @P?LAF GLRM @PMLAFG. @. "JTCMJ?P BSARQ AMLLCAR RM ?JTCMJ?P Q?AQ. A. (?Q CVAF?LEC @CRUCCL RFC JSLE ?LB @JMMB R?ICQ NJ?AC GL RFC ?JTCMJSQ. 2. Figure 39.13 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ GQ D?JQC >. *L RFC RGQQSCQ, P O BPMNQ ?Q @JMMB
2
?. #JMMB RP?TCJQ DPMK RFC JSLEQ RM RFC FC?PR RM @MBW RGQQSCQ, RFCL @?AI RM RFC FC?PR, RFCL RFC JSLEQ. @. #JMMB RP?TCJQ DPMK RFC JSLEQ RM RFC FC?PR RM @MBW RGQQSCQ, RFCL @?AI RM RFC JSLEQ, RFCL RFC FC?PR. A. P O GQ FGEFCP GL ?GP RF?L GL RFC JSLEQ.
2
3. Figure 39.20 5FC IGBLCWQ ?PC PCQNMLQG@JC DMP PCKMTGLE CVACQQ )+ GMLQ DPMK RFC @JMMB. *D RFC IGBLCWQ D?GJ, UF?R UMSJB F?NNCL RM @JMMB N) ?LB RM FCKMEJM@GL ?DDGLGRW DMP MVWECL
N?QQCQ DPMK RFC ?PRCPGCQ RM RFC TCGLQ, UFGJC P CO GLAPC?QCQ.

2
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4. 5FC PCQNGP?RMPW QWQRCK >>>>>>>>. >. NPMTGBCQ @MBW RGQQSCQ UGRF MVWECL ?. NPMTGBCQ @MBW RGQQSCQ UGRF MVWECL ?LB A?P@ML BGMVGBC @. CQR?@JGQFCQ FMU K?LW @PC?RFQ ?PC R?ICL NCP KGLSRC A. NPMTGBCQ RFC @MBW UGRF A?P@ML BGMVGBC 5. "GP GQ U?PKCB ?LB FSKGBGDGCB GL RFC L?Q?J N?QQ?ECQ. 5FGQ FCJNQ RM >>>>>>>>. >. U?PB MDD GLDCARGML ?. BCAPC?QC QCLQGRGTGRW BSPGLE @PC?RFGLE @. NPCTCLR B?K?EC RM RFC JSLEQ A. ?JJ MD RFC ?@MTC 6. 8FGAF GQ RFC MPBCP MD ?GPDJMU BSPGLE GLF?J?RGML >. L?Q?J A?TGRW, RP?AFC?, J?PWLV, @PMLAFG, @PMLAFGMJCQ, ?JTCMJG ?. L?Q?J A?TGRW, J?PWLV, RP?AFC?, @PMLAFG, @PMLAFGMJCQ, ?JTCMJG @. L?Q?J A?TGRW, J?PWLV, RP?AFC?, @PMLAFGMJCQ, @PMLAFG, ?JTCMJG A. L?Q?J A?TGRW, RP?AFC?, J?PWLV, @PMLAFG, @PMLAFGMJCQ, ?JTCMJG 7. 5FC GLQNGP?RMPW PCQCPTC TMJSKC KC?QSPCQ RFC >>>>>>>>. >. ?KMSLR MD ?GP PCK?GLGLE GL RFC JSLE ?DRCP ? K?VGK?J CVF?J?RGML ?. ?KMSLR MD ?GP RF?R RFC JSLE FMJBQ @. ?KMSLR MD ?GP RFC A?L @C DSPRFCP CVF?JCB ?DRCP ? LMPK?J @PC?RF A. ?KMSLR MD ?GP RF?R A?L @C DSPRFCP GLF?JCB ?DRCP ? LMPK?J @PC?RF 8. 0D RFC DMJJMUGLE, UFGAF BMCQ LMR CVNJ?GL UFW RFC N?PRG?J NPCQQSPC MD MVWECL GQ JMUCP GL RFC JSLE RF?L GL RFC CVRCPL?J ?GP
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1161
>. "GP GL RFC JSLE GQ FSKGBGDGCB; RFCPCDMPC, U?RCP T?NMP NPCQQSPC ?JRCPQ RFC NPCQQSPC. ?. $?P@ML BGMVGBC KGVCQ UGRF MVWECL. @. 0VWECL GQ KMTCB GLRM RFC @JMMB ?LB GQ FC?BCB RM RFC RGQQSCQ. A. -SLEQ CVCPR ? NPCQQSPC ML RFC ?GP RM PCBSAC RFC MVWECL NPCQQSPC. 9. 5FC RMR?J JSLE A?N?AGRW GQ A?JASJ?RCB SQGLE UFGAF MD RFC DMJJMUGLE DMPKSJ?Q >. PCQGBS?J TMJSKC + RGB?J TMJSKC + GLQNGP?RMPW PCQCPTC TMJSKC ?. PCQGBS?J TMJSKC + CVNGP?RMPW PCQCPTC TMJSKC + GLQNGP?RMPW PCQCPTC TMJSKC @. CVNGP?RMPW PCQCPTC TMJSKC + RGB?J TMJSKC + GLQNGP?RMPW PCQCPTC TMJSKC A. PCQGBS?J TMJSKC + CVNGP?RMPW PCQCPTC TMJSKC + RGB?J TMJSKC + GLQNGP?RMPW PCQCPTC TMJSKC 10. )MU UMSJB N?P?JWQGQ MD RFC BG?NFP?EK ?JRCP GLQNGP?RGML >. *R UMSJB NPCTCLR AMLRP?ARGML MD RFC GLRCPAMQR?J KSQAJCQ. ?. *R UMSJB NPCTCLR GLF?J?RGML @CA?SQC RFC GLRP?NJCSP?J NPCQQSPC UMSJB LMR AF?LEC. @. *R UMSJB BCAPC?QC RFC GLRP?NJCSP?J NPCQQSPC ?LB ?JJMU KMPC ?GP RM CLRCP RFC JSLEQ. A. *R UMSJB QJMU CVNGP?RGML @CA?SQC RFC JSLE UMSJB LMR PCJ?V. 11. 3CQRPGARGTC ?GPU?W BGQC?QCQ >>>>>>>>. >. GLAPC?QC RFC AMKNJG?LAC MD RFC JSLE
?. BCAPC?QC RFC AMKNJG?LAC MD RFC JSLE @. GLAPC?QC RFC JSLE TMJSKC A. BCAPC?QC RFC UMPI MD @PC?RFGLE 12. "JTCMJ?P TCLRGJ?RGML PCK?GLQ AMLQR?LR UFCL >>>>>>>>. >. RFC PCQNGP?RMPW P?RC GQ GLAPC?QCB UFGJC RFC TMJSKC MD ?GP NCP @PC?RF GQ BCAPC?QCB ?. RFC PCQNGP?RMPW P?RC ?LB RFC TMJSKC MD ?GP NCP @PC?RF ?PC GLAPC?QCB @. RFC PCQNGP?RMPW P?RC GQ BCAPC?QCB UFGJC GLAPC?QGLE RFC TMJSKC NCP @PC?RF A. @MRF ? ?LB A 13. 8FGAF MD RFC DMJJMUGLE UGJJ /05 D?AGJGR?RC RFC RP?LQDCP MD MVWECL RM RGQQSCQ >. BCAPC?QCB @MBW RCKNCP?RSPC ?. BCAPC?QCB N) MD RFC @JMMB @. GLAPC?QCB A?P@ML BGMVGBC A. GLAPC?QCB CVCPAGQC 14. 5FC K?HMPGRW MD A?P@ML BGMVGBC GL RFC @JMMB GQ RP?LQNMPRCB @W >>>>>>>>. >. @GLBGLE RM FCKMEJM@GL ?. BGQQMJSRGML GL RFC @JMMB @. AMLTCPQGML RM @GA?P@ML?RC A. @GLBGLE RM NJ?QK? NPMRCGLQ 15. 5FC K?HMPGRW MD MVWECL GL RFC @JMMB GQ RP?LQNMPRCB @W >>>>>>>>. >. BGQQMJSRGML GL RFC @JMMB ?. @CGLE A?PPGCB ?Q @GA?P@ML?RC GMLQ @. @GLBGLE RM @JMMB NJ?QK? A. @GLBGLE RM FCKMEJM@GL
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16. %CQAPG@C RFC DSLARGML MD RFCQC RCPKQ ?LB BCQAPG@C UFCPC RFCW ?PC JMA?RCB: K?GL @PMLAFSQ, RP?AFC?, ?JTCMJG, ?LB ?AGLSQ. 17. )MU BMCQ RFC QRPSARSPC MD ?JTCMJG K?VGKGXC E?Q CVAF?LEC 18. 8F?R BMCQ '&71/'7$ KC?QSPC 8F?R D?ARMPQ K?W ?DDCAR '&71/'7$ 19. 8F?R GQ RFC PC?QML DMP F?TGLE PCQGBS?J TMJSKC GL RFC JSLE 20. )MU A?L ? BCAPC?QC GL RFC NCPACLR MD MVWECL GL RFC ?GP ?DDCAR RFC KMTCKCLR MD MVWECL GL RFC @MBW 21. *D ? N?RGCLR F?Q GLAPC?QCB PCQGQR?LAC GL FGQ MP FCP JSLEQ, FMU A?L RFGQ BCRCARCB @W ? BMARMP 8F?R BMCQ RFGQ KC?L 22. )MU UMSJB GLAPC?QCB ?GPU?W PCQGQR?LAC ?DDCAR GLRP?NJCSP?J NPCQQSPC BSPGLE GLF?J?RGML 23. &VNJ?GL FMU ? NSLARSPC RM RFC RFMP?AGA A?TGRW (DPMK ? ILGDC UMSLB, DMP GLQR?LAC) AMSJB ?JRCP RFC ?@GJGRW RM GLF?JC. 24. 8FCL QMKCMLC GQ QR?LBGLE, EP?TGRW QRPCRAFCQ RFC @MRRMK MD RFC JSLE BMUL RMU?PB RFC DJMMP RM ? EPC?RCP CVRCLR RF?L RFC RMN MD RFC JSLE. 8F?R GKNJGA?RGML AMSJB RFGQ F?TC ML RFC DJMU MD ?GP GL RFC JSLEQ 8FCPC BMCQ E?Q CVAF?LEC MAASP GL RFC JSLEQ 25. 8F?R UMSJB F?NNCL GD LM A?P@MLGA ?LFWBP?QC UCPC NPCQCLR GL PCB @JMMB ACJJQ 26. )MU BMCQ RFC ?BKGLGQRP?RGML MD 100 NCPACLR MVWECL Q?TC ? N?RGCLR DPMK A?P@ML KMLMVGBC NMGQMLGLE 8FW UMSJBLsR EGTGLE A?P@ML BGMVGBC UMPI
1162
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1164
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1165
@MRF QGBCQ, ?Q GJJSQRP?RCB GL Figure 40.3b. &VAF?LEC MD DJSGBQ GQ ?QQGQRCB @W RFC NSJQGLE MD RFC HCJJWDGQF @MBW.
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1166
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1167
MD RFC @MBW ?LB GRQ MPE?LQ GLAJSBGLE RFC @P?GL (QWQRCKGA AGPASJ?RGML). *L ?KNFG@G?LQ, E?Q CVAF?LEC ?JQM MAASPQ RFPMSEF RFC QIGL BSPGLE NSJKML?PW AGPASJ?RGML ?LB GQ PCDCPPCB RM ?Q pulmocutaneous circulation. "Q QFMUL GL Figure 40.4b, ?KNFG@G?LQ F?TC ? RFPCC-AF?K@CPCB FC?PR RF?R F?Q RUM ?RPG? ?LB MLC TCLRPGAJC P?RFCP RF?L RFC RUM-AF?K@CPCB FC?PR MD DGQF. 5FC RUM atria (QSNCPGMP FC?PR AF?K@CPQ) PCACGTC @JMMB DPMK RFC RUM BGDDCPCLR AGPASGRQ (RFC JSLEQ ?LB RFC QWQRCKQ), ?LB RFCL RFCPC GQ QMKC KGVGLE MD RFC @JMMB GL RFC FC?PRsQ ventricle (GLDCPGMP FC?PR AF?K@CP), UFGAF PCBSACQ RFC CDDGAGCLAW MD MVWECL?RGML. 5FC ?BT?LR?EC RM RFGQ ?PP?LECKCLR GQ RF?R FGEF NPCQQSPC GL RFC TCQQCJQ NSQFCQ @JMMB RM RFC JSLEQ ?LB @MBW. 5FC KGVGLE GQ KGRGE?RCB @W ? PGBEC UGRFGL RFC TCLRPGAJC RF?R BGTCPRQ MVWECL-PGAF @JMMB RFPMSEF RFC QWQRCKGA AGPASJ?RMPW QWQRCK ?LB BCMVWECL?RCB @JMMB RM RFC NSJKMASR?LCMSQ AGPASGR. 'MP RFGQ PC?QML, ?KNFG@G?LQ ?PC MDRCL BCQAPG@CB ?Q F?TGLE double circulation. .MQR PCNRGJCQ ?JQM F?TC ? RFPCC-AF?K@CPCB FC?PR QGKGJ?P RM RFC ?KNFG@G?L FC?PR RF?R BGPCARQ @JMMB RM RFC NSJKML?PW ?LB QWQRCKGA AGPASGRQ, ?Q QFMUL GL Figure 40.4c. 5FC TCLRPGAJC GQ BGTGBCB KMPC CDDCARGTCJW @W ? N?PRG?J QCNRSK, UFGAF PCQSJRQ GL JCQQ KGVGLE MD MVWECL?RCB ?LB BCMVWECL?RCB @JMMB. 4MKC PCNRGJCQ (?JJGE?RMPQ ?LB APMAMBGJCQ) ?PC RFC KMQR NPGKGRGTC ?LGK?JQ RM CVFG@GR ? DMSP-AF?K@CPCB FC?PR. $PMAMBGJG?LQ F?TC ? SLGOSC AGPASJ?RMPW KCAF?LGQK UFCPC RFC FC?PR QFSLRQ @JMMB DPMK RFC JSLEQ RMU?PB RFC QRMK?AF ?LB MRFCP MPE?LQ BSPGLE JMLE NCPGMBQ MD QS@KCPECLAC, DMP GLQR?LAC, UFGJC RFC ?LGK?J U?GRQ DMP NPCW MP QR?WQ SLBCPU?RCP U?GRGLE DMP NPCW RM PMR. 0LC ?B?NR?RGML GLAJSBCQ RUM K?GL ?PRCPGCQ RF?R JC?TC RFC Q?KC N?PR MD RFC FC?PR: MLC R?ICQ @JMMB RM RFC JSLEQ ?LB RFC MRFCP NPMTGBCQ ?L ?JRCPL?RC PMSRC RM RFC QRMK?AF ?LB MRFCP N?PRQ MD RFC @MBW. 5UM MRFCP ?B?NR?RGMLQ GLAJSBC ? FMJC GL RFC FC?PR @CRUCCL RFC RUM TCLRPGAJCQ, A?JJCB RFC DMP?KCL MD 1?LGXX?, UFGAF ?JJMUQ @JMMB RM KMTC DPMK MLC QGBC MD RFC FC?PR RM RFC MRFCP, ?LB QNCAG?JGXCB AMLLCARGTC RGQQSC RF?R QJMUQ RFC @JMMB DJMU RM RFC JSLEQ. 5MECRFCP RFCQC ?B?NR?RGMLQ F?TC K?BC APMAMBGJCQ ?LB ?JJGE?RMPQ MLC MD RFC KMQR CTMJSRGML?PGJW QSAACQQDSJ ?LGK?J EPMSNQ ML C?PRF. *L K?KK?JQ ?LB @GPBQ, RFC FC?PR GQ ?JQM BGTGBCB GLRM DMSP AF?K@CPQ: RUM ?RPG? ?LB RUM TCLRPGAJCQ, ?Q GJJSQRP?RCB GL Figure 40.4d. 5FC MVWECL?RCB @JMMB GQ QCN?P?RCB DPMK RFC BCMVWECL?RCB @JMMB, UFGAF GKNPMTCQ RFC CDDGAGCLAW MD BMS@JC AGPASJ?RGML ?LB GQ NPM@?@JW PCOSGPCB DMP RFC U?PK-@JMMBCB JGDCQRWJC MD K?KK?JQ ?LB @GPBQ. 5FC DMSP-AF?K@CPCB FC?PR MD @GPBQ ?LB K?KK?JQ CTMJTCB GLBCNCLBCLRJW DPMK ? RFPCC-AF?K@CPCB FC?PR. 5FC GLBCNCLBCLR CTMJSRGML MD RFC Q?KC MP ? QGKGJ?P @GMJMEGA?J RP?GR GQ PCDCPPCB RM ?Q AMLTCPECLR CTMJSRGML.
40.2 R CFDGFE<EKJ F= K?< BCFF;

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z -GQR RFC @?QGA AMKNMLCLRQ MD RFC @JMMB z $MKN?PC PCB ?LB UFGRC @JMMB ACJJQ z %CQAPG@C @JMMB NJ?QK? ?LB QCPSK )CKMEJM@GL GQ PCQNMLQG@JC DMP BGQRPG@SRGLE MVWECL, ?LB RM ? JCQQCP CVRCLR, A?P@ML BGMVGBC, RFPMSEFMSR RFC AGPASJ?RMPW QWQRCKQ MD FSK?LQ, TCPRC@P?RCQ, ?LB K?LW GLTCPRC@P?RCQ. 5FC @JMMB GQ KMPC RF?L RFC NPMRCGLQ, RFMSEF. #JMMB GQ ?ARS?JJW ? RCPK SQCB RM BCQAPG@C RFC JGOSGB RF?R KMTCQ RFPMSEF RFC TCQQCJQ ?LB GLAJSBCQ plasma (RFC JGOSGB NMPRGML, UFGAF AMLR?GLQ U?RCP, NPMRCGLQ, Q?JRQ, JGNGBQ, ?LB EJSAMQC) ?LB RFC ACJJQ (PCB ?LB UFGRC ACJJQ) ?LB ACJJ DP?EKCLRQ A?JJCB platelets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sQ CVRP?ACJJSJ?P DJSGB ?LB CGEFR NCPACLR MD UCGEFR.
/?< -FC< F= BCFF; @E K?< BF;P

#JMMB, JGIC RFC FSK?L @JMMB GJJSQRP?RCB GL Figure 40.5 GQ GKNMPR?LR DMP PCESJ?RGML MD RFC @MBWsQ QWQRCKQ ?LB FMKCMQR?QGQ. #JMMB FCJNQ K?GLR?GL FMKCMQR?QGQ @W QR?@GJGXGLE N), RCKNCP?RSPC, MQKMRGA NPCQQSPC, ?LB @W CJGKGL?RGLE CVACQQ FC?R. #JMMB QSNNMPRQ EPMURF @W BGQRPG@SRGLE LSRPGCLRQ ?LB FMPKMLCQ, ?LB @W PCKMTGLE U?QRC. #JMMB NJ?WQ ? NPMRCARGTC PMJC @W RP?LQNMPRGLE AJMRRGLE D?ARMPQ ?LB NJ?RCJCRQ RM NPCTCLR @JMMB JMQQ ?LB RP?LQNMPRGLE RFC BGQC?QC-DGEFRGLE ?ECLRQ MP white blood cells RM QGRCQ MD GLDCARGML.
1168
#(!04%2 40 X 4(% #)2#5,!4/29 3934%-
!@>LI< 40.5 4EB @BIIP >KA @BIIRI>O @LJMLKBKQP LC ERJ>K ?ILLA >OB PELTK. 2BA ?ILLA @BIIP ABIFSBO LUVDBK QL QEB @BIIP >KA OBJLSB @>O?LK AFLUFAB. 7EFQB ?ILLA @BIIPuFK@IRAFKD KBRQOLMEFIP, JLKL@VQBP, IVJMEL@VQBP, BLPFKLMEFIP, >KA ?>PLMEFIPu>OB FKSLISBA FK QEB FJJRKB OBPMLKPB. 0I>QBIBQP CLOJ @ILQP QE>Q MOBSBKQ ?ILLA ILPP >CQBO FKGROV.
-<; BCFF; C<CCJ

Red blood cells, MP CPWRFPMAWRCQ (CPWRFPM- = tPCBu; -AWRC = tACJJu), ?PC QNCAG?JGXCB ACJJQ RF?R AGPASJ?RC RFPMSEF RFC @MBW BCJGTCPGLE MVWECL RM ACJJQ; RFCW ?PC DMPKCB DPMK QRCK ACJJQ GL RFC @MLC K?PPMU. *L K?KK?JQ, PCB @JMMB ACJJQ ?PC QK?JJ @GAMLA?TC ACJJQ RF?R ?R K?RSPGRW BM LMR AMLR?GL ? LSAJCSQ MP KGRMAFMLBPG? ?LB ?PC MLJW 7p8 ]K GL QGXC. *L @GPBQ ?LB LML-?TG?L PCNRGJCQ, ? LSAJCSQ GQ QRGJJ K?GLR?GLCB GL PCB @JMMB ACJJQ. 5FC PCB AMJMPGLE MD @JMMB AMKCQ DPMK RFC GPML-AMLR?GLGLE NPMRCGL FCKMEJM@GL, GJJSQRP?RCB GL Figure 40.6a. 5FC NPGLAGN?J HM@ MD RFGQ NPMRCGL GQ RM A?PPW MVWECL, @SR GR ?JQM RP?LQNMPRQ A?P@ML BGMVGBC ?Q UCJJ. )CKMEJM@GL GQ N?AICB GLRM PCB @JMMB ACJJQ ?R ? P?RC MD ?@MSR 250 KGJJGML KMJCASJCQ MD FCKMEJM@GL NCP ACJJ. &?AF FCKMEJM@GL KMJCASJC @GLBQ DMSP MVWECL KMJCASJCQ QM RF?R C?AF PCB @JMMB ACJJ A?PPGCQ MLC @GJJGML KMJCASJCQ MD MVWECL. 5FCPC ?PC ?NNPMVGK?RCJW 25 RPGJJGML PCB @JMMB ACJJQ GL RFC DGTC JGRCPQ MD @JMMB GL RFC FSK?L @MBW, UFGAF AMSJB A?PPW SN RM 25 QCVRGJJGML (25 `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igure 40.6b GQ DMSLB GL KMJJSQIQ, APSQR?AC?LQ, ?LB QMKC MD RFC ?PRFPMNMBQ. $FJMPMAPSMPGL, ? EPCCL-AMJMPCB, GPML-AMLR?GLGLE NGEKCLR GQ DMSLB GL DMSP D?KGJGCQ MD NMJWAF?CRC RS@CUMPKQ. )CKCPWRFPGL, ? PCB, GPML-AMLR?GLGLE NPMRCGL GQ DMSLB GL QMKC NMJWAF?CRC UMPKQ ?LB ?LLCJGBQ ?LB GQ GJJSQRP?RCB GL Figure 40.6c. %CQNGRC RFC L?KC, FCKCPWRFPGL BMCQ LMR AMLR?GL ? FCKC EPMSN ?LB GRQ MVWECL-A?PPWGLE A?N?AGRW GQ NMMP AMKN?PCB RM FCKMEJM@GL.
#(!04%2 40 X 4(% #)2#5,!4/29 3934%-
1169
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
5FC QK?JJ QGXC ?LB J?PEC QSPD?AC ?PC? MD PCB @JMMB ACJJQ ?JJMUQ DMP P?NGB BGDDSQGML MD MVWECL ?LB A?P@ML BGMVGBC ?APMQQ RFC NJ?QK? KCK@P?LC. *L RFC JSLEQ, A?P@ML BGMVGBC GQ PCJC?QCB ?LB MVWECL GQ R?ICL GL @W RFC @JMMB. *L RFC RGQQSCQ, MVWECL GQ PCJC?QCB DPMK RFC @JMMB ?LB A?P@ML BGMVGBC GQ @MSLB DMP RP?LQNMPR @?AI RM RFC JSLEQ. 4RSBGCQ F?TC DMSLB RF?R FCKMEJM@GL ?JQM @GLBQ LGRPMSQ MVGBC (/0). /0 GQ ? T?QMBGJ?RMP RF?R PCJ?VCQ RFC @JMMB TCQQCJQ ?LB A?NGJJ?PGCQ ?LB K?W FCJN UGRF E?Q CVAF?LEC ?LB RFC N?QQ?EC MD PCB @JMMB ACJJQ RFPMSEF L?PPMU TCQQCJQ. /GRPMEJWACPGL, ? FC?PR KCBGA?RGML DMP ?LEGL? ?LB FC?PR ?RR?AIQ, GQ AMLTCPRCB RM /0 RM FCJN PCJ?V RFC @JMMB TCQQCJQ ?LB GLAPC?QC MVWECL DJMU RFPMSEF RFC @MBW. " AF?P?ARCPGQRGA MD PCB @JMMB ACJJQ GQ RFCGP EJWAMJGNGB ?LB EJWAMNPMRCGL AM?RGLE; RFCQC ?PC JGNGBQ ?LB NPMRCGLQ RF?R F?TC A?P@MFWBP?RC KMJCASJCQ ?RR?AFCB. *L FSK?LQ, RFC QSPD?AC EJWAMNPMRCGLQ ?LB EJWAMJGNGBQ ML PCB @JMMB ACJJQ T?PW @CRUCCL GLBGTGBS?JQ, NPMBSAGLE RFC BGDDCPCLR @JMMB RWNCQ, QSAF ?Q ", #, ?LB 0. 3CB @JMMB ACJJQ F?TC ?L ?TCP?EC JGDC QN?L MD 120 B?WQ, ?R UFGAF RGKC RFCW ?PC @PMICL BMUL ?LB PCAWAJCB GL RFC JGTCP ?LB QNJCCL @W NF?EMAWRGA K?APMNF?ECQ, ? RWNC MD UFGRC @JMMB ACJJ.
2?@K< BCFF; C<CCJ

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igure 40.7 ?PC RFC EP?LSJMAWRCQ, UFGAF GLAJSBC RFC LCSRPMNFGJQ, CMQGLMNFGJQ, ?LB @?QMNFGJQ, ?LB RFC ?EP?LSJMAWRCQ, UFGAF GLAJSBC RFC KMLMAWRCQ ?LB JWKNFMAWRCQ.
1170
#(!04%2 40 X 4(% #)2#5,!4/29 3934%-
!@>LI< 40.7 (>) 'O>KRIL@VQBPuFK@IRAFKD KBRQOLMEFIP, BLPFKLMEFIP >KA ?>PLMEFIPu>OB @E>O>@QBOFWBA ?V > IL?BA KR@IBRP >KA DO>KRI>O FK@IRPFLKP FK QEB @VQLMI>PJ. 'O>KRIL@VQBP >OB QVMF@>IIV CFOPQOBPMLKABOP AROFKD FKGROV LO FKCB@QFLK. (?) !DO>KRIL@VQBP FK@IRAB IVJMEL@VQBP >KA JLKL@VQBP. ,VJMEL@VQBP, FK@IRAFKD " >KA 4 @BIIP, >OB OBPMLKPF?IB CLO >A>MQFSB FJJRKB OBPMLKPB. -LKL@VQBP AFCCBOBKQF>QB FKQL J>@OLME>DBP >KA ABKAOFQF@ @BIIP, TEF@E FK QROK OBPMLKA QL FKCB@QFLK LO FKGROV.
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sQ MUL ECLCRGA K?AFGLCPW. "DRCP RFC )*7 TGPSQ PCNJGA?RCQ, GR GQ RP?LQKGRRCB BGPCARJW DPMK RFC GLDCARCB 5 ACJJ RM K?APMNF?ECQ. 5FC NPCQCLAC MD )*7 A?L PCK?GL SLPCAMELGXCB DMP ?L CVRCLQGTC NCPGMB MD RGKC @CDMPC DSJJ BGQC?QC QWKNRMKQ BCTCJMN.
+C8K<C<KJ 8E; CF8>LC8K@FE !8:KFIJ

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igure 40.8b. 5FC NJSE MP AJMR J?QRQ DMP ? LSK@CP MD B?WQ ?LB QRMNQ RFC JMQQ MD @JMMB. 1J?RCJCRQ ?PC DMPKCB DPMK RFC BGQGLRCEP?RGML MD J?PECP ACJJQ A?JJCB KCE?I?PWMAWRCQ, JGIC RF?R QFMUL GL Figure 40.8a. 'MP C?AF KCE?I?PWMAWRC, 2000p3000 NJ?RCJCRQ ?PC DMPKCB UGRF 150,000 RM 400,000 NJ?RCJCRQ NPCQCLR GL C?AF AS@GA KGJJGKCRCP MD @JMMB. &?AF NJ?RCJCR GQ BGQA QF?NCB ?LB 2p4 mK GL BG?KCRCP. 5FCW AMLR?GL K?LW QK?JJ TCQGAJCQ @SR BM LMR AMLR?GL ? LSAJCSQ.
#(!04%2 40 X 4(% #)2#5,!4/29 3934%-
1171
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+C8JD8 8E; .<ILD

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1174
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5FC ?SRMLMKMSQ @C?RGLE MD A?PBG?A KSQAJC ACJJQ GQ PCESJ?RCB @W RFC FC?PRsQ GLRCPL?J N?ACK?ICP RF?R SQCQ CJCARPGA?J QGEL?JQ RM RGKC RFC @C?RGLE MD RFC FC?PR. 5FC CJCARPGA?J QGEL?JQ ?LB KCAF?LGA?J ?ARGMLQ, GJJSQRP?RCB GL Figure 40.14, ?PC GLRGK?RCJW GLRCPRUGLCB. 5FC GLRCPL?J N?ACK?ICP QR?PRQ ?R RFC sinoatrial (SA) node, UFGAF GQ JMA?RCB LC?P RFC U?JJ MD RFC PGEFR ?RPGSK. &JCARPGA?J AF?PECQ QNMLR?LCMSQJW NSJQC DPMK RFC 4" LMBC A?SQGLE RFC RUM ?RPG? RM AMLRP?AR GL SLGQML. 5FC NSJQC PC?AFCQ ? QCAMLB LMBC, A?JJCB RFC ?RPGMTCLRPGASJ?P ("7) LMBC, @CRUCCL RFC PGEFR ?RPGSK ?LB PGEFR TCLRPGAJC UFCPC GR N?SQCQ DMP ?NNPMVGK?RCJW 0.1 QCAMLB @CDMPC QNPC?BGLE RM RFC U?JJQ MD RFC TCLRPGAJCQ. 'PMK RFC "7 LMBC, RFC CJCARPGA?J GKNSJQC CLRCPQ RFC @SLBJC MD )GQ, RFCL RM RFC JCDR ?LB PGEFR @SLBJC @P?LAFCQ CVRCLBGLE RFPMSEF RFC GLRCPTCLRPGASJ?P QCNRSK. 'GL?JJW, RFC 1SPIGLHC DG@CPQ AMLBSAR RFC GKNSJQC DPMK RFC ?NCV MD RFC FC?PR SN RFC TCLRPGASJ?P KWMA?PBGSK, ?LB RFCL RFC TCLRPGAJCQ AMLRP?AR. 5FGQ N?SQC ?JJMUQ RFC ?RPG? RM CKNRW AMKNJCRCJW GLRM RFC TCLRPGAJCQ @CDMPC RFC TCLRPGAJCQ NSKN MSR RFC @JMMB. 5FC CJCARPGA?J GKNSJQCQ GL RFC FC?PR NPMBSAC CJCARPGA?J ASPPCLRQ RF?R DJMU RFPMSEF RFC @MBW ?LB A?L @C KC?QSPCB ML RFC QIGL SQGLE CJCARPMBCQ. 5FGQ GLDMPK?RGML A?L @C M@QCPTCB ?Q ?L electrocardiogram (ECG)q? PCAMPBGLE MD RFC CJCARPGA?J GKNSJQCQ MD RFC A?PBG?A KSQAJC.
!@>LI< 40.14 4EB ?B>QFKD LC QEB EB>OQ FP OBDRI>QBA ?V >K BIB@QOF@>I FJMRIPB QE>Q @>RPBP QEB @E>O>@QBOFPQF@ OB>AFKD LC >K %#'. 4EB PFDK>I FP FKFQF>QBA >Q QEB PFKL>QOF>I S>ISB. 4EB PFDK>I QEBK (>) PMOB>AP QL QEB >QOF>, @>RPFKD QEBJ QL @LKQO>@Q. 4EB PFDK>I FP (?) ABI>VBA >Q QEB >QOFLSBKQOF@RI>O KLAB ?BCLOB FQ FP M>PPBA LK QL QEB (@) EB>OQ >MBU. 4EB ABI>V >IILTP QEB >QOF> QL OBI>U ?BCLOB QEB (A) SBKQOF@IBP @LKQO>@Q. 4EB CFK>I M>OQ LC QEB %#' @V@IB MOBM>OBP QEB EB>OQ CLO QEB KBUQ ?B>Q.
1178
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7GQGR this site (http://openstaxcollege.org/l/electric_heart) RM QCC RFC FC?PRsQ tN?ACK?ICPu GL ?ARGML.
AIK<I@<J, 1<@EJ, 8E; C8G@CC8I@<J

5FC @JMMB DPMK RFC FC?PR GQ A?PPGCB RFPMSEF RFC @MBW @W ? AMKNJCV LCRUMPI MD @JMMB TCQQCJQ (Figure 40.15). Arteries R?IC @JMMB ?U?W DPMK RFC FC?PR. 5FC K?GL ?PRCPW GQ RFC ?MPR? RF?R @P?LAFCQ GLRM K?HMP ?PRCPGCQ RF?R R?IC @JMMB RM BGDDCPCLR JGK@Q ?LB MPE?LQ. 5FCQC K?HMP ?PRCPGCQ GLAJSBC RFC A?PMRGB ?PRCPW RF?R R?ICQ @JMMB RM RFC @P?GL, RFC @P?AFG?J ?PRCPGCQ RF?R R?IC @JMMB RM RFC ?PKQ, ?LB RFC RFMP?AGA ?PRCPW RF?R R?ICQ @JMMB RM RFC RFMP?V ?LB RFCL GLRM RFC FCN?RGA, PCL?J, ?LB E?QRPGA ?PRCPGCQ DMP RFC JGTCP, IGBLCW, ?LB QRMK?AF, PCQNCARGTCJW. 5FC GJG?A ?PRCPW R?ICQ @JMMB RM RFC JMUCP JGK@Q. 5FC K?HMP ?PRCPGCQ BGTCPEC GLRM KGLMP ?PRCPGCQ, ?LB RFCL QK?JJCP TCQQCJQ A?JJCB arterioles, RM PC?AF KMPC BCCNJW GLRM RFC KSQAJCQ ?LB MPE?LQ MD RFC @MBW.
!@>LI< 40.15 4EB J>GLO ERJ>K >OQBOFBP >KA SBFKP >OB PELTK. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V ->OF>K> 2RFW 6FII>OB>I)
"PRCPGMJCQ BGTCPEC GLRM A?NGJJ?PW @CBQ. Capillary beds AMLR?GL ? J?PEC LSK@CP (10 RM 100) MD capillaries RF?R @P?LAF ?KMLE RFC ACJJQ ?LB RGQQSCQ MD RFC @MBW. $?NGJJ?PGCQ ?PC L?PPMU-BG?KCRCP RS@CQ RF?R A?L DGR PCB @JMMB ACJJQ RFPMSEF GL QGLEJC DGJC ?LB ?PC RFC QGRCQ DMP RFC CVAF?LEC MD LSRPGCLRQ, U?QRC, ?LB MVWECL UGRF RGQQSCQ ?R RFC ACJJSJ?P JCTCJ. 'JSGB ?JQM APMQQCQ GLRM RFC GLRCPQRGRG?J QN?AC DPMK RFC A?NGJJ?PGCQ. 5FC A?NGJJ?PGCQ AMLTCPEC ?E?GL GLRM venules RF?R AMLLCAR RM KGLMP TCGLQ RF?R DGL?JJW AMLLCAR RM K?HMP TCGLQ RF?R R?IC @JMMB FGEF GL A?P@ML BGMVGBC @?AI RM RFC FC?PR. Veins ?PC @JMMB TCQQCJQ RF?R @PGLE @JMMB @?AI RM RFC FC?PR. 5FC K?HMP TCGLQ BP?GL @JMMB DPMK RFC Q?KC MPE?LQ ?LB JGK@Q RF?R RFC K?HMP ?PRCPGCQ QSNNJW. 'JSGB GQ ?JQM @PMSEFR @?AI RM RFC FC?PR TG? RFC JWKNF?RGA QWQRCK.
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1179
5FC QRPSARSPC MD RFC BGDDCPCLR RWNCQ MD @JMMB TCQQCJQ PCDJCARQ RFCGP DSLARGML MP J?WCPQ. 5FCPC ?PC RFPCC BGQRGLAR J?WCPQ, MP RSLGAQ, RF?R DMPK RFC U?JJQ MD @JMMB TCQQCJQ (Figure 40.16). 5FC DGPQR RSLGA GQ ? QKMMRF, GLLCP JGLGLE MD CLBMRFCJG?J ACJJQ RF?R ?PC GL AMLR?AR UGRF RFC PCB @JMMB ACJJQ. 5FC CLBMRFCJG?J RSLGA GQ AMLRGLSMSQ UGRF RFC CLBMA?PBGSK MD RFC FC?PR. *L A?NGJJ?PGCQ, RFGQ QGLEJC J?WCP MD ACJJQ GQ RFC JMA?RGML MD BGDDSQGML MD MVWECL ?LB A?P@ML BGMVGBC @CRUCCL RFC CLBMRFCJG?J ACJJQ ?LB PCB @JMMB ACJJQ, ?Q UCJJ ?Q RFC CVAF?LEC QGRC TG? CLBMAWRMQGQ ?LB CVMAWRMQGQ. 5FC KMTCKCLR MD K?RCPG?JQ ?R RFC QGRC MD A?NGJJ?PGCQ GQ PCESJ?RCB @W vasoconstriction, L?PPMUGLE MD RFC @JMMB TCQQCJQ, ?LB vasodilation, UGBCLGLE MD RFC @JMMB TCQQCJQ; RFGQ GQ GKNMPR?LR GL RFC MTCP?JJ PCESJ?RGML MD @JMMB NPCQQSPC. 7CGLQ ?LB ?PRCPGCQ @MRF F?TC RUM DSPRFCP RSLGAQ RF?R QSPPMSLB RFC CLBMRFCJGSK: RFC KGBBJC RSLGA GQ AMKNMQCB MD QKMMRF KSQAJC ?LB RFC MSRCPKMQR J?WCP GQ AMLLCARGTC RGQQSC (AMJJ?ECL ?LB CJ?QRGA DG@CPQ). 5FC CJ?QRGA AMLLCARGTC RGQQSC QRPCRAFCQ ?LB QSNNMPRQ RFC @JMMB TCQQCJQ, ?LB RFC QKMMRF KSQAJC J?WCP FCJNQ PCESJ?RC @JMMB DJMU @W ?JRCPGLE T?QASJ?P PCQGQR?LAC RFPMSEF T?QMAMLQRPGARGML ?LB T?QMBGJ?RGML. 5FC ?PRCPGCQ F?TC RFGAICP QKMMRF KSQAJC ?LB AMLLCARGTC RGQQSC RF?L RFC TCGLQ RM ?AAMKKMB?RC RFC FGEFCP NPCQQSPC ?LB QNCCB MD DPCQFJW NSKNCB @JMMB. 5FC TCGLQ ?PC RFGLLCP U?JJCB ?Q RFC NPCQQSPC ?LB P?RC MD DJMU ?PC KSAF JMUCP. *L ?BBGRGML, TCGLQ ?PC QRPSARSP?JJW BGDDCPCLR RF?L ?PRCPGCQ GL RF?R TCGLQ F?TC T?JTCQ RM NPCTCLR RFC @?AIDJMU MD @JMMB. #CA?SQC TCGLQ F?TC RM UMPI ?E?GLQR EP?TGRW RM ECR @JMMB @?AI RM RFC FC?PR, AMLRP?ARGML MD QICJCR?J KSQAJC ?QQGQRQ UGRF RFC DJMU MD @JMMB @?AI RM RFC FC?PR.
!@>LI< 40.16 !OQBOFBP >KA SBFKP @LKPFPQ LC QEOBB I>VBOP: >K LRQBO QRKF@> BUQBOK>, > JFAAIB QRKF@> JBAF>, >KA >K FKKBO QRKF@> FKQFJ>. #>MFII>OFBP @LKPFPQ LC > PFKDIB I>VBO LC BMFQEBIF>I @BIIP, QEB QRKF@> FKQFJ>. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .#), .)()
40.4 R BCFF; !CFN 8E; BCFF; +I<JJLI< -<>LC8K@FE

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C RFC QWQRCK MD @JMMB DJMU RFPMSEF RFC @MBW z %CQAPG@C FMU @JMMB NPCQQSPC GQ PCESJ?RCB Blood pressure (BP) GQ RFC NPCQQSPC CVCPRCB @W @JMMB ML RFC U?JJQ MD ? @JMMB TCQQCJ RF?R FCJNQ RM NSQF @JMMB RFPMSEF RFC @MBW. 4WQRMJGA @JMMB NPCQQSPC KC?QSPCQ RFC ?KMSLR MD NPCQQSPC RF?R @JMMB CVCPRQ ML TCQQCJQ UFGJC RFC FC?PR GQ @C?RGLE. 5FC MNRGK?J QWQRMJGA @JMMB NPCQQSPC GQ 120 KK)E. %G?QRMJGA @JMMB NPCQQSPC KC?QSPCQ RFC NPCQQSPC GL RFC TCQQCJQ @CRUCCL FC?PR@C?RQ. 5FC MNRGK?J BG?QRMJGA @JMMB NPCQQSPC GQ 80 KK)E. .?LW D?ARMPQ A?L ?DDCAR @JMMB NPCQQSPC, QSAF ?Q FMPKMLCQ, QRPCQQ, CVCPAGQC, C?RGLE, QGRRGLE, ?LB QR?LBGLE. #JMMB DJMU RFPMSEF RFC @MBW GQ PCESJ?RCB @W RFC QGXC MD @JMMB TCQQCJQ, @W RFC ?ARGML MD QKMMRF KSQAJC, @W MLC-U?W T?JTCQ, ?LB @W RFC DJSGB NPCQQSPC MD RFC @JMMB GRQCJD.
#FN BCFF; !CFNJ /?IFL>? K?< BF;P

#JMMB GQ NSQFCB RFPMSEF RFC @MBW @W RFC ?ARGML MD RFC NSKNGLE FC?PR. 8GRF C?AF PFWRFKGA NSKN, @JMMB GQ NSQFCB SLBCP FGEF NPCQQSPC ?LB TCJMAGRW ?U?W DPMK RFC FC?PR, GLGRG?JJW ?JMLE RFC K?GL ?PRCPW, RFC ?MPR?.
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*L RFC ?MPR?, RFC @JMMB RP?TCJQ ?R 30 AK/QCA. "Q @JMMB KMTCQ GLRM RFC ?PRCPGCQ, ?PRCPGMJCQ, ?LB SJRGK?RCJW RM RFC A?NGJJ?PW @CBQ, RFC P?RC MD KMTCKCLR QJMUQ BP?K?RGA?JJW RM ?@MSR 0.026 AK/QCA, MLC-RFMSQ?LB RGKCQ QJMUCP RF?L RFC P?RC MD KMTCKCLR GL RFC ?MPR?. 8FGJC RFC BG?KCRCP MD C?AF GLBGTGBS?J ?PRCPGMJC ?LB A?NGJJ?PW GQ D?P L?PPMUCP RF?L RFC BG?KCRCP MD RFC ?MPR?, ?LB ?AAMPBGLE RM RFC J?U MD AMLRGLSGRW, DJSGB QFMSJB RP?TCJ D?QRCP RFPMSEF ? L?PPMUCP BG?KCRCP RS@C, RFC P?RC GQ ?ARS?JJW QJMUCP BSC RM RFC MTCP?JJ BG?KCRCP MD ?JJ RFC AMK@GLCB A?NGJJ?PGCQ @CGLE D?P EPC?RCP RF?L RFC BG?KCRCP MD RFC GLBGTGBS?J ?MPR?. 5FC QJMU P?RC MD RP?TCJ RFPMSEF RFC A?NGJJ?PW @CBQ, UFGAF PC?AF ?JKMQR CTCPW ACJJ GL RFC @MBW, ?QQGQRQ UGRF E?Q ?LB LSRPGCLR CVAF?LEC ?LB ?JQM NPMKMRCQ RFC BGDDSQGML MD DJSGB GLRM RFC GLRCPQRGRG?J QN?AC. "DRCP RFC @JMMB F?Q N?QQCB RFPMSEF RFC A?NGJJ?PW @CBQ RM RFC TCLSJCQ, TCGLQ, ?LB DGL?JJW RM RFC K?GL TCL?C A?T?C, RFC P?RC MD DJMU GLAPC?QCQ ?E?GL @SR GQ QRGJJ KSAF QJMUCP RF?L RFC GLGRG?J P?RC GL RFC ?MPR?. #JMMB NPGK?PGJW KMTCQ GL RFC TCGLQ @W RFC PFWRFKGA KMTCKCLR MD QKMMRF KSQAJC GL RFC TCQQCJ U?JJ ?LB @W RFC ?ARGML MD RFC QICJCR?J KSQAJC ?Q RFC @MBW KMTCQ. #CA?SQC KMQR TCGLQ KSQR KMTC @JMMB ?E?GLQR RFC NSJJ MD EP?TGRW, @JMMB GQ NPCTCLRCB DPMK DJMUGLE @?AIU?PB GL RFC TCGLQ @W MLC-U?W T?JTCQ. #CA?SQC QICJCR?J KSQAJC AMLRP?ARGML ?GBQ GL TCLMSQ @JMMB DJMU, GR GQ GKNMPR?LR RM ECR SN ?LB KMTC DPCOSCLRJW ?DRCP JMLE NCPGMBQ MD QGRRGLE QM RF?R @JMMB UGJJ LMR NMMJ GL RFC CVRPCKGRGCQ. #JMMB DJMU RFPMSEF RFC A?NGJJ?PW @CBQ GQ PCESJ?RCB BCNCLBGLE ML RFC @MBWs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igure 40.17. *D RFC QNFGLARCPQ ?PC MNCL, RFC @JMMB UGJJ DJMU GLRM RFC ?QQMAG?RCB @P?LAFCQ MD RFC A?NGJJ?PW @JMMB. *D ?JJ MD RFC QNFGLARCPQ ?PC AJMQCB, RFCL RFC @JMMB UGJJ DJMU BGPCARJW DPMK RFC ?PRCPGMJC RM RFC TCLSJC RFPMSEF RFC RFMPMSEFD?PC AF?LLCJ (QCC Figure 40.17). 5FCQC KSQAJCQ ?JJMU RFC @MBW RM NPCAGQCJW AMLRPMJ UFCL A?NGJJ?PW @CBQ PCACGTC @JMMB DJMU. "R ?LW EGTCL KMKCLR MLJW ?@MSR 5-10% MD MSP A?NGJJ?PW @CBQ ?ARS?JJW F?TC @JMMB DJMUGLE RFPMSEF RFCK.
!@>LI< 40.17 (>) 0OB@>MFII>OV PMEFK@QBOP >OB OFKDP LC PJLLQE JRP@IB QE>Q OBDRI>QB QEB CILT LC ?ILLA QEOLRDE @>MFII>OFBP; QEBV EBIM @LKQOLI QEB IL@>QFLK LC ?ILLA CILT QL TEBOB FQ FP KBBABA. (?) 6>ISBP FK QEB SBFKP MOBSBKQ ?ILLA COLJ JLSFKD ?>@HT>OA. (@OBAFQ >: JLAFCF@>QFLK LC TLOH ?V .#))
6>OF@LPB SBFKP >OB SBFKP QE>Q ?B@LJB BKI>ODBA ?B@>RPB QEB S>ISBP KL ILKDBO @ILPB MOLMBOIV, >IILTFKD ?ILLA QL CILT ?>@HT>OA. 6>OF@LPB SBFKP >OB LCQBK JLPQ MOLJFKBKQ LK QEB IBDP. 7EV AL VLR QEFKH QEFP FP QEB @>PB
7GQGR this site (http://openstaxcollege.org/l/circulation) RM QCC RFC AGPASJ?RMPW QWQRCKsQ @JMMB DJMU.
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1181
1PMRCGLQ ?LB MRFCP J?PEC QMJSRCQ A?LLMR JC?TC RFC A?NGJJ?PGCQ. 5FC JMQQ MD RFC U?RCPW NJ?QK? APC?RCQ ? FWNCPMQKMRGA QMJSRGML UGRFGL RFC A?NGJJ?PGCQ, CQNCAG?JJW LC?P RFC TCLSJCQ. 5FGQ A?SQCQ ?@MSR 85% MD RFC NJ?QK? RF?R JC?TCQ RFC A?NGJJ?PGCQ RM CTCLRS?JJW BGDDSQCQ @?AI GLRM RFC A?NGJJ?PGCQ LC?P RFC TCLSJCQ. 5FC PCK?GLGLE 15% MD @JMMB NJ?QK? BP?GLQ MSR DPMK RFC GLRCPQRGRG?J DJSGB GLRM LC?P@W JWKNF?RGA TCQQCJQ (Figure 40.18). 5FC DJSGB GL RFC JWKNF GQ QGKGJ?P GL AMKNMQGRGML RM RFC GLRCPQRGRG?J DJSGB. 5FC JWKNF DJSGB N?QQCQ RFPMSEF JWKNF LMBCQ @CDMPC GR PCRSPLQ RM RFC FC?PR TG? RFC TCL? A?T?. Lymph nodes ?PC QNCAG?JGXCB MPE?LQ RF?R DGJRCP RFC JWKNF @W NCPAMJ?RGML RFPMSEF ? K?XC MD AMLLCARGTC RGQQSC DGJJCB UGRF UFGRC @JMMB ACJJQ. 5FC UFGRC @JMMB ACJJQ PCKMTC GLDCARGMSQ ?ECLRQ, QSAF ?Q @?ARCPG? ?LB TGPSQCQ, RM AJC?L RFC JWKNF @CDMPC GR PCRSPLQ RM RFC @JMMBQRPC?K. "DRCP GR GQ AJC?LCB, RFC JWKNF PCRSPLQ RM RFC FC?PR @W RFC ?ARGML MD QKMMRF KSQAJC NSKNGLE, QICJCR?J KSQAJC ?ARGML, ?LB MLC-U?W T?JTCQ HMGLGLE RFC PCRSPLGLE @JMMB LC?P RFC HSLARGML MD RFC TCL?C A?T?C CLRCPGLE RFC PGEFR ?RPGSK MD RFC FC?PR.
!@>LI< 40.18 &IRFA COLJ QEB @>MFII>OFBP JLSBP FKQL QEB FKQBOPQFQF>I PM>@B >KA IVJME @>MFII>OFBP ?V AFCCRPFLK ALTK > MOBPPROB DO>AFBKQ >KA >IPL ?V LPJLPFP. /RQ LC 7,200 IFQBOP LC CIRFA MRJMBA ?V QEB >SBO>DB EB>OQ FK > A>V, LSBO 1,500 IFQBOP FP CFIQBOBA. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .#), .)()
1<IK<9I8K< D@M<IJ@KP @E BCFF; C@I:LC8K@FE

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1182
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BCFF; +I<JJLI<
5FC NPCQQSPC MD RFC @JMMB DJMU GL RFC @MBW GQ NPMBSACB @W RFC FWBPMQR?RGA NPCQQSPC MD RFC DJSGB (@JMMB) ?E?GLQR RFC U?JJQ MD RFC @JMMB TCQQCJQ. 'JSGB UGJJ KMTC DPMK ?PC?Q MD FGEF RM JMU FWBPMQR?RGA NPCQQSPCQ. *L RFC ?PRCPGCQ, RFC FWBPMQR?RGA NPCQQSPC LC?P RFC FC?PR GQ TCPW FGEF ?LB @JMMB DJMUQ RM RFC ?PRCPGMJCQ UFCPC RFC P?RC MD DJMU GQ QJMUCB @W RFC L?PPMU MNCLGLEQ MD RFC ?PRCPGMJCQ. %SPGLE QWQRMJC, UFCL LCU @JMMB GQ CLRCPGLE RFC ?PRCPGCQ, RFC ?PRCPW U?JJQ QRPCRAF RM ?AAMKKMB?RC RFC GLAPC?QC MD NPCQQSPC MD RFC CVRP? @JMMB; BSPGLE BG?QRMJC, RFC U?JJQ PCRSPL RM LMPK?J @CA?SQC MD RFCGP CJ?QRGA NPMNCPRGCQ. 5FC @JMMB NPCQQSPC MD RFC QWQRMJC NF?QC ?LB RFC BG?QRMJC NF?QC, EP?NFCB GL Figure 40.19, EGTCQ RFC RUM NPCQQSPC PC?BGLEQ DMP @JMMB NPCQQSPC. 'MP CV?KNJC, 120/80 GLBGA?RCQ ? PC?BGLE MD 120 KK )E BSPGLE RFC QWQRMJC ?LB 80 KK )E BSPGLE BG?QRMJC. 5FPMSEFMSR RFC A?PBG?A AWAJC, RFC @JMMB AMLRGLSCQ RM CKNRW GLRM RFC ?PRCPGMJCQ ?R ? PCJ?RGTCJW CTCL P?RC. 5FGQ PCQGQR?LAC RM @JMMB DJMU GQ A?JJCB peripheral resistance.
!@>LI< 40.19 "ILLA MOBPPROB FP OBI>QBA QL QEB ?ILLA SBIL@FQV FK QEB >OQBOFBP >KA >OQBOFLIBP. )K QEB @>MFII>OFBP >KA SBFKP, QEB ?ILLA MOBPPROB @LKQFKRBP QL AB@B>PB ?RQ SBIL@FQV FK@OB>PBP.
BCFF; +I<JJLI< -<>LC8K@FE

$?PBG?A MSRNSR GQ RFC TMJSKC MD @JMMB NSKNCB @W RFC FC?PR GL MLC KGLSRC. *R GQ A?JASJ?RCB @W KSJRGNJWGLE RFC LSK@CP MD FC?PR AMLRP?ARGMLQ RF?R MAASP NCP KGLSRC (FC?PR P?RC) RGKCQ RFC stroke volume (RFC TMJSKC MD @JMMB NSKNCB GLRM RFC ?MPR? NCP AMLRP?ARGML MD RFC JCDR TCLRPGAJC). 5FCPCDMPC, A?PBG?A MSRNSR A?L @C GLAPC?QCB @W GLAPC?QGLE FC?PR P?RC, ?Q UFCL CVCPAGQGLE. )MUCTCP, A?PBG?A MSRNSR A?L ?JQM @C GLAPC?QCB @W GLAPC?QGLE QRPMIC TMJSKC, QSAF ?Q GD RFC FC?PR AMLRP?ARQ UGRF EPC?RCP QRPCLERF. 4RPMIC TMJSKC A?L ?JQM @C GLAPC?QCB @W QNCCBGLE @JMMB AGPASJ?RGML RFPMSEF RFC @MBW QM RF?R KMPC @JMMB CLRCPQ RFC FC?PR @CRUCCL AMLRP?ARGMLQ. %SPGLE FC?TW CVCPRGML, RFC @JMMB TCQQCJQ PCJ?V ?LB GLAPC?QC GL BG?KCRCP, MDDQCRRGLE RFC GLAPC?QCB FC?PR P?RC ?LB CLQSPGLE ?BCOS?RC MVWECL?RCB @JMMB ECRQ RM RFC KSQAJCQ. 4RPCQQ RPGEECPQ ? BCAPC?QC GL RFC BG?KCRCP MD RFC @JMMB TCQQCJQ, AMLQCOSCLRJW GLAPC?QGLE @JMMB NPCQQSPC. 5FCQC AF?LECQ A?L ?JQM @C A?SQCB @W LCPTC QGEL?JQ MP FMPKMLCQ, ?LB CTCL QR?LBGLE SN MP JWGLE BMUL A?L F?TC ? EPC?R CDDCAR ML @JMMB NPCQQSPC.
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& 4 / -(.
8E>@E8 N?GL A?SQCB @W N?PRG?J @JMAI?EC MD RFC AMPML?PW ?PRCPGCQ @W RFC @SGJBSN MD NJ?OSC ?LB J?AI MD MVWECL RM RFC FC?PR KSQAJC 8FIK8 K?HMP ?PRCPW MD RFC @MBW RF?R R?ICQ @JMMB ?U?W DPMK RFC FC?PR 8IK<I@FC< QK?JJ TCQQCJ RF?R AMLLCARQ ?L ?PRCPW RM ? A?NGJJ?PW @CB 8IK<IP @JMMB TCQQCJ RF?R R?ICQ @JMMB ?U?W DPMK RFC FC?PR 8K?<IFJ:C<IFJ@J @SGJBSN MD D?RRW NJ?OSCQ GL RFC AMPML?PW ?PRCPGCQ GL RFC FC?PR 8KI@FM<EKI@:LC8I M8CM< MLC-U?W KCK@P?LMSQ DJ?N MD AMLLCARGTC RGQQSC @CRUCCL RFC ?RPGSK ?LB RFC TCLRPGAJC GL RFC PGEFR QGBC MD RFC FC?PR; ?JQM ILMUL ?Q RPGASQNGB T?JTC 8KI@LD (NJSP?J: ?RPG?) AF?K@CP MD RFC FC?PR RF?R PCACGTCQ @JMMB DPMK RFC TCGLQ ?LB QCLBQ @JMMB RM RFC TCLRPGAJCQ 9@:LJG@; M8CM< (?JQM, KGRP?J T?JTC; JCDR ?RPGMTCLRPGASJ?P T?JTC) MLC-U?W KCK@P?LMSQ DJ?N @CRUCCL RFC ?RPGSK ?LB RFC TCLRPGAJC GL RFC JCDR QGBC MD RFC FC?PR 9CFF; GI<JJLI< (B+) NPCQQSPC MD @JMMB GL RFC ?PRCPGCQ RF?R FCJNQ RM NSQF @JMMB RFPMSEF RFC @MBW :8G@CC8IP 9<; J?PEC LSK@CP MD A?NGJJ?PGCQ RF?R AMLTCPEC RM R?IC @JMMB RM ? N?PRGASJ?P MPE?L MP RGQQSC :8G@CC8IP QK?JJCQR @JMMB TCQQCJ RF?R ?JJMUQ RFC N?QQ?EC MD GLBGTGBS?J @JMMB ACJJQ ?LB RFC QGRC MD BGDDSQGML MD MVWECL ?LB LSRPGCLR CVAF?LEC :8I;@8: :P:C< DGJJGLE ?LB CKNRWGLE RFC FC?PR MD @JMMB @W CJCARPGA?J QGEL?JQ RF?R A?SQC RFC FC?PR KSQAJCQ RM AMLRP?AR ?LB PCJ?V :8I;@8: FLKGLK RFC TMJSKC MD @JMMB NSKNCB @W RFC FC?PR GL MLC KGLSRC ?Q ? NPMBSAR MD FC?PR P?RC KSJRGNJGCB @W QRPMIC TMJSKC :8I;@FDPF:PK< QNCAG?JGXCB FC?PR KSQAJC ACJJ RF?R GQ QRPG?RCB @SR AMLRP?ARQ GLTMJSLR?PGJW JGIC QKMMRF KSQAJC :CFJ<; :@I:LC8KFIP JPJK<D QWQRCK GL UFGAF RFC @JMMB GQ QCN?P?RCB DPMK RFC @MBGJW GLRCPQRGRG?J DJSGB ?LB AMLR?GLCB GL @JMMB TCQQCJQ :FIFE8IP 8IK<IP TCQQCJ RF?R QSNNJGCQ RFC FC?PR RGQQSC UGRF @JMMB :FIFE8IP M<@E TCQQCJ RF?R R?ICQ @JMMB ?U?W DPMK RFC FC?PR RGQQSC @?AI RM RFC AF?K@CPQ GL RFC FC?PR ;@8JKFC< PCJ?V?RGML NF?QC MD RFC A?PBG?A AWAJC UFCL RFC FC?PR GQ PCJ?VCB ?LB RFC TCLRPGAJCQ ?PC DGJJGLE UGRF @JMMB ;FL9C< :@I:LC8K@FE DJMU MD @JMMB GL RUM AGPASGRQ: RFC NSJKML?PW AGPASGR RFPMSEF RFC JSLEQ ?LB RFC QWQRCKGA AGPASGR RFPMSEF RFC MPE?LQ ?LB @MBW <C<:KIF:8I;@F>I8D ( C") PCAMPBGLE MD RFC CJCARPGA?J GKNSJQCQ MD RFC A?PBG?A KSQAJC <E;F:8I;@LD GLLCPKMQR J?WCP MD RGQQSC GL RFC FC?PR <G@:8I;@LD MSRCPKMQR RGQQSC J?WCP MD RFC FC?PR >@CC :@I:LC8K@FE AGPASJ?RMPW QWQRCK RF?R GQ QNCAGDGA RM ?LGK?JQ UGRF EGJJQ DMP E?Q CVAF?LEC; RFC @JMMB DJMUQ RFPMSEF RFC EGJJQ DMP MVWECL?RGML ?<DF:F<C A?TGRW GLRM UFGAF @JMMB GQ NSKNCB GL ?L MNCL AGPASJ?RMPW QWQRCK ?<DFCPDG? KGVRSPC MD @JMMB ?LB GLRCPQRGRG?J DJSGB RF?R GQ DMSLB GL GLQCARQ ?LB MRFCP ?PRFPMNMBQ ?Q UCJJ ?Q KMQR KMJJSQIQ @E=<I@FI M<E8 :8M8 BP?GLQ @JMMB DPMK RFC TCGLQ RF?R AMKC DPMK RFC JMUCP MPE?LQ ?LB RFC JCEQ
1184
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p8 mK) @GAMLA?TC ACJJ UGRFMSR KGRMAFMLBPG? (?LB GL K?KK?JQ UGRFMSR LSAJCG) RF?R GQ N?AICB UGRF FCKMEJM@GL, EGTGLE RFC ACJJ GRQ PCB AMJMP; RP?LQNMPRQ MVWECL RFPMSEF RFC @MBW J<D@CLE8I M8CM< KCK@P?LMSQ DJ?N MD AMLLCARGTC RGQQSC @CRUCCL RFC ?MPR? ?LB ? TCLRPGAJC MD RFC FC?PR (RFC ?MPRGA MP NSJKML?PW QCKGJSL?P T?JTCQ) J<ILD NJ?QK? UGRFMSR RFC AM?ESJ?RGML D?ARMPQ J@EF8KI@8C (.A) EF;< RFC FC?PRsQ GLRCPL?J N?ACK?ICP; JMA?RCB LC?P RFC U?JJ MD RFC PGEFR ?RPGSK JKIFB< MFCLD<> - RFC TMJSKC MD @JMMB NSKNCB GLRM RFC ?MPR? NCP AMLRP?ARGML MD RFC JCDR TCLRPGAJC JLG<I@FI M<E8 :8M8 BP?GLQ @JMMB DPMK RFC HSESJ?P TCGL RF?R AMKCQ DPMK RFC @P?GL ?LB DPMK RFC TCGLQ RF?R AMKC DPMK RFC ?PKQ JPJK<D@: :@I:LC8K@FE DJMU MD @JMMB ?U?W DPMK RFC FC?PR RM RFC @P?GL, JGTCP, IGBLCWQ, QRMK?AF, ?LB MRFCP MPE?LQ, RFC JGK@Q, ?LB RFC KSQAJCQ MD RFC @MBW, ?LB RFCL RFC PCRSPL MD RFGQ @JMMB RM RFC FC?PR JPJKFC< AMLRP?ARGML NF?QC MD A?PBG?A AWAJC UFCL RFC TCLRPGAJCQ ?PC NSKNGLE @JMMB GLRM RFC ?PRCPGCQ KI@:LJG@; M8CM< MLC-U?W KCK@P?LMSQ DJ?N MD AMLLCARGTC RGQQSC @CRUCCL RFC ?RPGSK ?LB RFC TCLRPGAJC GL RFC PGEFR QGBC MD RFC FC?PR; ?JQM ILMUL ?Q ?RPGMTCLRPGASJ?P T?JTC LE@;@I<:K@FE8C :@I:LC8K@FE DJMU MD @JMMB GL ? QGLEJC AGPASGR; MAASPQ GL DGQF UFCPC RFC @JMMB DJMUQ RFPMSEF RFC EGJJQ, RFCL N?QR RFC MPE?LQ ?LB RFC PCQR MD RFC @MBW, @CDMPC PCRSPLGLE RM RFC FC?PR M8JF:FEJKI@:K@FE L?PPMUGLE MD ? @JMMB TCQQCJ M8JF;@C8K@FE UGBCLGLE MD ? @JMMB TCQQCJ M<@E @JMMB TCQQCJ RF?R @PGLEQ @JMMB @?AI RM RFC FC?PR
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1185
M<E8 :8M8 K?HMP TCGL MD RFC @MBW PCRSPLGLE @JMMB DPMK RFC SNNCP ?LB JMUCP N?PRQ MD RFC @MBW; QCC RFC QSNCPGMP TCL? A?T? ?LB GLDCPGMP TCL? A?T? M<EKI@:C< (FC?PR) J?PEC GLDCPGMP AF?K@CP MD RFC FC?PR RF?R NSKNQ @JMMB GLRM ?PRCPGCQ M<ELC< @JMMB TCQQCJ RF?R AMLLCARQ ? A?NGJJ?PW @CB RM ? TCGL N?@K< 9CFF; :<CC J?PEC (30 mK) ACJJ UGRF LSAJCG MD UFGAF RFCPC ?PC K?LW RWNCQ UGRF BGDDCPCLR PMJCQ GLAJSBGLE RFC NPMRCARGML MD RFC @MBW DPMK TGPSQCQ ?LB @?ARCPG?, ?LB AJC?LGLE SN BC?B ACJJQ ?LB MRFCP U?QRC
C#A+/ - .0((A-4
40.1 *M<IM@<N F= K?< C@I:LC8KFIP .PJK<D
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
40.2 CFDGFE<EKJ F= K?< BCFF;

4NCAGDGA AMKNMLCLRQ MD RFC @JMMB GLAJSBC PCB @JMMB ACJJQ, UFGRC @JMMB ACJJQ, NJ?RCJCRQ, ?LB RFC NJ?QK?, UFGAF AMLR?GLQ AM?ESJ?RGML D?ARMPQ ?LB QCPSK. #JMMB GQ GKNMPR?LR DMP PCESJ?RGML MD RFC @MBWs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
40.3 (8DD8C@8E #<8IK 8E; BCFF; 1<JJ<CJ

5FC FC?PR KSQAJC NSKNQ @JMMB RFPMSEF RFPCC BGTGQGMLQ MD RFC AGPASJ?RMPW QWQRCK: AMPML?PW, NSJKML?PW, ?LB QWQRCKGA. 5FCPC GQ MLC ?RPGSK ?LB MLC TCLRPGAJC ML RFC PGEFR QGBC ?LB MLC ?RPGSK ?LB MLC TCLRPGAJC ML RFC JCDR QGBC. 5FC NSKNGLE MD RFC FC?PR GQ ? DSLARGML MD A?PBGMKWMAWRCQ, BGQRGLARGTC KSQAJC ACJJQ RF?R ?PC QRPG?RCB JGIC QICJCR?J KSQAJC @SR NSKN PFWRFKGA?JJW ?LB GLTMJSLR?PGJW JGIC QKMMRF KSQAJC. 5FC GLRCPL?J N?ACK?ICP QR?PRQ ?R RFC QGLM?RPG?J LMBC, UFGAF GQ JMA?RCB LC?P RFC U?JJ MD RFC PGEFR ?RPGSK. &JCARPGA?J AF?PECQ NSJQC DPMK RFC 4" LMBC A?SQGLE RFC RUM ?RPG? RM AMLRP?AR GL SLGQML; RFCL RFC NSJQC PC?AFCQ RFC ?RPGMTCLRPGASJ?P LMBC @CRUCCL RFC PGEFR ?RPGSK ?LB PGEFR TCLRPGAJC. " N?SQC GL RFC CJCARPGA QGEL?J ?JJMUQ RFC ?RPG? RM CKNRW AMKNJCRCJW GLRM RFC TCLRPGAJCQ @CDMPC RFC TCLRPGAJCQ NSKN MSR RFC @JMMB. 5FC @JMMB DPMK RFC FC?PR GQ A?PPGCB RFPMSEF RFC @MBW @W ? AMKNJCV LCRUMPI MD @JMMB TCQQCJQ; ?PRCPGCQ R?IC @JMMB ?U?W DPMK RFC FC?PR, ?LB TCGLQ @PGLE @JMMB @?AI RM RFC FC?PR.
40.4 BCFF; !CFN 8E; BCFF; +I<JJLI< -<>LC8K@FE

#JMMB NPGK?PGJW KMTCQ RFPMSEF RFC @MBW @W RFC PFWRFKGA KMTCKCLR MD QKMMRF KSQAJC GL RFC TCQQCJ U?JJ ?LB @W RFC ?ARGML MD RFC QICJCR?J KSQAJC ?Q RFC @MBW KMTCQ. #JMMB GQ NPCTCLRCB DPMK DJMUGLE @?AIU?PB GL RFC TCGLQ @W MLC-U?W T?JTCQ. #JMMB DJMU RFPMSEF RFC A?NGJJ?PW @CBQ GQ AMLRPMJJCB @W NPCA?NGJJ?PW QNFGLARCPQ RM GLAPC?QC ?LB BCAPC?QC DJMU BCNCLBGLE ML RFC @MBWsQ LCCBQ ?LB GQ BGPCARCB @W LCPTC ?LB FMPKMLC QGEL?JQ. -WKNF TCQQCJQ R?IC DJSGB RF?R F?Q JC?ICB MSR MD RFC @JMMB RM RFC JWKNF LMBCQ UFCPC GR GQ AJC?LCB @CDMPC PCRSPLGLE RM RFC FC?PR. %SPGLE QWQRMJC, @JMMB CLRCPQ RFC ?PRCPGCQ, ?LB RFC ?PRCPW U?JJQ QRPCRAF RM ?AAMKKMB?RC RFC CVRP? @JMMB. %SPGLE BG?QRMJC, RFC ?PRCPW U?JJQ PCRSPL RM LMPK?J.
1186
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5FC @JMMB NPCQQSPC MD RFC QWQRMJC NF?QC ?LB RFC BG?QRMJC NF?QC EGTCQ RFC RUM NPCQQSPC PC?BGLEQ DMP @JMMB NPCQQSPC.
A-/ C*)) C/$*) ,0 ./$*).

1. Figure 40.10 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR RFC AGPASJ?RMPW QWQRCK GQ D?JQC >. #JMMB GL RFC NSJKML?PW TCGL GQ BCMVWECL?RCB. ?. #JMMB GL RFC GLDCPGMP TCL? A?T? GQ BCMVWECL?RCB. @. #JMMB GL RFC NSJKML?PW ?PRCPW GQ BCMVWECL?RCB. A. #JMMB GL RFC ?MPR? GQ MVWECL?RCB. 2. Figure 40.11 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR RFC FC?PR GQ D?JQC >. 5FC KGRP?J T?JTC QCN?P?RCQ RFC JCDR TCLRPGAJC DPMK RFC JCDR ?RPGSK. ?. #JMMB RP?TCJQ RFPMSEF RFC @GASQNGB T?JTC RM RFC JCDR ?RPGSK. @. #MRF RFC ?MPRGA ?LB RFC NSJKML?PW T?JTCQ ?PC QCKGJSL?P T?JTCQ. A. 5FC KGRP?J T?JTC GQ ?L ?RPGMTCLRPGASJ?P T?JTC. 3. Figure 40.17 7?PGAMQC TCGLQ ?PC TCGLQ RF?R @CAMKC CLJ?PECB @CA?SQC RFC T?JTCQ LM JMLECP AJMQC NPMNCPJW, ?JJMUGLE @JMMB RM DJMU @?AIU?PB. 7?PGAMQC TCGLQ ?PC MDRCL KMQR NPMKGLCLR ML RFC JCEQ. 8FW BM WMS RFGLI RFGQ GQ RFC A?QC
- 1$ 2 ,0 ./$*).
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sQ GLRCPL?J N?ACK?ICP @C?RQ @W: >. ?L GLRCPL?J GKNJ?LR RF?R QCLBQ ?L CJCARPGA?J GKNSJQC RFPMSEF RFC FC?PR ?. RFC CVAGR?RGML MD A?PBG?A KSQAJC ACJJQ ?R RFC QGLM?RPG?J LMBC DMJJMUCB @W RFC ?RPGMTCLRPGASJ?P LMBC @. RFC CVAGR?RGML MD A?PBG?A KSQAJC ACJJQ ?R RFC ?RPGMTCLRPGASJ?P LMBC DMJJMUCB @W RFC QGLM?RPG?J LMBC A. RFC ?ARGML MD RFC QGLSQ 12. %SPGLE RFC QWQRMJGA NF?QC MD RFC A?PBG?A AWAJC, RFC FC?PR GQ >>>>>>>>. >. AMLRP?ARGLE ?. PCJ?VGLE @. AMLRP?ARGLE ?LB PCJ?VGLE A. DGJJGLE UGRF @JMMB 13. $?PBGMKWMAWRCQ ?PC QGKGJ?P RM QICJCR?J KSQAJC @CA?SQC: >. RFCW @C?R GLTMJSLR?PGJW ?. RFCW ?PC SQCB DMP UCGEFR JGDRGLE @. RFCW NSJQC PFWRFKGA?JJW A. RFCW ?PC QRPG?RCB 14. )MU BM ?PRCPGCQ BGDDCP DPMK TCGLQ >. "PRCPGCQ F?TC RFGAICP QKMMRF KSQAJC J?WCPQ RM ?AAMKKMB?RC RFC AF?LECQ GL NPCQQSPC DPMK RFC FC?PR. ?. "PRCPGCQ A?PPW @JMMB.
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>. ? FGEF A?PBG?A MSRNSR ?LB FGEF NCPGNFCP?J PCQGQR?LAC ?. ? FGEF A?PBG?A MSRNSR ?LB JMU NCPGNFCP?J PCQGQR?LAC @. ? JMU A?PBG?A MSRNSR ?LB FGEF NCPGNFCP?J PCQGQR?LAC A. ? JMU A?PBG?A MSRNSR ?LB JMU NCPGNFCP?J PCQGQR?LAC
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16. %CQAPG@C ? AJMQCB AGPASJ?RMPW QWQRCK. 17. %CQAPG@C QWQRCKGA AGPASJ?RGML. 18. %CQAPG@C RFC A?SQC MD BGDDCPCLR @JMMB RWNC EPMSNQ. 19. -GQR QMKC MD RFC DSLARGMLQ MD @JMMB GL RFC @MBW. 20. )MU BMCQ RFC JWKNF?RGA QWQRCK UMPI UGRF @JMMB DJMU 21. %CQAPG@C RFC A?PBG?A AWAJC. 22. 8F?R F?NNCLQ GL A?NGJJ?PGCQ 23. )MU BMCQ @JMMB NPCQQSPC AF?LEC BSPGLE FC?TW CVCPAGQC
1188
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
1190
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41.1 R *JDFI<>LC8K@FE 8E; *JDFK@: B8C8E:<

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CDGLC MQKMQGQ ?LB CVNJ?GL GRQ PMJC UGRFGL KMJCASJCQ z &VNJ?GL UFW MQKMPCESJ?RGML ?LB MQKMRGA @?J?LAC ?PC GKNMPR?LR @MBW DSLARGMLQ z %CQAPG@C ?ARGTC RP?LQNMPR KCAF?LGQKQ z &VNJ?GL MQKMJ?PGRW ?LB RFC U?W GL UFGAF GR GQ KC?QSPCB z %CQAPG@C MQKMPCESJ?RMPQ MP MQKMAMLDMPKCPQ ?LB FMU RFCQC RMMJQ ?JJMU ?LGK?JQ RM ?B?NR RM BGDDCPCLR CLTGPMLKCLRQ 0QKMQGQ GQ RFC BGDDSQGML MD U?RCP ?APMQQ ? KCK@P?LC GL PCQNMLQC RM osmotic pressure A?SQCB @W ?L GK@?J?LAC MD KMJCASJCQ ML CGRFCP QGBC MD RFC KCK@P?LC. Osmoregulation GQ RFC NPMACQQ MD K?GLRCL?LAC MD Q?JR ?LB U?RCP @?J?LAC ( osmotic balance) ?APMQQ KCK@P?LCQ UGRFGL RFC @MBWsQ DJSGBQ, UFGAF ?PC AMKNMQCB MD U?RCP, NJSQ CJCARPMJWRCQ ?LB LML-CJCARPMJWRCQ. "L electrolyte GQ ? QMJSRC RF?R BGQQMAG?RCQ GLRM GMLQ UFCL BGQQMJTCB GL U?RCP. " non-electrolyte, GL AMLRP?QR, BMCQLsR BGQQMAG?RC GLRM GMLQ BSPGLE U?RCP BGQQMJSRGML. #MRF CJCARPMJWRCQ ?LB LML-CJCARPMJWRCQ AMLRPG@SRC RM RFC MQKMRGA @?J?LAC. 5FC @MBWsQ DJSGBQ GLAJSBC @JMMB NJ?QK?, RFC AWRMQMJ UGRFGL ACJJQ, ?LB GLRCPQRGRG?J DJSGB, RFC DJSGB RF?R CVGQRQ GL RFC QN?ACQ @CRUCCL ACJJQ ?LB RGQQSCQ MD RFC @MBW. 5FC KCK@P?LCQ MD RFC @MBW (QSAF ?Q RFC NJCSP?J, QCPMSQ, ?LB ACJJ KCK@P?LCQ) ?PC semi-permeable membranes. 4CKG-NCPKC?@JC KCK@P?LCQ ?PC NCPKC?@JC (MP NCPKGQQGTC) RM ACPR?GL RWNCQ MD QMJSRCQ ?LB U?RCP. 4MJSRGMLQ ML RUM QGBCQ MD ? QCKG-NCPKC?@JC KCK@P?LC RCLB RM COS?JGXC GL QMJSRC AMLACLRP?RGML @W KMTCKCLR MD QMJSRCQ ?LB/MP U?RCP ?APMQQ RFC KCK@P?LC. "Q QCCL GL Figure 41.2, ? ACJJ NJ?ACB GL U?RCP RCLBQ RM QUCJJ BSC RM E?GL MD U?RCP DPMK RFC FWNMRMLGA MP tJMU Q?JRu CLTGPMLKCLR. " ACJJ NJ?ACB GL ? QMJSRGML UGRF FGEFCP Q?JR AMLACLRP?RGML, ML RFC MRFCP F?LB, RCLBQ RM K?IC RFC KCK@P?LC QFPGTCJ SN BSC RM JMQQ MD U?RCP GLRM RFC FWNCPRMLGA MP tFGEF Q?JRu CLTGPMLKCLR. *QMRMLGA ACJJQ F?TC ?L COS?J AMLACLRP?RGML MD QMJSRCQ GLQGBC ?LB MSRQGBC RFC ACJJ; RFGQ COS?JGXCQ RFC MQKMRGA NPCQQSPC ML CGRFCP QGBC MD RFC ACJJ KCK@P?LC UFGAF GQ ? QCKG-NCPKC?@JC KCK@P?LC.
!@>LI< 41.2 #BIIP MI>@BA FK > EVMBOQLKF@ BKSFOLKJBKQ QBKA QL PEOFKH ARB QL ILPP LC T>QBO. )K > EVMLQLKF@ BKSFOLKJBKQ, @BIIP QBKA QL PTBII ARB QL FKQ>HB LC T>QBO. 4EB ?ILLA J>FKQ>FKP >K FPLQLKF@ BKSFOLKJBKQ PL QE>Q @BIIP KBFQEBO PEOFKH KLO PTBII. (@OBAFQ: ->OF>K> 2RFW 6FII>OB>I)
5FC @MBW BMCQ LMR CVGQR GL GQMJ?RGML. 5FCPC GQ ? AMLQR?LR GLNSR MD U?RCP ?LB CJCARPMJWRCQ GLRM RFC QWQRCK. 8FGJC MQKMPCESJ?RGML GQ ?AFGCTCB ?APMQQ KCK@P?LCQ UGRFGL RFC @MBW, CVACQQ CJCARPMJWRCQ ?LB U?QRCQ ?PC RP?LQNMPRCB RM RFC IGBLCWQ ?LB CVAPCRCB, FCJNGLE RM K?GLR?GL MQKMRGA @?J?LAC.
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/I8EJGFIK F= C<:KIFCPK<J 8:IFJJ C<CC (<D9I8E<J

&JCARPMJWRCQ, QSAF ?Q QMBGSK AFJMPGBC, GMLGXC GL U?RCP, KC?LGLE RF?R RFCW BGQQMAG?RC GLRM RFCGP AMKNMLCLR GMLQ. *L U?RCP, QMBGSK AFJMPGBC (/?$J), BGQQMAG?RCQ GLRM RFC QMBGSK GML (/?+) ?LB RFC AFJMPGBC GML ($Jp). 5FC KMQR GKNMPR?LR GMLQ, UFMQC AMLACLRP?RGMLQ ?PC TCPW AJMQCJW PCESJ?RCB GL @MBW DJSGBQ, ?PC RFC A?RGMLQ QMBGSK (/?+), NMR?QQGSK (,+), A?JAGSK ($?+2), K?ELCQGSK (.E+2), ?LB RFC ?LGMLQ AFJMPGBC ($J-), A?P@ML?RC ($03-2), @GA?P@ML?RC ()$03-), ?LB NFMQNF?RC(103-). &JCARPMJWRCQ ?PC JMQR DPMK RFC @MBW BSPGLE SPGL?RGML ?LB NCPQNGP?RGML. 'MP RFGQ PC?QML, ?RFJCRCQ ?PC CLAMSP?ECB RM PCNJ?AC CJCARPMJWRCQ ?LB DJSGBQ BSPGLE NCPGMBQ MD GLAPC?QCB ?ARGTGRW ?LB NCPQNGP?RGML. 0QKMRGA NPCQQSPC GQ GLDJSCLACB @W RFC AMLACLRP?RGML MD QMJSRCQ GL ? QMJSRGML. *R GQ BGPCARJW NPMNMPRGML?J RM RFC LSK@CP MD QMJSRC ?RMKQ MP KMJCASJCQ ?LB LMR BCNCLBCLR ML RFC QGXC MD RFC QMJSRC KMJCASJCQ. #CA?SQC CJCARPMJWRCQ BGQQMAG?RC GLRM RFCGP AMKNMLCLR GMLQ, RFCW, GL CQQCLAC, ?BB KMPC QMJSRC N?PRGAJCQ GLRM RFC QMJSRGML ?LB F?TC ? EPC?RCP CDDCAR ML MQKMRGA NPCQQSPC, NCP K?QQ RF?L AMKNMSLBQ RF?R BM LMR BGQQMAG?RC GL U?RCP, QSAF ?Q EJSAMQC. 8?RCP A?L N?QQ RFPMSEF KCK@P?LCQ @W N?QQGTC BGDDSQGML. *D CJCARPMJWRC GMLQ AMSJB N?QQGTCJW BGDDSQC ?APMQQ KCK@P?LCQ, GR UMSJB @C GKNMQQG@JC RM K?GLR?GL QNCAGDGA AMLACLRP?RGMLQ MD GMLQ GL C?AF DJSGB AMKN?PRKCLR RFCPCDMPC RFCW PCOSGPC QNCAG?J KCAF?LGQKQ RM APMQQ RFC QCKG-NCPKC?@JC KCK@P?LCQ GL RFC @MBW. 5FGQ KMTCKCLR A?L @C ?AAMKNJGQFCB @W D?AGJGR?RCB BGDDSQGML ?LB ?ARGTC RP?LQNMPR. '?AGJGR?RCB BGDDSQGML PCOSGPCQ NPMRCGL-@?QCB AF?LLCJQ DMP KMTGLE RFC QMJSRC. "ARGTC RP?LQNMPR PCOSGPCQ CLCPEW GL RFC DMPK MD "51 AMLTCPQGML, A?PPGCP NPMRCGLQ, MP NSKNQ GL MPBCP RM KMTC GMLQ ?E?GLQR RFC AMLACLRP?RGML EP?BGCLR.
CFE:<GK F= *JDFC8C@KP 8E; (@CC@<HL@M8C<EK

*L MPBCP RM A?JASJ?RC MQKMRGA NPCQQSPC, GR GQ LCACQQ?PW RM SLBCPQR?LB FMU QMJSRC AMLACLRP?RGMLQ ?PC KC?QSPCB. 5FC SLGR DMP KC?QSPGLE QMJSRCQ GQ RFC mole. 0LC KMJC GQ BCDGLCB ?Q RFC EP?K KMJCASJ?P UCGEFR MD RFC QMJSRC. 'MP CV?KNJC, RFC KMJCASJ?P UCGEFR MD QMBGSK AFJMPGBC GQ 58.44. 5FSQ, MLC KMJC MD QMBGSK AFJMPGBC UCGEFQ 58.44 EP?KQ. 5FC molarity MD ? QMJSRGML GQ RFC LSK@CP MD KMJCQ MD QMJSRC NCP JGRCP MD QMJSRGML. 5FC molality MD ? QMJSRGML GQ RFC LSK@CP MD KMJCQ MD QMJSRC NCP IGJMEP?K MD QMJTCLR. *D RFC QMJTCLR GQ U?RCP, MLC IGJMEP?K MD U?RCP GQ COS?J RM MLC JGRCP MD U?RCP. 8FGJC KMJ?PGRW ?LB KMJ?JGRW ?PC SQCB RM CVNPCQQ RFC AMLACLRP?RGML MD QMJSRGMLQ, CJCARPMJWRC AMLACLRP?RGMLQ ?PC SQS?JJW CVNPCQQCB GL RCPKQ MD KGJJGCOSGT?JCLRQ NCP JGRCP (K&O/-): RFC K&O/- GQ COS?J RM RFC GML AMLACLRP?RGML (GL KGJJGKMJCQ) KSJRGNJGCB @W RFC LSK@CP MD CJCARPGA?J AF?PECQ ML RFC GML. 5FC SLGR MD KGJJGCOSGT?JCLR R?ICQ GLRM AMLQGBCP?RGML RFC GMLQ NPCQCLR GL RFC QMJSRGML (QGLAC CJCARPMJWRCQ DMPK GMLQ GL ?OSCMSQ QMJSRGMLQ) ?LB RFC AF?PEC ML RFC GMLQ. 5FSQ, DMP GMLQ RF?R F?TC ? AF?PEC MD MLC, MLC KGJJGCOSGT?JCLR GQ COS?J RM MLC KGJJGKMJC. 'MP GMLQ RF?R F?TC ? AF?PEC MD RUM (JGIC A?JAGSK), MLC KGJJGCOSGT?JCLR GQ COS?J RM 0.5 KGJJGKMJCQ. "LMRFCP SLGR DMP RFC CVNPCQQGML MD CJCARPMJWRC AMLACLRP?RGML GQ RFC KGJJGMQKMJC (K0QK), UFGAF GQ RFC LSK@CP MD KGJJGCOSGT?JCLRQ MD QMJSRC NCP IGJMEP?K MD QMJTCLR. #MBW DJSGBQ ?PC SQS?JJW K?GLR?GLCB UGRFGL RFC P?LEC MD 280 RM 300 K0QK.
*JDFI<>LC8KFIJ 8E; *JDF:FE=FID<IJ

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osmoregulatory KCAF?LGQKQ RM QSPTGTC GL ?JJ IGLBQ MD ?OS?RGA CLTGPMLKCLRQ. 8FCL RFCW JGTC GL DPCQF U?RCP, RFCGP @MBGCQ RCLB RM R?IC SN U?RCP @CA?SQC RFC CLTGPMLKCLR GQ PCJ?RGTCJW FWNMRMLGA, ?Q GJJSQRP?RCB GL Figure 41.3a. *L QSAF FWNMRMLGA CLTGPMLKCLRQ, RFCQC DGQF BM LMR BPGLI KSAF U?RCP. *LQRC?B, RFCW N?QQ ? JMR MD TCPW BGJSRC SPGLC, ?LB RFCW ?AFGCTC CJCARPMJWRC @?J?LAC @W ?ARGTC RP?LQNMPR MD Q?JRQ RFPMSEF RFC EGJJQ. 8FCL RFCW KMTC RM ? FWNCPRMLGA K?PGLC CLTGPMLKCLR, RFCQC DGQF QR?PR BPGLIGLE QC? U?RCP; RFCW CVAPCRC RFC CVACQQ Q?JRQ RFPMSEF RFCGP EGJJQ ?LB RFCGP SPGLC, ?Q GJJSQRP?RCB GL Figure 41.3b. .MQR K?PGLC GLTCPRC@P?RCQ, ML RFC MRFCP F?LB, K?W @C GQMRMLGA UGRF QC? U?RCP ( osmoconformers). 5FCGP @MBW DJSGB AMLACLRP?RGMLQ AMLDMPK RM AF?LECQ GL QC?U?RCP AMLACLRP?RGML. $?PRGJ?EGLMSQ DGQFCQs Q?JR AMKNMQGRGML MD RFC @JMMB GQ QGKGJ?P RM @MLW DGQFCQ; FMUCTCP, RFC @JMMB MD QF?PIQ AMLR?GLQ RFC MPE?LGA AMKNMSLBQ SPC? ?LB RPGKCRFWJ?KGLC MVGBC (5."0). 5FGQ BMCQ LMR KC?L RF?R RFCGP CJCARPMJWRC AMKNMQGRGML GQ QGKGJ?P RM RF?R MD QC? U?RCP. 5FCW ?AFGCTC GQMRMLGAGRW UGRF RFC QC? @W QRMPGLE J?PEC AMLACLRP?RGMLQ MD SPC?. 5FCQC ?LGK?JQ RF?R QCAPCRC SPC? ?PC A?JJCB SPCMRCJGA ?LGK?JQ. 5."0 QR?@GJGXCQ NPMRCGLQ GL RFC NPCQCLAC MD FGEF SPC? JCTCJQ, NPCTCLRGLE RFC BGQPSNRGML MD NCNRGBC @MLBQ RF?R
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KSJRGNJC NWP?KGB?J RGQQSC K?QQCQ, A?JJCB RFC renal pyramids. *L @CRUCCL RFC NWP?KGBQ ?PC QN?ACQ A?JJCB renal columns RFPMSEF UFGAF RFC @JMMB TCQQCJQ N?QQ. 5FC RGNQ MD RFC NWP?KGBQ, A?JJCB PCL?J N?NGJJ?C, NMGLR RMU?PB RFC PCL?J NCJTGQ. 5FCPC ?PC, ML ?TCP?EC, CGEFR PCL?J NWP?KGBQ GL C?AF IGBLCW. 5FC PCL?J NWP?KGBQ ?JMLE UGRF RFC ?BHMGLGLE AMPRGA?J PCEGML ?PC A?JJCB RFC lobes of the kidney. 5FC PCL?J NCJTGQ JC?BQ RM RFC ureter ML RFC MSRQGBC MD RFC IGBLCW. 0L RFC GLQGBC MD RFC IGBLCW, RFC PCL?J NCJTGQ @P?LAFCQ MSR GLRM RUM MP RFPCC CVRCLQGMLQ A?JJCB RFC K?HMP calyces, UFGAF DSPRFCP @P?LAF GLRM RFC KGLMP A?JWACQ. 5FC SPCRCPQ ?PC SPGLC-@C?PGLE RS@CQ RF?R CVGR RFC IGBLCW ?LB CKNRW GLRM RFC urinary bladder.
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7EF@E LC QEB CLIILTFKD PQ>QBJBKQP >?LRQ QEB KBMEOLK FP C>IPB >. 4EB @LIIB@QFKD AR@Q BJMQFBP FKQL QEB AFPQ>I @LKSLIRQBA QR?RIB. ?. 4EB "LTJ>KwP @>MPRIB PROOLRKAP QEB DILJBORIRP. @. 4EB ILLM LC (BKIB FP ?BQTBBK QEB MOLUFJ>I >KA AFPQ>I @LKSLIRQBA QR?RIBP. A. 4EB ILLM LC (BKIB BJMQFBP FKQL QEB AFPQ>I @LKSLIRQBA QR?RIB. +3'1 C475:8)1+ 5FC PCL?J AMPNSQAJC, JMA?RCB GL RFC PCL?J AMPRCV, GQ K?BC SN MD ? LCRUMPI MD A?NGJJ?PGCQ ILMUL ?Q RFC glomerulus ?LB RFC A?NQSJC, ? ASN-QF?NCB AF?K@CP RF?R QSPPMSLBQ GR, A?JJCB RFC EJMKCPSJ?P MP Bowman's capsule. +3'1 ":(:1+ 5FC PCL?J RS@SJC GQ ? JMLE ?LB AMLTMJSRCB QRPSARSPC RF?R CKCPECQ DPMK RFC EJMKCPSJSQ ?LB A?L @C BGTGBCB GLRM RFPCC N?PRQ @?QCB ML DSLARGML. 5FC DGPQR N?PR GQ A?JJCB RFC proximal convoluted tubule (PCT) BSC RM GRQ NPMVGKGRW RM RFC EJMKCPSJSQ; GR QR?WQ GL RFC PCL?J AMPRCV. 5FC QCAMLB N?PR GQ A?JJCB RFC loop of Henle, MP LCNFPGRGA JMMN, @CA?SQC GR DMPKQ ? JMMN (UGRF descending ?LB ascending limbs) RF?R EMCQ RFPMSEF RFC PCL?J KCBSJJ?. 5FC RFGPB N?PR MD RFC PCL?J RS@SJC GQ A?JJCB RFC distal convoluted tubule (DCT) ?LB RFGQ N?PR GQ ?JQM PCQRPGARCB RM RFC PCL?J AMPRCV. 5FC %$5, UFGAF GQ RFC J?QR N?PR MD RFC LCNFPML, AMLLCARQ ?LB CKNRGCQ GRQ AMLRCLRQ GLRM AMJJCARGLE BSARQ RF?R JGLC RFC KCBSJJ?PW NWP?KGBQ. 5FC AMJJCARGLE BSARQ ?K?QQ AMLRCLRQ DPMK KSJRGNJC LCNFPMLQ ?LB DSQC RMECRFCP ?Q RFCW CLRCP RFC N?NGJJ?C MD RFC PCL?J KCBSJJ?. C'5/11'7> N+9<470 </9./3 9.+ N+5.743 5FC A?NGJJ?PW LCRUMPI RF?R MPGEGL?RCQ DPMK RFC PCL?J ?PRCPGCQ QSNNJGCQ RFC LCNFPML UGRF @JMMB RF?R LCCBQ RM @C DGJRCPCB. 5FC @P?LAF RF?R CLRCPQ RFC EJMKCPSJSQ GQ A?JJCB RFC afferent arteriole. 5FC @P?LAF RF?R CVGRQ RFC EJMKCPSJSQ GQ A?JJCB RFC efferent arteriole. 8GRFGL RFC EJMKCPSJSQ, RFC LCRUMPI MD A?NGJJ?PGCQ GQ A?JJCB RFC EJMKCPSJ?P A?NGJJ?PW @CB. 0LAC RFC CDDCPCLR ?PRCPGMJC CVGRQ RFC EJMKCPSJSQ, GR DMPKQ RFC peritubular capillary network, UFGAF QSPPMSLBQ ?LB GLRCP?ARQ UGRF N?PRQ MD RFC PCL?J RS@SJC. *L AMPRGA?J LCNFPMLQ, RFC NCPGRS@SJ?P A?NGJJ?PW LCRUMPI QSPPMSLBQ RFC 1$5 ?LB %$5. *L HSVR?KCBSJJ?PW LCNFPMLQ, RFC NCPGRS@SJ?P A?NGJJ?PW LCRUMPI DMPKQ ? LCRUMPI ?PMSLB RFC JMMN MD )CLJC ?LB GQ A?JJCB RFC vasa recta.
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
G142+7:1'7 F/197'9/43 (JMKCPSJ?P DGJRP?RGML DGJRCPQ MSR KMQR MD RFC QMJSRCQ BSC RM FGEF @JMMB NPCQQSPC ?LB QNCAG?JGXCB KCK@P?LCQ GL RFC ?DDCPCLR ?PRCPGMJC. 5FC @JMMB NPCQQSPC GL RFC EJMKCPSJSQ GQ K?GLR?GLCB GLBCNCLBCLR MD D?ARMPQ RF?R ?DDCAR QWQRCKGA @JMMB NPCQQSPC. 5FC tJC?IWu AMLLCARGMLQ @CRUCCL RFC CLBMRFCJG?J ACJJQ MD RFC EJMKCPSJ?P A?NGJJ?PW LCRUMPI ?JJMU QMJSRCQ RM N?QQ RFPMSEF C?QGJW. "JJ QMJSRCQ GL RFC EJMKCPSJ?P A?NGJJ?PGCQ, CVACNR DMP K?APMKMJCASJCQ JGIC NPMRCGLQ, N?QQ RFPMSEF @W N?QQGTC BGDDSQGML. 5FCPC GQ LM CLCPEW PCOSGPCKCLR ?R RFGQ QR?EC MD RFC DGJRP?RGML NPMACQQ. Glomerular filtration rate (GFR) GQ RFC TMJSKC MD EJMKCPSJ?P DGJRP?RC DMPKCB NCP KGLSRC @W RFC IGBLCWQ. ('3 GQ PCESJ?RCB @W KSJRGNJC KCAF?LGQKQ ?LB GQ ?L GKNMPR?LR GLBGA?RMP MD IGBLCW DSLARGML.
5M JC?PL KMPC ?@MSR RFC T?QASJ?P QWQRCK MD IGBLCWQ, (http://openstaxcollege.org/l/kidneys) ?LB RFC QRCNQ MD @JMMB DJMU.
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,LLM AFROBQF@P >OB AORDP PLJBQFJBP RPBA QL QOB>Q EVMBOQBKPFLK. 4EBPB AORDP FKEF?FQ QEB OB>?PLOMQFLK LC .>+ >KA #I- FLKP ?V QEB >P@BKAFKD IFJ? LC QEB ILLM LC (BKIB. ! PFAB BCCB@Q FP QE>Q QEBV FK@OB>PB ROFK>QFLK. 7EV AL VLR QEFKH QEFP FP QEB @>PB #W RFC RGKC RFC DGJRP?RC PC?AFCQ RFC %$5, KMQR MD RFC SPGLC ?LB QMJSRCQ F?TC @CCL PC?@QMP@CB. *D RFC @MBW PCOSGPCQ ?BBGRGML?J U?RCP, ?JJ MD GR A?L @C PC?@QMP@CB ?R RFGQ NMGLR. 'SPRFCP PC?@QMPNRGML GQ AMLRPMJJCB @W FMPKMLCQ, UFGAF UGJJ @C BGQASQQCB GL ? J?RCP QCARGML. &VAPCRGML MD U?QRCQ MAASPQ BSC RM J?AI MD PC?@QMPNRGML AMK@GLCB UGRF RS@SJ?P QCAPCRGML. 6LBCQGP?@JC NPMBSARQ JGIC KCR?@MJGA U?QRCQ, SPC?, SPGA ?AGB, ?LB ACPR?GL BPSEQ, ?PC CVAPCRCB @W RS@SJ?P QCAPCRGML. .MQR MD RFC RS@SJ?P QCAPCRGML F?NNCLQ GL RFC %$5, @SR QMKC MAASPQ GL RFC C?PJW N?PR MD RFC AMJJCARGLE BSAR. ,GBLCWQ ?JQM K?GLR?GL ?L ?AGB-@?QC @?J?LAC @W QCAPCRGLE CVACQQ )+ GMLQ.
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL FMU T?ASMJCQ, NPCQCLR GL KGAPMMPE?LGQKQ, UMPI RM CVAPCRC U?QRC z %CQAPG@C RFC U?W GL UFGAF DJ?KC ACJJQ ?LB LCNFPGBG? GL UMPKQ NCPDMPK CVAPCRMPW DSLARGMLQ ?LB K?GLR?GL MQKMRGA @?J?LAC z &VNJ?GL FMU GLQCARQ SQC .?JNGEFG?L RS@SJCQ RM CVAPCRC U?QRCQ ?LB K?GLR?GL MQKMRGA @?J?LAC .GAPMMPE?LGQKQ ?LB GLTCPRC@P?RC ?LGK?JQ SQC KMPC NPGKGRGTC ?LB QGKNJC KCAF?LGQKQ RM ECR PGB MD RFCGP KCR?@MJGA U?QRCQ RF?L RFC K?KK?JG?L QWQRCK MD IGBLCW ?LB SPGL?PW DSLARGML. 5FPCC CVAPCRMPW QWQRCKQ CTMJTCB GL MPE?LGQKQ @CDMPC AMKNJCV IGBLCWQ: T?ASMJCQ, DJ?KC ACJJQ, ?LB .?JNGEFG?L RS@SJCQ.
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1200
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1202
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4MJSRC AMLACLRP?RGMLQ ?APMQQ ? QCKG-NCPKC?@JC KCK@P?LCQ GLDJSCLAC RFC KMTCKCLR MD U?RCP ?LB QMJSRCQ ?APMQQ RFC KCK@P?LC. *R GQ RFC LSK@CP MD QMJSRC KMJCASJCQ ?LB LMR RFC KMJCASJ?P QGXC RF?R GQ GKNMPR?LR GL MQKMQGQ. 0QKMPCESJ?RGML ?LB MQKMRGA @?J?LAC ?PC GKNMPR?LR @MBGJW DSLARGMLQ, PCQSJRGLE GL U?RCP ?LB Q?JR @?J?LAC. /MR ?JJ QMJSRCQ A?L N?QQ RFPMSEF ? QCKG-NCPKC?@JC KCK@P?LC. 0QKMQGQ GQ RFC KMTCKCLR MD U?RCP ?APMQQ RFC KCK@P?LC. 0QKMQGQ MAASPQ RM COS?JGXC RFC LSK@CP MD QMJSRC KMJCASJCQ ?APMQQ ? QCKG-NCPKC?@JC KCK@P?LC @W RFC KMTCKCLR MD U?RCP RM RFC QGBC MD FGEFCP QMJSRC AMLACLRP?RGML. '?AGJGR?RCB BGDDSQGML SRGJGXCQ NPMRCGL AF?LLCJQ RM KMTC QMJSRC KMJCASJCQ DPMK ?PC?Q MD FGEFCP RM JMUCP AMLACLRP?RGML UFGJC ?ARGTC RP?LQNMPR KCAF?LGQKQ ?PC PCOSGPCB RM KMTC QMJSRCQ ?E?GLQR AMLACLRP?RGML EP?BGCLRQ. 0QKMJ?PGRW GQ KC?QSPCB GL SLGRQ MD KGJJGCOSGT?JCLRQ MP KGJJGMQKMJCQ, @MRF MD UFGAF R?IC GLRM AMLQGBCP?RGML RFC LSK@CP MD QMJSRC N?PRGAJCQ ?LB RFC AF?PEC ML RFCK. 'GQF RF?R JGTC GL DPCQF U?RCP MP Q?JRU?RCP ?B?NR @W @CGLE MQKMPCESJ?RMPQ MP MQKMAMLDMPKCPQ.
41.2 /?< &@;E<PJ 8E; *JDFI<>LC8KFIP *I>8EJ

5FC IGBLCWQ ?PC RFC K?GL MQKMPCESJ?RMPW MPE?LQ GL K?KK?JG?L QWQRCKQ; RFCW DSLARGML RM DGJRCP @JMMB ?LB K?GLR?GL RFC MQKMJ?PGRW MD @MBW DJSGBQ ?R 300 K0QK. 5FCW ?PC QSPPMSLBCB @W RFPCC J?WCPQ ?LB ?PC K?BC SN GLRCPL?JJW MD RFPCC BGQRGLAR PCEGMLQqRFC AMPRCV, KCBSJJ?, ?LB NCJTGQ. 5FC @JMMB TCQQCJQ RF?R RP?LQNMPR @JMMB GLRM ?LB MSR MD RFC IGBLCWQ ?PGQC DPMK ?LB KCPEC UGRF RFC ?MPR? ?LB GLDCPGMP TCL? A?T?, PCQNCARGTCJW. 5FC PCL?J ?PRCPGCQ @P?LAF MSR DPMK RFC ?MPR? ?LB CLRCP RFC IGBLCW UFCPC RFCW DSPRFCP BGTGBC GLRM QCEKCLR?J, GLRCPJM@?P, ?PAS?RC, ?LB AMPRGA?J P?BG?RC ?PRCPGCQ. 5FC LCNFPML GQ RFC DSLARGML?J SLGR MD RFC IGBLCW, UFGAF ?ARGTCJW DGJRCPQ @JMMB ?LB ECLCP?RCQ SPGLC. 5FC LCNFPML GQ K?BC SN MD RFC PCL?J AMPNSQAJC ?LB PCL?J RS@SJC. $MPRGA?J LCNFPMLQ ?PC DMSLB GL RFC PCL?J AMPRCV, UFGJC HSVR?KCBSJJ?PW LCNFPMLQ ?PC DMSLB GL RFC PCL?J AMPRCV AJMQC RM RFC PCL?J KCBSJJ?. 5FC LCNFPML DGJRCPQ ?LB CVAF?LECQ U?RCP ?LB QMJSRCQ UGRF RUM QCRQ MD @JMMB TCQQCJQ ?LB RFC RGQQSC DJSGB GL RFC IGBLCWQ.
1210
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5FCPC ?PC RFPCC QRCNQ GL RFC DMPK?RGML MD SPGLC: EJMKCPSJ?P DGJRP?RGML, UFGAF MAASPQ GL RFC EJMKCPSJSQ; RS@SJ?P PC?@QMPNRGML, UFGAF MAASPQ GL RFC PCL?J RS@SJCQ; ?LB RS@SJ?P QCAPCRGML, UFGAF ?JQM MAASPQ GL RFC PCL?J RS@SJCQ.
41.3 O:I<K@FE .PJK<DJ

.?LW QWQRCKQ F?TC CTMJTCB DMP CVAPCRGLE U?QRCQ RF?R ?PC QGKNJCP RF?L RFC IGBLCW ?LB SPGL?PW QWQRCKQ MD TCPRC@P?RC ?LGK?JQ. 5FC QGKNJCQR QWQRCK GQ RF?R MD AMLRP?ARGJC T?ASMJCQ NPCQCLR GL KGAPMMPE?LGQKQ. 'J?KC ACJJQ ?LB LCNFPGBG? GL UMPKQ NCPDMPK CVAPCRMPW DSLARGMLQ ?LB K?GLR?GL MQKMRGA @?J?LAC. 4MKC GLQCARQ F?TC CTMJTCB .?JNGEFG?L RS@SJCQ RM CVAPCRC U?QRCQ ?LB K?GLR?GL MQKMRGA @?J?LAC.
41.4 )@KIF><EFLJ 28JK<J

"KKMLG? GQ RFC U?QRC NPMBSACB @W KCR?@MJGQK MD LGRPMECL-AMLR?GLGLE AMKNMSLBQ JGIC NPMRCGLQ ?LB LSAJCGA ?AGBQ. 8FGJC ?OS?RGA ?LGK?JQ A?L C?QGJW CVAPCRC ?KKMLG? GLRM RFCGP U?RCPW QSPPMSLBGLEQ, RCPPCQRPG?J ?LGK?JQ F?TC CTMJTCB QNCAG?J KCAF?LGQKQ RM CJGKGL?RC RFC RMVGA ?KKMLG? DPMK RFCGP QWQRCKQ. 6PC? GQ RFC K?HMP @WNPMBSAR MD ?KKMLG? KCR?@MJGQK GL TCPRC@P?RC ?LGK?JQ. 6PGA ?AGB GQ RFC K?HMP @WNPMBSAR MD ?KKMLG? KCR?@MJGQK GL @GPBQ, RCPPCQRPG?J ?PRFPMNMBQ, ?LB PCNRGJCQ.
41.5 #FIDFE8C CFEKIFC F= *JDFI<>LC8KFIP !LE:K@FEJ

)MPKML?J ASCQ FCJN RFC IGBLCWQ QWLAFPMLGXC RFC MQKMRGA LCCBQ MD RFC @MBW. )MPKMLCQ JGIC CNGLCNFPGLC, LMPCNGLCNFPGLC, PCLGL-?LEGMRCLQGL, ?JBMQRCPMLC, ?LRG-BGSPCRGA FMPKMLC, ?LB ?RPG?J L?RPGSPCRGA NCNRGBC FCJN PCESJ?RC RFC LCCBQ MD RFC @MBW ?Q UCJJ ?Q RFC AMKKSLGA?RGML @CRUCCL RFC BGDDCPCLR MPE?L QWQRCKQ.
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1. Figure 41.5 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR RFC IGBLCW GQ D?JQC >. 5FC PCL?J NCJTGQ BP?GLQ GLRM RFC SPCRCP. ?. 5FC PCL?J NWP?KGBQ ?PC GL RFC KCBSJJ?. @. 5FC AMPRCV AMTCPQ RFC A?NQSJC. A. /CNFPMLQ ?PC GL RFC PCL?J AMPRCV. 2. Figure 41.6 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR RFC LCNFPML GQ D?JQC >. 5FC AMJJCARGLE BSAR CKNRGCQ GLRM RFC BGQR?J AMLTMJSRCB RS@SJC. ?. 5FC #MUK?LsQ A?NQSJC QSPPMSLBQ RFC EJMKCPSJSQ. @. 5FC JMMN MD )CLJC GQ @CRUCCL RFC NPMVGK?J ?LB BGQR?J AMLTMJSRCB RS@SJCQ. A. 5FC JMMN MD )CLJC CKNRGCQ GLRM RFC BGQR?J AMLTMJSRCB RS@SJC. 3. Figure 41.8 -MMN BGSPCRGAQ ?PC BPSEQ QMKCRGKCQ SQCB RM RPC?R FWNCPRCLQGML. 5FCQC BPSEQ GLFG@GR RFC PC?@QMPNRGML MD /?+ ?LB $J- GMLQ @W RFC ?QACLBGLE JGK@ MD RFC JMMN MD )CLJC. " QGBC CDDCAR GQ RF?R RFCW GLAPC?QC SPGL?RGML. 8FW BM WMS RFGLI RFGQ GQ RFC A?QC
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4. 8FCL ? BCFWBP?RCB FSK?L N?RGCLR LCCBQ RM @C EGTCL DJSGBQ GLRP?TCLMSQJW, FC MP QFC GQ EGTCL: >. U?RCP, UFGAF GQ FWNMRMLGA UGRF PCQNCAR RM @MBW DJSGBQ ?. Q?JGLC ?R ? AMLACLRP?RGML RF?R GQ GQMRMLGA UGRF PCQNCAR RM @MBW DJSGBQ @. EJSAMQC @CA?SQC GR GQ ? LML-CJCARPMJWRC A. @JMMB 5. 5FC QMBGSK GML GQ ?R RFC FGEFCQR AMLACLRP?RGML GL: >. GLRP?ACJJSJ?P DJSGB ?. CVRP?ACJJSJ?P DJSGB @. @JMMB NJ?QK? A. LMLC MD RFC ?@MTC 6. $CJJQ GL ? FWNCPRMLGA QMJSRGML RCLB RM: >. QFPGLI BSC RM U?RCP JMQQ ?. QUCJJ BSC RM U?RCP E?GL @. QR?W RFC Q?KC QGXC BSC RM U?RCP KMTGLE GLRM ?LB MSR MD RFC ACJJ ?R RFC Q?KC P?RC A. LMLC MD RFC ?@MTC 7. 5FC K?ASJ? BCLQ? GQ/?PC: >. NPCQCLR GL RFC PCL?J KCBSJJ?. ?. BCLQC RGQQSC NPCQCLR GL RFC MSRCP J?WCP MD RFC IGBLCW. @. ACJJQ NPCQCLR GL RFC %$5 ?LB AMJJCARGLE RS@SJCQ. A. NPCQCLR GL @JMMB A?NGJJ?PGCQ. 8. 5FC MQKMJ?PGRW MD @MBW DJSGBQ GQ K?GLR?GLCB ?R >>>>>>>>. >. 100 K0QK ?. 300 K0QK @. 1000 K0QK A. GR GQ LMR AMLQR?LRJW K?GLR?GLCB
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1211
9. 5FC EJ?LB JMA?RCB ?R RFC RMN MD RFC IGBLCW GQ RFC >>>>>>>> EJ?LB. >. ?BPCL?J ?. NGRSGR?PW @. RFWPMGB A. RFWKSQ 10. "ARGTC RP?LQNMPR MD ,+ GL .?JNGEFG?L RS@SJCQ CLQSPCQ RF?R: >. U?RCP DMJJMUQ ,+ RM K?IC SPGLC ?. MQKMRGA @?J?LAC GQ K?GLR?GLCB @CRUCCL U?QRC K?RRCP ?LB @MBGJW DJSGBQ @. @MRF ? ?LB @ A. LCGRFCP ? LMP @ 11. $MLRP?ARGJC T?ASMJCQ GL KGAPMMPE?LGQKQ: >. CVAJSQGTCJW NCPDMPK ?L CVAPCRMPW DSLARGML ?. A?L NCPDMPK K?LW DSLARGMLQ, MLC MD UFGAF GQ CVAPCRGML MD KCR?@MJGA U?QRCQ @. MPGEGL?RC DPMK RFC ACJJ KCK@P?LC A. @MRF @ ?LB A 12. 'J?KC ACJJQ ?PC NPGKGRGTC CVAPCRMPW MPE?LQ DMSLB GL >>>>>>>>. >. ?PRFPMNMBQ ?. ?LLCJGBQ @. K?KK?JQ A. DJ?RUMPKQ 13. #6/ GQ >>>>>>>>.
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@JMMB SPC? LGRPMECL @JMMB SPGA ?AGB LGRPMECL ?L GLBGA?RMP MD @JMMB TMJSKC ?L GLBGA?RMP MD @JMMB NPCQQSPC
14. )SK?L @CGLEQ ?AASKSJ?RC >>>>>>>> @CDMPC CVAPCRGLE LGRPMECLMSQ U?QRC. >. LGRPMECL ?. ?KKMLG? @. SPC? A. SPGA ?AGB 15. 3CLGL GQ K?BC @W >>>>>>>>. >. EP?LSJ?P ACJJQ MD RFC HSVR?EJMKCPSJ?P ?NN?P?RSQ ?. RFC IGBLCWQ @. RFC LCNFPMLQ A. "JJ MD RFC ?@MTC. 16. 1?RGCLRQ UGRF "BBGQML'Q BGQC?QC >>>>>>>>. >. ?. @. A. PCR?GL U?RCP PCR?GL Q?JRQ JMQC Q?JRQ ?LB U?RCP F?TC RMM KSAF ?JBMQRCPMLC
17. 8FGAF FMPKMLC CJGAGRQ RFC tDGEFR MP DJGEFRu PCQNMLQC >. CNGLCNFPGLC ?. KGLCP?JAMPRGAMGBQ @. ?LRG-BGSPCRGA FMPKMLC A. RFWPMVGLC
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18. 8FW GQ CVAPCRGML GKNMPR?LR GL MPBCP RM ?AFGCTC MQKMRGA @?J?LAC 19. 8FW BM CJCARPMJWRC GMLQ KMTC ?APMQQ KCK@P?LCQ @W ?ARGTC RP?LQNMPR 20. 8FW ?PC RFC JMMN MD )CLJC ?LB T?Q? PCAR? GKNMPR?LR DMP RFC DMPK?RGML MD AMLACLRP?RCB SPGLC 21. %CQAPG@C RFC QRPSARSPC MD RFC IGBLCW. 22. 8FW KGEFR QNCAG?JGXCB MPE?LQ F?TC CTMJTCB DMP CVAPCRGML MD U?QRCQ 23. &VNJ?GL RUM BGDDCPCLR CVAPCRMPW QWQRCKQ MRFCP RF?L RFC IGBLCWQ. 24. *L RCPKQ MD CTMJSRGML, UFW KGEFR RFC SPC? AWAJC F?TC CTMJTCB GL MPE?LGQKQ 25. $MKN?PC ?LB AMLRP?QR RFC DMPK?RGML MD SPC? ?LB SPGA ?AGB. 26. %CQAPG@C FMU FMPKMLCQ PCESJ?RC @JMMB NPCQQSPC, @JMMB TMJSKC, ?LB IGBLCW DSLARGML. 27. )MU BMCQ RFC PCLGL-?LEGMRCLQGL-?JBMQRCPMLC KCAF?LGQK DSLARGML 8FW GQ GR AMLRPMJJCB @W RFC IGBLCWQ
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5FC CLTGPMLKCLR AMLQGQRQ MD LSKCPMSQ pathogens, UFGAF ?PC ?ECLRQ, SQS?JJW KGAPMMPE?LGQKQ, RF?R A?SQC BGQC?QCQ GL RFCGP FMQRQ. " host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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C NFWQGA?J ?LB AFCKGA?J GKKSLC @?PPGCPQ z &VNJ?GL GKKCBG?RC ?LB GLBSACB GLL?RC GKKSLC PCQNMLQCQ z %GQASQQ L?RSP?J IGJJCP ACJJQ z %CQAPG@C K?HMP FGQRMAMKN?RG@GJGRW AJ?QQ * KMJCASJCQ z 4SKK?PGXC FMU RFC NPMRCGLQ GL ? AMKNJCKCLR QWQRCK DSLARGML RM BCQRPMW CVRP?ACJJSJ?P N?RFMECLQ 5FC GKKSLC QWQRCK AMKNPGQCQ @MRF GLL?RC ?LB ?B?NRGTC GKKSLC PCQNMLQCQ. Innate immunity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#CDMPC ?LW GKKSLC D?ARMPQ ?PC RPGEECPCB, RFC QIGL DSLARGMLQ ?Q ? AMLRGLSMSQ, GKN?QQ?@JC @?PPGCP RM NMRCLRG?JJW GLDCARGMSQ N?RFMECLQ. 1?RFMECLQ ?PC IGJJCB MP GL?ARGT?RCB ML RFC QIGL @W BCQGAA?RGML (BPWGLE MSR) ?LB @W RFC QIGLsQ ?AGBGRW. *L ?BBGRGML, @CLCDGAG?J KGAPMMPE?LGQKQ RF?R AMCVGQR ML RFC QIGL AMKNCRC UGRF GLT?BGLE N?RFMECLQ, NPCTCLRGLE GLDCARGML. 3CEGMLQ MD RFC @MBW RF?R ?PC LMR NPMRCARCB @W QIGL (QSAF ?Q RFC CWCQ ?LB KSASQ KCK@P?LCQ) F?TC ?JRCPL?RGTC KCRFMBQ MD BCDCLQC, QSAF ?Q RC?PQ ?LB KSASQ QCAPCRGMLQ RF?R RP?N ?LB PGLQC ?U?W N?RFMECLQ, ?LB AGJG? GL RFC L?Q?J N?QQ?ECQ ?LB PCQNGP?RMPW RP?AR RF?R NSQF RFC KSASQ UGRF RFC N?RFMECLQ MSR MD RFC @MBW. 5FPMSEFMSR RFC @MBW ?PC MRFCP BCDCLQCQ, QSAF ?Q RFC JMU N) MD RFC QRMK?AF (UFGAF GLFG@GRQ RFC EPMURF MD N?RFMECLQ), @JMMB NPMRCGLQ RF?R @GLB ?LB BGQPSNR @?ARCPG?J ACJJ KCK@P?LCQ, ?LB RFC NPMACQQ MD SPGL?RGML (UFGAF DJSQFCQ N?RFMECLQ DPMK RFC SPGL?PW RP?AR). %CQNGRC RFCQC @?PPGCPQ, N?RFMECLQ K?W CLRCP RFC @MBW RFPMSEF QIGL ?@P?QGMLQ MP NSLARSPCQ, MP @W AMJJCARGLE ML KSAMQ?J QSPD?ACQ GL J?PEC LSK@CPQ RF?R MTCPAMKC RFC KSASQ MP AGJG?. 4MKC N?RFMECLQ F?TC CTMJTCB QNCAGDGA KCAF?LGQKQ RF?R ?JJMU RFCK RM MTCPAMKC NFWQGA?J ?LB AFCKGA?J @?PPGCPQ. 8FCL N?RFMECLQ BM CLRCP RFC @MBW, RFC GLL?RC GKKSLC QWQRCK PCQNMLBQ UGRF GLDJ?KK?RGML, N?RFMECL CLESJDKCLR, ?LB QCAPCRGML MD GKKSLC D?ARMPQ ?LB NPMRCGLQ.
+8K?F><E -<:F>E@K@FE
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pathogen-associated molecular patterns (PAMPs) ML RFC N?RFMECLsQ QSPD?AC. 1".1Q ?PC A?P@MFWBP?RC, NMJWNCNRGBC, ?LB LSAJCGA ?AGB tQGEL?RSPCQu RF?R ?PC CVNPCQQCB @W TGPSQCQ, @?ARCPG?, ?LB N?P?QGRCQ @SR UFGAF BGDDCP DPMK KMJCASJCQ ML FMQR ACJJQ. 5FC GKKSLC QWQRCK F?Q QNCAGDGA ACJJQ, BCQAPG@CB GL Figure 42.2 ?LB QFMUL GL Figure 42.3, UGRF PCACNRMPQ RF?R PCAMELGXC RFCQC 1".1Q. " macrophage GQ ? J?PEC NF?EMAWRGA ACJJ RF?R CLESJDQ DMPCGEL N?PRGAJCQ ?LB N?RFMECLQ. .?APMNF?ECQ PCAMELGXC 1".1Q TG? AMKNJCKCLR?PW pattern recognition receptors (PRRs). 133Q ?PC KMJCASJCQ ML K?APMNF?ECQ ?LB BCLBPGRGA ACJJQ UFGAF ?PC GL AMLR?AR UGRF RFC CVRCPL?J CLTGPMLKCLR. " monocyte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
#(!04%2 42 X 4(% )--5.% 3934%-
1215
!@>LI< 42.2 4EB @E>O>@QBOFPQF@P >KA IL@>QFLK LC @BIIP FKSLISBA FK QEB FKK>QB FJJRKB PVPQBJ >OB ABP@OF?BA. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .)()
1216
#(!04%2 42 X 4(% )--5.% 3934%-
!@>LI< 42.3 #BIIP LC QEB ?ILLA FK@IRAB (1) JLKL@VQBP, (2) IVJMEL@VQBP, (3) KBRQOLMEFIP, (4) OBA ?ILLA @BIIP, >KA (5) MI>QBIBQP. .LQB QEB SBOV PFJFI>O JLOMELILDFBP LC QEB IBRHL@VQBP (1, 2, 3). (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V "OR@B 7BQWBI, (>OOV 3@E>BCBO, .#); P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
C>940/3+ +1+'8+ A,,+)9 5FC @GLBGLE MD 133Q UGRF 1".1Q RPGEECPQ RFC PCJC?QC MD AWRMIGLCQ, UFGAF QGEL?J RF?R ? N?RFMECL GQ NPCQCLR ?LB LCCBQ RM @C BCQRPMWCB ?JMLE UGRF ?LW GLDCARCB ACJJQ. " cytokine GQ ? AFCKGA?J KCQQCLECP RF?R PCESJ?RCQ ACJJ BGDDCPCLRG?RGML (DMPK ?LB DSLARGML), NPMJGDCP?RGML (NPMBSARGML), ?LB ECLC CVNPCQQGML RM ?DDCAR GKKSLC PCQNMLQCQ. "R JC?QR 40 RWNCQ MD AWRMIGLCQ CVGQR GL FSK?LQ RF?R BGDDCP GL RCPKQ MD RFC ACJJ RWNC RF?R NPMBSACQ RFCK, RFC ACJJ RWNC RF?R PCQNMLBQ RM RFCK, ?LB RFC AF?LECQ RFCW NPMBSAC. 0LC RWNC AWRMIGLC, GLRCPDCPML, GQ GJJSQRP?RCB GL Figure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interferon GQ RM GLFG@GR TGP?J PCNJGA?RGML. 5FCW ?JQM F?TC MRFCP GKNMPR?LR DSLARGMLQ, QSAF ?Q RSKMP QSPTCGJJ?LAC. *LRCPDCPMLQ UMPI @W QGEL?JGLE LCGEF@MPGLE SLGLDCARCB ACJJQ RM BCQRPMW 3/" ?LB PCBSAC NPMRCGL QWLRFCQGQ, QGEL?JGLE LCGEF@MPGLE GLDCARCB ACJJQ RM SLBCPEM ?NMNRMQGQ (NPMEP?KKCB ACJJ BC?RF), ?LB ?ARGT?RGLE GKKSLC ACJJQ. *L PCQNMLQC RM GLRCPDCPMLQ, SLGLDCARCB ACJJQ ?JRCP RFCGP ECLC CVNPCQQGML, UFGAF GLAPC?QCQ RFC ACJJQs PCQGQR?LAC RM GLDCARGML. 0LC CDDCAR MD GLRCPDCPML-GLBSACB ECLC CVNPCQQGML GQ ? QF?PNJW PCBSACB ACJJSJ?P NPMRCGL QWLRFCQGQ. 7GP?JJW GLDCARCB ACJJQ NPMBSAC KMPC TGPSQCQ @W QWLRFCQGXGLE J?PEC OS?LRGRGCQ MD TGP?J NPMRCGLQ. 5FSQ, @W PCBSAGLE NPMRCGL QWLRFCQGQ, ? ACJJ @CAMKCQ PCQGQR?LR RM TGP?J GLDCARGML.
#(!04%2 42 X 4(% )--5.% 3934%-
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!@>LI< 42.4 )KQBOCBOLKP >OB @VQLHFKBP QE>Q >OB OBIB>PBA ?V > @BII FKCB@QBA TFQE > SFORP. 2BPMLKPB LC KBFDE?LOFKD @BIIP QL FKQBOCBOLK EBIMP PQBJ QEB FKCB@QFLK.
P.'-4)>948/8 '3* I3,1'22'9/43 5FC DGPQR AWRMIGLCQ RM @C NPMBSACB ?PC NPM-GLDJ?KK?RMPW; RF?R GQ, RFCW CLAMSP?EC inflammation, RFC JMA?JGXCB PCBLCQQ, QUCJJGLE, FC?R, ?LB N?GL RF?R PCQSJR DPMK RFC KMTCKCLR MD JCSIMAWRCQ ?LB DJSGB RFPMSEF GLAPC?QGLEJW NCPKC?@JC A?NGJJ?PGCQ RM ? QGRC MD GLDCARGML. 5FC NMNSJ?RGML MD JCSIMAWRCQ RF?R ?PPGTCQ ?R ?L GLDCARGML QGRC BCNCLBQ ML RFC L?RSPC MD RFC GLDCARGLE N?RFMECL. #MRF K?APMNF?ECQ ?LB BCLBPGRGA ACJJQ CLESJD N?RFMECLQ ?LB ACJJSJ?P BC@PGQ RFPMSEF NF?EMAWRMQGQ. " neutrophil GQ ?JQM ? NF?EMAWRGA JCSIMAWRC RF?R CLESJDQ ?LB BGECQRQ N?RFMECLQ. /CSRPMNFGJQ, QFMUL GL Figure 42.3, ?PC RFC KMQR ?@SLB?LR JCSIMAWRCQ MD RFC GKKSLC QWQRCK. /CSRPMNFGJQ F?TC ? LSAJCSQ UGRF RUM RM DGTC JM@CQ, ?LB RFCW AMLR?GL MPE?LCJJCQ, A?JJCB JWQMQMKCQ, RF?R BGECQR CLESJDCB N?RFMECLQ. "L eosinophil GQ ? JCSIMAWRC RF?R UMPIQ UGRF MRFCP CMQGLMNFGJQ RM QSPPMSLB ? N?P?QGRC; GR GQ GLTMJTCB GL RFC ?JJCPEGA PCQNMLQC ?LB GL NPMRCARGML ?E?GLQR FCJKGLRFCQ (N?P?QGRGA UMPKQ). /CSRPMNFGJQ ?LB CMQGLMNFGJQ ?PC N?PRGASJ?PJW GKNMPR?LR JCSIMAWRCQ RF?R CLESJD J?PEC N?RFMECLQ, QSAF ?Q @?ARCPG? ?LB DSLEG. " mast cell GQ ? JCSIMAWRC RF?R NPMBSACQ GLDJ?KK?RMPW KMJCASJCQ, QSAF ?Q FGQR?KGLC, GL PCQNMLQC RM J?PEC N?RFMECLQ. " basophil GQ ? JCSIMAWRC RF?R, JGIC ? LCSRPMNFGJ, PCJC?QCQ AFCKGA?JQ RM QRGKSJ?RC RFC GLDJ?KK?RMPW PCQNMLQC ?Q GJJSQRP?RCB GL Figure 42.5. #?QMNFGJQ ?PC ?JQM GLTMJTCB GL ?JJCPEW ?LB FWNCPQCLQGRGTGRW PCQNMLQCQ ?LB GLBSAC QNCAGDGA RWNCQ MD GLDJ?KK?RMPW PCQNMLQCQ. &MQGLMNFGJQ ?LB @?QMNFGJQ NPMBSAC ?BBGRGML?J GLDJ?KK?RMPW KCBG?RMPQ RM PCAPSGR KMPC JCSIMAWRCQ. " FWNCPQCLQGRGTC GKKSLC PCQNMLQC RM F?PKJCQQ ?LRGECLQ, QSAF ?Q GL NMJJCL, MDRCL GLTMJTCQ RFC PCJC?QC MD FGQR?KGLC @W @?QMNFGJQ ?LB K?QR ACJJQ.
!@>LI< 42.5 )K OBPMLKPB QL > @RQ, J>PQ @BIIP PB@OBQB EFPQ>JFKBP QE>Q @>RPB KB>O?V @>MFII>OFBP QL AFI>QB. .BRQOLMEFIP >KA JLKL@VQBP IB>SB QEB @>MFII>OFBP. -LKL@VQBP J>QROB FKQL J>@OLME>DBP. .BRQOLMEFIP, ABKAOFQF@ @BIIP >KA J>@OLME>DBP OBIB>PB @EBJF@>IP QL PQFJRI>QB QEB FKCI>JJ>QLOV OBPMLKPB. .BRQOLMEFIP >KA J>@OLME>DBP >IPL @LKPRJB FKS>AFKD ?>@QBOF> ?V ME>DL@VQLPFP.
1218
#(!04%2 42 X 4(% )--5.% 3934%-
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
8?RAF RFGQ 23-QCAMLB QRMN-KMRGML video (http://openstaxcollege.org/l/conidia) QFMUGLE ? LCSRPMNFGJ RF?R QC?PAFCQ DMP ?LB CLESJDQ DSLESQ QNMPCQ BSPGLE ?L CJ?NQCB RGKC MD ?@MSR 79 KGLSRCQ.
)8KLI8C &@CC<I C<CCJ

Lymphocytes ?PC JCSIMAWRCQ RF?R ?PC FGQRMJMEGA?JJW GBCLRGDG?@JC @W RFCGP J?PEC, B?PIJW QR?GLGLE LSAJCG; RFCW ?PC QK?JJ ACJJQ UGRF TCPW JGRRJC AWRMNJ?QK, ?Q QFMUL GL Figure 42.6. *LDCARCB ACJJQ ?PC GBCLRGDGCB ?LB BCQRPMWCB @W natural killer (NK) cells, JWKNFMAWRCQ RF?R A?L IGJJ ACJJQ GLDCARCB UGRF TGPSQCQ MP RSKMP ACJJQ (?@LMPK?J ACJJQ RF?R SLAMLRPMJJ?@JW BGTGBC ?LB GLT?BC MRFCP RGQQSC). 5 ACJJQ ?LB # ACJJQ MD RFC ?B?NRGTC GKKSLC QWQRCK ?JQM ?PC AJ?QQGDGCB ?Q JWKNFMAWRCQ. T cells ?PC JWKNFMAWRCQ RF?R K?RSPC GL RFC RFWKSQ EJ?LB, ?LB B cells ?PC JWKNFMAWRCQ RF?R K?RSPC GL RFC @MLC K?PPMU. /, ACJJQ GBCLRGDW GLRP?ACJJSJ?P GLDCARGMLQ, CQNCAG?JJW DPMK TGPSQCQ, @W RFC ?JRCPCB CVNPCQQGML MD major histocompatibility class (MHC) I molecules ML RFC QSPD?AC MD GLDCARCB ACJJQ. .)$ * KMJCASJCQ ?PC NPMRCGLQ ML RFC QSPD?ACQ MD ?JJ LSAJC?RCB ACJJQ, RFSQ RFCW ?PC QA?PAC ML PCB @JMMB ACJJQ ?LB NJ?RCJCRQ UFGAF ?PC LML-LSAJC?RCB. 5FC DSLARGML MD .)$ * KMJCASJCQ GQ RM BGQNJ?W DP?EKCLRQ MD NPMRCGLQ DPMK RFC GLDCARGMSQ ?ECLRQ UGRFGL RFC ACJJ RM 5-ACJJQ; FC?JRFW ACJJQ UGJJ @C GELMPCB, UFGJC tLML-QCJDu MP DMPCGEL NPMRCGLQ UGJJ @C ?RR?AICB @W RFC GKKSLC QWQRCK. .)$ ** KMJCASJCQ ?PC DMSLB K?GLJW ML ACJJQ AMLR?GLGLE ?LRGECLQ (tLML-QCJD NPMRCGLQu) ?LB ML JWKNFMAWRCQ. MHC II molecules GLRCP?AR UGRF FCJNCP 5-ACJJQ RM RPGEECP RFC ?NNPMNPG?RC GKKSLC PCQNMLQC, UFGAF K?W GLAJSBC RFC GLDJ?KK?RMPW PCQNMLQC.
!@>LI< 42.6 ,VJMEL@VQBP, PR@E >P .+ @BIIP, >OB @E>O>@QBOFWBA ?V QEBFO I>ODB KR@IBF QE>Q >@QFSBIV >?PLO? 7OFDEQ PQ>FK >KA QEBOBCLOB >MMB>O A>OH @LILOBA RKABO > JF@OLP@LMB.
"L GLDCARCB ACJJ (MP ? RSKMP ACJJ) GQ SQS?JJW GLA?N?@JC MD QWLRFCQGXGLE ?LB BGQNJ?WGLE .)$ * KMJCASJCQ ?NNPMNPG?RCJW. 5FC KCR?@MJGA PCQMSPACQ MD ACJJQ GLDCARCB @W QMKC TGPSQCQ NPMBSAC NPMRCGLQ RF?R GLRCPDCPC UGRF .)$ * NPMACQQGLE ?LB/MP RP?DDGAIGLE RM RFC ACJJ QSPD?AC. 5FC PCBSACB .)$ * ML FMQR ACJJQ T?PGCQ DPMK TGPSQ RM TGPSQ ?LB PCQSJRQ DPMK ?ARGTC GLFG@GRMPQ @CGLE NPMBSACB @W RFC TGPSQCQ. 5FGQ NPMACQQ A?L BCNJCRC FMQR .)$ * KMJCASJCQ ML RFC ACJJ QSPD?AC, UFGAF /, ACJJQ BCRCAR ?Q tSLFC?JRFWu MP t?@LMPK?Ju UFGJC QC?PAFGLE DMP ACJJSJ?P .)$ * KMJCASJCQ. 4GKGJ?PJW, RFC BP?K?RGA?JJW ?JRCPCB ECLC
#(!04%2 42 X 4(% )--5.% 3934%-
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CVNPCQQGML MD RSKMP ACJJQ JC?BQ RM CVNPCQQGML MD CVRPCKCJW BCDMPKCB MP ?@QCLR .)$ * KMJCASJCQ RF?R ?JQM QGEL?J tSLFC?JRFWu MP t?@LMPK?J.u /, ACJJQ ?PC ?JU?WQ ?ARGTC; ?L GLRCP?ARGML UGRF LMPK?J, GLR?AR .)$ * KMJCASJCQ ML ? FC?JRFW ACJJ BGQ?@JCQ RFC IGJJGLE QCOSCLAC, ?LB RFC /, ACJJ KMTCQ ML. "DRCP RFC /, ACJJ BCRCARQ ?L GLDCARCB MP RSKMP ACJJ, GRQ AWRMNJ?QK QCAPCRCQ EP?LSJCQ AMKNPGQCB MD perforin, ? BCQRPSARGTC NPMRCGL RF?R APC?RCQ ? NMPC GL RFC R?PECR ACJJ. (P?LXWKCQ ?PC PCJC?QCB ?JMLE UGRF RFC NCPDMPGL GL RFC GKKSLMJMEGA?J QWL?NQC. " granzyme GQ ? NPMRC?QC RF?R BGECQRQ ACJJSJ?P NPMRCGLQ ?LB GLBSACQ RFC R?PECR ACJJ RM SLBCPEM NPMEP?KKCB ACJJ BC?RF, MP ?NMNRMQGQ. 1F?EMAWRGA ACJJQ RFCL BGECQR RFC ACJJ BC@PGQ JCDR @CFGLB. /, ACJJQ ?PC AMLQR?LRJW N?RPMJJGLE RFC @MBW ?LB ?PC ?L CDDCARGTC KCAF?LGQK DMP AMLRPMJJGLE NMRCLRG?J GLDCARGMLQ ?LB NPCTCLRGLE A?LACP NPMEPCQQGML.
CFDGC<D<EK
"L ?PP?W MD ?NNPMVGK?RCJW 20 RWNCQ MD QMJS@JC NPMRCGLQ, A?JJCB ? complement system, DSLARGMLQ RM BCQRPMW CVRP?ACJJSJ?P N?RFMECLQ. $CJJQ MD RFC JGTCP ?LB K?APMNF?ECQ QWLRFCQGXC AMKNJCKCLR NPMRCGLQ AMLRGLSMSQJW; RFCQC NPMRCGLQ ?PC ?@SLB?LR GL RFC @JMMB QCPSK ?LB ?PC A?N?@JC MD PCQNMLBGLE GKKCBG?RCJW RM GLDCARGLE KGAPMMPE?LGQKQ. 5FC AMKNJCKCLR QWQRCK GQ QM L?KCB @CA?SQC GR GQ AMKNJCKCLR?PW RM RFC ?LRG@MBW PCQNMLQC MD RFC ?B?NRGTC GKKSLC QWQRCK. $MKNJCKCLR NPMRCGLQ @GLB RM RFC QSPD?ACQ MD KGAPMMPE?LGQKQ ?LB ?PC N?PRGASJ?PJW ?RRP?ARCB RM N?RFMECLQ RF?R ?PC ?JPC?BW @MSLB @W ?LRG@MBGCQ. #GLBGLE MD AMKNJCKCLR NPMRCGLQ MAASPQ GL ? QNCAGDGA ?LB FGEFJW PCESJ?RCB QCOSCLAC, UGRF C?AF QSAACQQGTC NPMRCGL @CGLE ?ARGT?RCB @W AJC?T?EC ?LB/MP QRPSARSP?J AF?LECQ GLBSACB SNML @GLBGLE MD RFC NPCACBGLE NPMRCGL(Q). "DRCP RFC DGPQR DCU AMKNJCKCLR NPMRCGLQ @GLB, ? A?QA?BC MD QCOSCLRG?J @GLBGLE CTCLRQ DMJJMUQ GL UFGAF RFC N?RFMECL P?NGBJW @CAMKCQ AM?RCB GL AMKNJCKCLR NPMRCGLQ. $MKNJCKCLR NPMRCGLQ NCPDMPK QCTCP?J DSLARGMLQ. 5FC NPMRCGLQ QCPTC ?Q ? K?PICP RM GLBGA?RC RFC NPCQCLAC MD ? N?RFMECL RM NF?EMAWRGA ACJJQ, QSAF ?Q K?APMNF?ECQ ?LB # ACJJQ, ?LB CLF?LAC CLESJDKCLR; RFGQ NPMACQQ GQ A?JJCB opsonization. $CPR?GL AMKNJCKCLR NPMRCGLQ A?L AMK@GLC RM DMPK ?RR?AI AMKNJCVCQ RF?R MNCL NMPCQ GL KGAPM@G?J ACJJ KCK@P?LCQ. 5FCQC QRPSARSPCQ BCQRPMW N?RFMECLQ @W A?SQGLE RFCGP AMLRCLRQ RM JC?I, ?Q GJJSQRP?RCB GL Figure 42.7.
1220
#(!04%2 42 X 4(% )--5.% 3934%-
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
42.2 R A;8GK@M< $DDLE< -<JGFEJ<

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL ?B?NRGTC GKKSLGRW z $MKN?PC ?LB AMLRP?QR ?B?NRGTC ?LB GLL?RC GKKSLGRW z %CQAPG@C ACJJ-KCBG?RCB GKKSLC PCQNMLQC ?LB FSKMP?J GKKSLC PCQNMLQC z %CQAPG@C GKKSLC RMJCP?LAC
#(!04%2 42 X 4(% )--5.% 3934%-
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5FC ?B?NRGTC, MP ?AOSGPCB, GKKSLC PCQNMLQC R?ICQ B?WQ MP CTCL UCCIQ RM @CAMKC CQR?@JGQFCBqKSAF JMLECP RF?L RFC GLL?RC PCQNMLQC; FMUCTCP, ?B?NRGTC GKKSLGRW GQ KMPC QNCAGDGA RM N?RFMECLQ ?LB F?Q KCKMPW. Adaptive immunity GQ ?L GKKSLGRW RF?R MAASPQ ?DRCP CVNMQSPC RM ?L ?LRGECL CGRFCP DPMK ? N?RFMECL MP ? T?AAGL?RGML. 5FGQ N?PR MD RFC GKKSLC QWQRCK GQ ?ARGT?RCB UFCL RFC GLL?RC GKKSLC PCQNMLQC GQ GLQSDDGAGCLR RM AMLRPMJ ?L GLDCARGML. *L D?AR, UGRFMSR GLDMPK?RGML DPMK RFC GLL?RC GKKSLC QWQRCK, RFC ?B?NRGTC PCQNMLQC AMSJB LMR @C KM@GJGXCB. 5FCPC ?PC RUM RWNCQ MD ?B?NRGTC PCQNMLQCQ: RFC cellmediated immune response, UFGAF GQ A?PPGCB MSR @W 5 ACJJQ, ?LB RFC humoral immune response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6LJGIC /, ACJJQ MD RFC GLL?RC GKKSLC QWQRCK, # ACJJQ (# JWKNFMAWRCQ) ?PC ? RWNC MD UFGRC @JMMB ACJJ RF?R EGTCQ PGQC RM ?LRG@MBGCQ, UFCPC?Q 5 ACJJQ (5 JWKNFMAWRCQ) ?PC ? RWNC MD UFGRC @JMMB ACJJ RF?R NJ?WQ ?L GKNMPR?LR PMJC GL RFC GKKSLC PCQNMLQC. 5 ACJJQ ?PC ? ICW AMKNMLCLR GL RFC ACJJ-KCBG?RCB PCQNMLQCq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antigen GQ ? DMPCGEL MP tLML-QCJDu K?APMKMJCASJC RF?R PC?ARQ UGRF ACJJQ MD RFC GKKSLC QWQRCK. /MR ?JJ ?LRGECLQ UGJJ NPMTMIC ? PCQNMLQC. 'MP GLQR?LAC, GLBGTGBS?JQ NPMBSAC GLLSKCP?@JC tQCJDu ?LRGECLQ ?LB ?PC AMLQR?LRJW CVNMQCB RM F?PKJCQQ DMPCGEL ?LRGECLQ, QSAF ?Q DMMB NPMRCGLQ, NMJJCL, MP BSQR AMKNMLCLRQ. 5FC QSNNPCQQGML MD GKKSLC PCQNMLQCQ RM F?PKJCQQ K?APMKMJCASJCQ GQ FGEFJW PCESJ?RCB ?LB RWNGA?JJW NPCTCLRQ NPMACQQCQ RF?R AMSJB @C B?K?EGLE RM RFC FMQR, ILMUL ?Q RMJCP?LAC. 5FC GLL?RC GKKSLC QWQRCK AMLR?GLQ ACJJQ RF?R BCRCAR NMRCLRG?JJW F?PKDSJ ?LRGECLQ, ?LB RFCL GLDMPK RFC ?B?NRGTC GKKSLC PCQNMLQC ?@MSR RFC NPCQCLAC MD RFCQC ?LRGECLQ. "L antigen-presenting cell (APC) GQ ?L GKKSLC ACJJ RF?R BCRCARQ, CLESJDQ, ?LB GLDMPKQ RFC ?B?NRGTC GKKSLC PCQNMLQC ?@MSR ?L GLDCARGML. 8FCL ? N?RFMECL GQ BCRCARCB, RFCQC "1$Q UGJJ NF?EMAWRMQC RFC N?RFMECL ?LB BGECQR GR RM DMPK K?LW BGDDCPCLR DP?EKCLRQ MD RFC ?LRGECL. "LRGECL DP?EKCLRQ UGJJ RFCL @C RP?LQNMPRCB RM RFC QSPD?AC MD RFC "1$, UFCPC RFCW UGJJ QCPTC ?Q ?L GLBGA?RMP RM MRFCP GKKSLC ACJJQ. Dendritic cells ?PC GKKSLC ACJJQ RF?R NPMACQQ ?LRGECL K?RCPG?J; RFCW ?PC NPCQCLR GL RFC QIGL (-?LECPF?LQ ACJJQ) ?LB RFC JGLGLE MD RFC LMQC, JSLEQ, QRMK?AF, ?LB GLRCQRGLCQ. 4MKCRGKCQ ? BCLBPGRGA ACJJ NPCQCLRQ ML RFC QSPD?AC MD MRFCP ACJJQ RM GLBSAC ?L GKKSLC PCQNMLQC, RFSQ DSLARGMLGLE ?Q ?L ?LRGECL-NPCQCLRGLE ACJJ. .?APMNF?ECQ ?JQM DSLARGML ?Q "1$Q. #CDMPC ?ARGT?RGML ?LB BGDDCPCLRG?RGML, # ACJJQ A?L ?JQM DSLARGML ?Q "1$Q. "DRCP NF?EMAWRMQGQ @W "1$Q, RFC NF?EMAWRGA TCQGAJC DSQCQ UGRF ?L GLRP?ACJJSJ?P JWQMQMKC DMPKGLE NF?EMJWQMQMKC. 8GRFGL RFC NF?EMJWQMQMKC, RFC AMKNMLCLRQ ?PC @PMICL BMUL GLRM DP?EKCLRQ; RFC DP?EKCLRQ ?PC RFCL JM?BCB MLRM .)$ AJ?QQ * MP .)$ AJ?QQ ** KMJCASJCQ ?LB ?PC RP?LQNMPRCB RM RFC ACJJ QSPD?AC DMP ?LRGECL NPCQCLR?RGML, ?Q GJJSQRP?RCB GL Figure 42.8. /MRC RF?R 5 JWKNFMAWRCQ A?LLMR NPMNCPJW PCQNMLB RM RFC ?LRGECL SLJCQQ GR GQ NPMACQQCB ?LB CK@CBBCB GL ?L .)$ ** KMJCASJC. "1$Q CVNPCQQ .)$ ML RFCGP QSPD?ACQ, ?LB UFCL AMK@GLCB UGRF ? DMPCGEL ?LRGECL, RFCQC AMKNJCVCQ QGEL?J ? tLMLQCJDu GLT?BCP. 0LAC RFC DP?EKCLR MD ?LRGECL GQ CK@CBBCB GL RFC .)$ ** KMJCASJC, RFC GKKSLC ACJJ A?L PCQNMLB. )CJNCP 5- ACJJQ ?PC MLC MD RFC K?GL JWKNFMAWRCQ RF?R PCQNMLB RM ?LRGECL-NPCQCLRGLE ACJJQ. 3CA?JJ RF?R ?JJ MRFCP LSAJC?RCB ACJJQ MD RFC @MBW CVNPCQQCB .)$ * KMJCASJCQ, UFGAF QGEL?J tFC?JRFWu MP tLMPK?J.u
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5FGQ animation (http://openstaxcollege.org/l/immune_system) DPMK 3MAICDCJJCP 6LGTCPQGRW QFMUQ FMU BCLBPGRGA ACJJQ ?AR ?Q QCLRGLCJQ GL RFC @MBW'Q GKKSLC QWQRCK.
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-WKNFMAWRCQ GL FSK?L AGPASJ?RGLE @JMMB ?PC ?NNPMVGK?RCJW 80 RM 90 NCPACLR 5 ACJJQ, QFMUL GL Figure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tQCJDu ?LRGECLQ.
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epitopes, NPCQCLRCB @W "1$Q, GJJSQRP?RCB GL Figure 42.10. /MRC RF?R PCAMELGRGML MAASPQ ?R ? QNCAGDGA CNGRMNC P?RFCP RF?L ML RFC CLRGPC ?LRGECL; DMP RFGQ PC?QML, CNGRMNCQ ?PC ILMUL ?Q t?LRGECLGA BCRCPKGL?LRQ.u *L RFC ?@QCLAC MD GLDMPK?RGML DPMK "1$Q, 5 ?LB # ACJJQ PCK?GL GL?ARGTC, MP L?bTC, ?LB ?PC SL?@JC RM NPCN?PC ?L GKKSLC PCQNMLQC. 5FC PCOSGPCKCLR DMP GLDMPK?RGML DPMK RFC "1$Q MD GLL?RC GKKSLGRW RM RPGEECP # ACJJ MP 5 ACJJ ?ARGT?RGML GJJSQRP?RCQ RFC CQQCLRG?J L?RSPC MD RFC GLL?RC GKKSLC PCQNMLQC RM RFC DSLARGMLGLE MD RFC CLRGPC GKKSLC QWQRCK.
!@>LI< 42.10 !K >KQFDBK FP > J>@OLJLIB@RIB QE>Q OB>@QP TFQE @LJMLKBKQP LC QEB FJJRKB PVPQBJ. ! DFSBK >KQFDBK J>V @LKQ>FK PBSBO>I JLQFCP QE>Q >OB OB@LDKFWBA ?V FJJRKB @BIIP. %>@E JLQFC FP >K BMFQLMB. )K QEFP CFDROB, QEB BKQFOB PQOR@QROB FP >K >KQFDBK, >KA QEB LO>KDB, P>IJLK >KA DOBBK @LJMLKBKQP MOLGB@QFKD COLJ FQ OBMOBPBKQ MLQBKQF>I BMFQLMBP.
/?bTC 5 ACJJQ A?L CVNPCQQ MLC MD RUM BGDDCPCLR KMJCASJCQ, $%4 MP $%8, ML RFCGP QSPD?AC, ?Q QFMUL GL Figure 42.11, ?LB ?PC ?AAMPBGLEJW AJ?QQGDGCB ?Q $%4+ MP $%8+ ACJJQ. 5FCQC KMJCASJCQ ?PC GKNMPR?LR @CA?SQC RFCW PCESJ?RC FMU ? 5 ACJJ UGJJ GLRCP?AR UGRF ?LB PCQNMLB RM ?L "1$. /?bTC $%4+ ACJJQ @GLB "1$Q TG? RFCGP ?LRGECL-CK@CBBCB .)$ ** KMJCASJCQ ?LB ?PC QRGKSJ?RCB RM @CAMKC helper T (TH) lymphocytes, ACJJQ RF?R EM ML RM QRGKSJ?RC # ACJJQ (MP AWRMRMVGA 5 ACJJQ) BGPCARJW MP QCAPCRC AWRMIGLCQ RM GLDMPK KMPC ?LB T?PGMSQ R?PECR ACJJQ ?@MSR RFC N?RFMECLGA RFPC?R. *L AMLRP?QR, $%8+ ACJJQ CLE?EC ?LRGECLCK@CBBCB .)$ * KMJCASJCQ ML "1$Q ?LB ?PC QRGKSJ?RCB RM @CAMKC cytotoxic T lymphocytes (CTLs), UFGAF BGPCARJW IGJJ GLDCARCB ACJJQ @W ?NMNRMQGQ ?LB CKGR AWRMIGLCQ RM ?KNJGDW RFC GKKSLC PCQNMLQC. 5FC RUM NMNSJ?RGMLQ MD 5 ACJJQ F?TC BGDDCPCLR KCAF?LGQKQ MD GKKSLC NPMRCARGML, @SR @MRF @GLB .)$ KMJCASJCQ TG? RFCGP ?LRGECL PCACNRMPQ A?JJCB 5 ACJJ PCACNRMPQ (5$3Q). 5FC $%4 MP $%8 QSPD?AC KMJCASJCQ BGDDCPCLRG?RC UFCRFCP RFC 5$3 UGJJ CLE?EC ?L .)$ ** MP ?L .)$ * KMJCASJC. #CA?SQC RFCW ?QQGQR GL @GLBGLE QNCAGDGAGRW, RFC $%4 ?LB $%8 KMJCASJCQ ?PC BCQAPG@CB ?Q AMPCACNRMPQ.
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!@>LI< 42.11 .>eSB #$4+ 4 @BIIP BKD>DB -(# )) JLIB@RIBP LK >KQFDBK-MOBPBKQFKD @BIIP (!0#P) >KA ?B@LJB >@QFS>QBA. #ILKBP LC QEB >@QFS>QBA EBIMBO 4 @BII, FK QROK, >@QFS>QB " @BIIP >KA #$8+ 4 @BIIP, TEF@E ?B@LJB @VQLQLUF@ 4 @BIIP. #VQLQLUF@ 4 @BIIP HFII FKCB@QBA @BIIP.
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!@>LI< 42.13 " @BII OB@BMQLOP >OB BJ?BAABA FK QEB JBJ?O>KBP LC " @BIIP >KA ?FKA > S>OFBQV LC >KQFDBKP QEOLRDE QEBFO S>OF>?IB OBDFLKP. 4EB PFDK>I QO>KPAR@QFLK OBDFLK QO>KPCBOP QEB PFDK>I FKQL QEB @BII.
5 ?LB # ACJJQ BGDDCP GL MLC DSLB?KCLR?J U?W: UFCPC?Q 5 ACJJQ @GLB ?LRGECLQ RF?R F?TC @CCL BGECQRCB ?LB CK@CBBCB GL .)$ KMJCASJCQ @W "1$Q, # ACJJQ DSLARGML ?Q "1$Q RF?R @GLB GLR?AR ?LRGECLQ RF?R F?TC LMR @CCL NPMACQQCB. "JRFMSEF 5 ?LB # ACJJQ @MRF PC?AR UGRF KMJCASJCQ RF?R ?PC RCPKCB t?LRGECLQ,u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bTC $%8+ 5 ACJJQ RF?R CVNPCQQ AMKNJCKCLR?PW 5$3Q, RFC $%8+ 5 ACJJQ @CAMKC ?ARGT?RCB RM NPMJGDCP?RC ?AAMPBGLE RM AJML?J QCJCARGML. 5FCQC PCQSJRGLE $5-Q RFCL GBCLRGDW LML"1$Q BGQNJ?WGLE RFC Q?KC .)$ *-CK@CBBCB ?LRGECLQ (DMP CV?KNJC, TGP?J NPMRCGLQ)q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igure 42.14. *D RFC /, ACJJ A?LLMR GBCLRGDW RFC tKGQQGLE QCJDu N?RRCPL MD BMUL-PCESJ?RCB .)$ * KMJCASJCQ, RFCL RFC $5- A?L GBCLRGDW GR @W RFC AMKNJCV MD .)$ * UGRF DMPCGEL ?LRGECLQ, UFGAF QGEL?JQ t?JRCPCB QCJD.u 4GKGJ?PJW, GD RFC $5- A?LLMR BCRCAR ?LRGECL-CK@CBBCB .)$ * @CA?SQC RFC PCACNRMPQ ?PC BCNJCRCB DPMK RFC ACJJ QSPD?AC, /, ACJJQ UGJJ BCQRPMW RFC ACJJ GLQRC?B. $5-Q ?JQM CKGR AWRMIGLCQ, QSAF ?Q GLRCPDCPMLQ, RF?R ?JRCP QSPD?AC NPMRCGL CVNPCQQGML GL MRFCP GLDCARCB ACJJQ, QSAF RF?R RFC GLDCARCB ACJJQ A?L @C C?QGJW GBCLRGDGCB ?LB BCQRPMWCB. .MPCMTCP, RFCQC GLRCPDCPMLQ A?L ?JQM NPCTCLR TGP?JJW GLDCARCB ACJJQ DPMK PCJC?QGLE TGPSQ N?PRGAJCQ.
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!@>LI< 42.14 .>QRO>I HFIIBO (.+) @BIIP OB@LDKFWB QEB -(# ) OB@BMQLO LK EB>IQEV @BIIP. )C -(# ) FP >?PBKQ, QEB @BII FP IVPBA.
">PBA LK TE>Q VLR HKLT >?LRQ -(# OB@BMQLOP, TEV AL VLR QEFKH >K LOD>K QO>KPMI>KQBA COLJ >K FK@LJM>QF?IB ALKLO QL > OB@FMFBKQ TFII ?B OBGB@QBA 1J?QK? ACJJQ ?LB $5-Q ?PC AMJJCARGTCJW A?JJCB effector cells: RFCW PCNPCQCLR BGDDCPCLRG?RCB TCPQGMLQ MD RFCGP L?b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ucosa-associated lymphoid tissue (MALT), GJJSQRP?RCB GL Figure 42.15, GQ ? AMJJCARGML MD JWKNF?RGA RGQQSC RF?R AMK@GLCQ UGRF CNGRFCJG?J RGQQSC JGLGLE RFC KSAMQ? RFPMSEFMSR RFC @MBW. 5FGQ RGQQSC DSLARGMLQ ?Q RFC GKKSLC @?PPGCP ?LB PCQNMLQC GL ?PC?Q MD RFC @MBW UGRF BGPCAR AMLR?AR RM RFC CVRCPL?J CLTGPMLKCLR. 5FC QWQRCKGA ?LB KSAMQ?J GKKSLC QWQRCKQ SQC K?LW MD RFC Q?KC ACJJ RWNCQ. 'MPCGEL N?PRGAJCQ RF?R K?IC RFCGP U?W RM ."-5 ?PC R?ICL SN @W ?@QMPNRGTC CNGRFCJG?J ACJJQ A?JJCB . ACJJQ ?LB BCJGTCPCB RM "1$Q JMA?RCB BGPCARJW @CJMU RFC KSAMQ?J RGQQSC. . ACJJQ DSLARGML GL RFC RP?LQNMPR BCQAPG@CB, ?LB ?PC JMA?RCB GL RFC 1CWCPsQ N?RAF, ? JWKNFMGB LMBSJC. "1$Q MD RFC KSAMQ?J GKKSLC QWQRCK ?PC NPGK?PGJW BCLBPGRGA ACJJQ, UGRF # ACJJQ ?LB K?APMNF?ECQ F?TGLE KGLMP PMJCQ. 1PMACQQCB ?LRGECLQ BGQNJ?WCB ML "1$Q ?PC BCRCARCB @W 5 ACJJQ GL RFC ."-5 ?LB ?R T?PGMSQ KSAMQ?J GLBSARGML QGRCQ, QSAF ?Q RFC RMLQGJQ, ?BCLMGBQ, ?NNCLBGV, MP RFC KCQCLRCPGA JWKNF LMBCQ MD RFC GLRCQRGLC. "ARGT?RCB 5 ACJJQ RFCL KGEP?RC RFPMSEF RFC JWKNF?RGA QWQRCK ?LB GLRM RFC AGPASJ?RMPW QWQRCK RM KSAMQ?J QGRCQ MD GLDCARGML.
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#(!04%2 42 X 4(% )--5.% 3934%-
!@>LI< 42.15 4EB QLMLILDV >KA CRK@QFLK LC FKQBPQFK>I -!,4 FP PELTK. 0>QELDBKP >OB Q>HBK RM ?V @BIIP FK QEB FKQBPQFK>I BMFQEBIFRJ >KA BU@OBQBA FKQL > ML@HBQ CLOJBA ?V QEB FKKBO PROC>@B LC QEB @BII. 4EB ML@HBQ @LKQ>FKP >KQFDBK-MOBPBKQFKD @BIIP PR@E >P ABKAOFQF@ @BIIP, TEF@E BKDRIC QEB >KQFDBKP, QEBK MOBPBKQ QEBJ TFQE -(# )) JLIB@RIBP LK QEB @BII PROC>@B. 4EB ABKAOFQF@ @BIIP JFDO>QB QL >K RKABOIVFKD QFPPRB @>IIBA > 0BVBOwP M>Q@E. !KQFDBK-MOBPBKQFKD @BIIP, 4 @BIIP, >KA " @BIIP >DDOBD>QB TFQEFK QEB 0BVBOwP M>Q@E, CLOJFKD LOD>KFWBA IVJMELFA CLIIF@IBP. 4EBOB, PLJB 4 @BIIP >KA " @BIIP >OB >@QFS>QBA. /QEBO >KQFDBK-IL>ABA ABKAOFQF@ @BIIP JFDO>QB QEOLRDE QEB IVJME>QF@ PVPQBJ TEBOB QEBV >@QFS>QB " @BIIP, 4 @BIIP, >KA MI>PJ> @BIIP FK QEB IVJME KLABP. 4EB >@QFS>QBA @BIIP QEBK OBQROK QL -!,4 QFPPRB BCCB@QLO PFQBP. )D! >KA LQEBO >KQF?LAFBP >OB PB@OBQBA FKQL QEB FKQBPQFK>I IRJBK.
."-5 GQ ? APSAG?J AMKNMLCLR MD ? DSLARGML?J GKKSLC QWQRCK @CA?SQC KSAMQ?J QSPD?ACQ, QSAF ?Q RFC L?Q?J N?QQ?ECQ, ?PC RFC DGPQR RGQQSCQ MLRM UFGAF GLF?JCB MP GLECQRCB N?RFMECLQ ?PC BCNMQGRCB. 5FC KSAMQ?J RGQQSC GLAJSBCQ RFC KMSRF, NF?PWLV, ?LB CQMNF?ESQ, ?LB RFC E?QRPMGLRCQRGL?J, PCQNGP?RMPW, ?LB SPMECLGR?J RP?ARQ. 5FC GKKSLC QWQRCK F?Q RM @C PCESJ?RCB RM NPCTCLR U?QRCDSJ, SLLCACQQ?PW PCQNMLQCQ RM F?PKJCQQ QS@QR?LACQ, ?LB KMPC GKNMPR?LRJW QM RF?R GR BMCQ LMR ?RR?AI tQCJD.u 5FC ?AOSGPCB ?@GJGRW RM NPCTCLR ?L SLLCACQQ?PW MP F?PKDSJ GKKSLC PCQNMLQC RM ? BCRCARCB DMPCGEL QS@QR?LAC ILMUL LMR RM A?SQC BGQC?QC GQ BCQAPG@CB ?Q immune tolerance. *KKSLC RMJCP?LAC GQ APSAG?J DMP K?GLR?GLGLE KSAMQ?J FMKCMQR?QGQ EGTCL RFC RPCKCLBMSQ LSK@CP MD DMPCGEL QS@QR?LACQ (QSAF ?Q DMMB NPMRCGLQ) RF?R "1$Q MD RFC MP?J A?TGRW, NF?PWLV, ?LB E?QRPMGLRCQRGL?J KSAMQ? CLAMSLRCP. *KKSLC RMJCP?LAC GQ @PMSEFR ?@MSR @W QNCAG?JGXCB "1$Q GL RFC JGTCP, JWKNF LMBCQ, QK?JJ GLRCQRGLC, ?LB JSLE RF?R NPCQCLR F?PKJCQQ ?LRGECLQ RM ?L CVACNRGML?JJW BGTCPQC NMNSJ?RGML MD regulatory T (Treg) cells, QNCAG?JGXCB JWKNFMAWRCQ RF?R QSNNPCQQ JMA?J GLDJ?KK?RGML ?LB GLFG@GR RFC QCAPCRGML MD QRGKSJ?RMPW GKKSLC D?ARMPQ. 5FC AMK@GLCB PCQSJR MD 5PCE ACJJQ GQ RM NPCTCLR GKKSLMJMEGA ?ARGT?RGML ?LB GLDJ?KK?RGML GL SLBCQGPCB RGQQSC AMKN?PRKCLRQ ?LB RM ?JJMU RFC GKKSLC QWQRCK RM DMASQ ML N?RFMECLQ GLQRC?B. *L ?BBGRGML RM NPMKMRGLE GKKSLC RMJCP?LAC MD F?PKJCQQ ?LRGECLQ, MRFCP QS@QCRQ MD 5PCE ACJJQ ?PC GLTMJTCB GL RFC NPCTCLRGML MD RFC autoimmune response, UFGAF GQ ?L GL?NNPMNPG?RC GKKSLC PCQNMLQC RM FMQR ACJJQ MP QCJD-?LRGECLQ. "LMRFCP 5PCE AJ?QQ QSNNPCQQCQ GKKSLC PCQNMLQCQ RM F?PKDSJ N?RFMECLQ ?DRCP RFC GLDCARGML F?Q AJC?PCB RM KGLGKGXC FMQR ACJJ B?K?EC GLBSACB @W GLDJ?KK?RGML ?LB ACJJ JWQGQ.
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$DDLEFCF>@:8C (<DFIP
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bTC NMNSJ?RGMLQ BGDDCPCLRG?RCQ GLRM # ?LB 5 KCKMPW ACJJQ UGRF RFC Q?KC ?LRGECL QNCAGDGAGRGCQ, ?Q GJJSQRP?RCB GL Figure 42.16. " memory cell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
!@>LI< 42.16 !CQBO FKFQF>IIV ?FKAFKD >K >KQFDBK QL QEB " @BII OB@BMQLO ("#2), > " @BII FKQBOK>IFWBP QEB >KQFDBK >KA MOBPBKQP FQ LK -(# )). ! EBIMBO 4 @BII OB@LDKFWBP QEB -(# ))t>KQFDBK @LJMIBU >KA >@QFS>QBP QEB " @BII. !P > OBPRIQ, JBJLOV " @BIIP >KA MI>PJ> @BIIP >OB J>AB.
4EB 2E >KQFDBK FP CLRKA LK 2E-MLPFQFSB OBA ?ILLA @BIIP. !K 2E-KBD>QFSB CBJ>IB @>K RPR>IIV @>OOV >K 2E-MLPFQFSB CBQRP QL QBOJ TFQELRQ AFCCF@RIQV. (LTBSBO, FC PEB E>P > PB@LKA 2EMLPFQFSB CBQRP, EBO ?LAV J>V I>RK@E >K FJJRKB >QQ>@H QE>Q @>RPBP EBJLIVQF@ AFPB>PB LC QEB KBT?LOK. 7EV AL VLR QEFKH EBJLIVQF@ AFPB>PB FP LKIV > MOL?IBJ AROFKD QEB PB@LKA LO PR?PBNRBKQ MOBDK>K@FBP
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*D RFC N?RFMECL GQ LCTCP CLAMSLRCPCB ?E?GL BSPGLE RFC GLBGTGBS?JsQ JGDCRGKC, # ?LB 5 KCKMPW ACJJQ UGJJ AGPASJ?RC DMP ? DCU WC?PQ MP CTCL QCTCP?J BCA?BCQ ?LB UGJJ EP?BS?JJW BGC MDD, F?TGLE LCTCP DSLARGMLCB ?Q CDDCARMP ACJJQ. )MUCTCP, GD RFC FMQR GQ PC-CVNMQCB RM RFC Q?KC N?RFMECL RWNC, AGPASJ?RGLE KCKMPW ACJJQ UGJJ GKKCBG?RCJW BGDDCPCLRG?RC GLRM NJ?QK? ACJJQ ?LB $5-Q UGRFMSR GLNSR DPMK "1$Q MP 5) ACJJQ. 0LC PC?QML RFC ?B?NRGTC GKKSLC PCQNMLQC GQ BCJ?WCB GQ @CA?SQC GR R?ICQ RGKC DMP L?bTC # ?LB 5 ACJJQ UGRF RFC ?NNPMNPG?RC ?LRGECL QNCAGDGAGRGCQ RM @C GBCLRGDGCB ?LB ?ARGT?RCB. 6NML PCGLDCARGML, RFGQ QRCN GQ QIGNNCB, ?LB RFC PCQSJR GQ ? KMPC P?NGB NPMBSARGML MD GKKSLC BCDCLQCQ. .CKMPW # ACJJQ RF?R BGDDCPCLRG?RC GLRM NJ?QK? ACJJQ MSRNSR RCLQ RM FSLBPCBQ-DMJB EPC?RCP ?LRG@MBW ?KMSLRQ RF?L UCPC QCAPCRCB BSPGLE RFC NPGK?PW PCQNMLQC, ?Q RFC EP?NF GL Figure 42.17 GJJSQRP?RCQ. 5FGQ P?NGB ?LB BP?K?RGA ?LRG@MBW PCQNMLQC K?W QRMN RFC GLDCARGML @CDMPC GR A?L CTCL @CAMKC CQR?@JGQFCB, ?LB RFC GLBGTGBS?J K?W LMR PC?JGXC RFCW F?B @CCL CVNMQCB.
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#(!04%2 42 X 4(% )--5.% 3934%-
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"JRFMSEF RFC GKKSLC QWQRCK GQ AF?P?ARCPGXCB @W AGPASJ?RGLE ACJJQ RFPMSEFMSR RFC @MBW, RFC PCESJ?RGML, K?RSP?RGML, ?LB GLRCPAMKKSLGA?RGML MD GKKSLC D?ARMPQ MAASP ?R QNCAGDGA QGRCQ. 5FC @JMMB AGPASJ?RCQ GKKSLC ACJJQ, NPMRCGLQ, ?LB MRFCP D?ARMPQ RFPMSEF RFC @MBW. "NNPMVGK?RCJW 0.1 NCPACLR MD ?JJ ACJJQ GL RFC @JMMB ?PC JCSIMAWRCQ, UFGAF CLAMKN?QQ KMLMAWRCQ (RFC NPCASPQMP MD K?APMNF?ECQ) ?LB JWKNFMAWRCQ. 5FC K?HMPGRW MD ACJJQ GL RFC @JMMB ?PC CPWRFPMAWRCQ (PCB @JMMB ACJJQ). Lymph GQ ? U?RCPW DJSGB RF?R @?RFCQ RGQQSCQ ?LB MPE?LQ UGRF NPMRCARGTC UFGRC @JMMB ACJJQ ?LB BMCQ LMR AMLR?GL CPWRFPMAWRCQ. $CJJQ MD RFC GKKSLC QWQRCK A?L RP?TCJ @CRUCCL RFC BGQRGLAR JWKNF?RGA ?LB @JMMB AGPASJ?RMPW QWQRCKQ, UFGAF ?PC QCN?P?RCB @W GLRCPQRGRG?J QN?AC, @W ? NPMACQQ A?JJCB CVRP?T?Q?RGML (N?QQGLE RFPMSEF RM QSPPMSLBGLE RGQQSC). 5FC ACJJQ MD RFC GKKSLC QWQRCK MPGEGL?RC DPMK FCK?RMNMGCRGA QRCK ACJJQ GL RFC @MLC K?PPMU. $WRMIGLCQ QRGKSJ?RC RFCQC QRCK ACJJQ RM BGDDCPCLRG?RC GLRM GKKSLC ACJJQ. # ACJJ K?RSP?RGML MAASPQ GL RFC @MLC K?PPMU, UFCPC?Q L?bTC 5 ACJJQ RP?LQGR DPMK RFC @MLC K?PPMU RM RFC RFWKSQ DMP K?RSP?RGML. *L RFC RFWKSQ, GKK?RSPC 5 ACJJQ RF?R CVNPCQQ 5$3Q AMKNJCKCLR?PW RM QCJD-?LRGECLQ ?PC BCQRPMWCB. 5FGQ NPMACQQ FCJNQ NPCTCLR ?SRMGKKSLC PCQNMLQCQ. 0L K?RSP?RGML, 5 ?LB # JWKNFMAWRCQ AGPASJ?RC RM T?PGMSQ BCQRGL?RGMLQ. -WKNF LMBCQ QA?RRCPCB RFPMSEFMSR RFC @MBW, ?Q GJJSQRP?RCB GL Figure 42.20, FMSQC J?PEC NMNSJ?RGMLQ MD 5 ?LB # ACJJQ, BCLBPGRGA ACJJQ, ?LB K?APMNF?ECQ. -WKNF E?RFCPQ ?LRGECLQ ?Q GR BP?GLQ DPMK RGQQSCQ. 5FCQC ?LRGECLQ RFCL ?PC DGJRCPCB RFPMSEF JWKNF LMBCQ @CDMPC RFC JWKNF GQ PCRSPLCB RM AGPASJ?RGML. "1$Q GL RFC JWKNF LMBCQ A?NRSPC ?LB NPMACQQ ?LRGECLQ ?LB GLDMPK LC?P@W JWKNFMAWRCQ ?@MSR NMRCLRG?J N?RFMECLQ.
!@>LI< 42.20 (>) ,VJME>QF@ SBPPBIP @>OOV > @IB>O CIRFA @>IIBA IVJME QEOLRDELRQ QEB ?LAV. 4EB IFNRFA BKQBOP (?) IVJME KLABP QEOLRDE >CCBOBKQ SBPPBIP. ,VJME KLABP >OB CFIIBA TFQE IVJMEL@VQBP QE>Q MRODB FKCB@QFKD @BIIP. 4EB IVJME QEBK BUFQP QEOLRDE BCCBOBKQ SBPPBIP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .)(, .#))
5FC QNJCCL FMSQCQ # ?LB 5 ACJJQ, K?APMNF?ECQ, BCLBPGRGA ACJJQ, ?LB /, ACJJQ. 5FC QNJCCL, QFMUL GL Figure 42.21, GQ RFC QGRC UFCPC "1$Q RF?R F?TC RP?NNCB DMPCGEL N?PRGAJCQ GL RFC @JMMB A?L AMKKSLGA?RC UGRF JWKNFMAWRCQ. "LRG@MBGCQ ?PC QWLRFCQGXCB ?LB QCAPCRCB @W ?ARGT?RCB NJ?QK? ACJJQ GL RFC QNJCCL, ?LB RFC QNJCCL DGJRCPQ DMPCGEL QS@QR?LACQ ?LB ?LRG@MBW-AMKNJCVCB N?RFMECLQ DPMK RFC @JMMB. 'SLARGML?JJW, RFC QNJCCL GQ RM RFC @JMMB ?Q JWKNF LMBCQ ?PC RM RFC JWKNF.
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z &VNJ?GL APMQQ-PC?ARGTGRW z %CQAPG@C RFC QRPSARSPC ?LB DSLARGML MD ?LRG@MBGCQ z %GQASQQ ?LRG@MBW NPMBSARGML "L antibody, ?JQM ILMUL ?Q ?L GKKSLMEJM@SJGL (*E), GQ ? NPMRCGL RF?R GQ NPMBSACB @W NJ?QK? ACJJQ ?DRCP QRGKSJ?RGML @W ?L ?LRGECL. "LRG@MBGCQ ?PC RFC DSLARGML?J @?QGQ MD FSKMP?J GKKSLGRW. "LRG@MBGCQ MAASP GL RFC @JMMB, GL E?QRPGA ?LB KSASQ QCAPCRGMLQ, ?LB GL @PC?QR KGJI. "LRG@MBGCQ GL RFCQC @MBGJW DJSGBQ A?L @GLB N?RFMECLQ ?LB K?PI RFCK DMP BCQRPSARGML @W NF?EMAWRCQ @CDMPC RFCW A?L GLDCAR ACJJQ.
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"L ?LRG@MBW KMJCASJC GQ AMKNPGQCB MD DMSP NMJWNCNRGBCQ: RUM GBCLRGA?J FC?TW AF?GLQ (J?PEC NCNRGBC SLGRQ) RF?R ?PC N?PRG?JJW @MSLB RM C?AF MRFCP GL ? t:u DMPK?RGML, UFGAF ?PC DJ?LICB @W RUM GBCLRGA?J JGEFR AF?GLQ (QK?JJ NCNRGBC SLGRQ), ?Q GJJSQRP?RCB GL Figure 42.22. #MLBQ @CRUCCL RFC AWQRCGLC ?KGLM ?AGBQ GL RFC ?LRG@MBW KMJCASJC ?RR?AF RFC NMJWNCNRGBCQ RM C?AF MRFCP. 5FC ?PC?Q UFCPC RFC ?LRGECL GQ PCAMELGXCB ML RFC ?LRG@MBW ?PC T?PG?@JC BMK?GLQ ?LB RFC ?LRG@MBW @?QC GQ AMKNMQCB MD AMLQR?LR BMK?GLQ. *L ECPK-JGLC # ACJJQ, RFC T?PG?@JC PCEGML MD RFC JGEFR AF?GL ECLC F?Q 40 T?PG?@JC (7) ?LB DGTC HMGLGLE (+) QCEKCLRQ. "L CLXWKC A?JJCB %/" PCAMK@GL?QC P?LBMKJW CVAGQCQ KMQR MD RFCQC QCEKCLRQ MSR MD RFC ECLC, ?LB QNJGACQ MLC 7 QCEKCLR RM MLC + QCEKCLR. %SPGLE 3/" NPMACQQGLE, ?JJ @SR MLC 7 ?LB + QCEKCLR ?PC QNJGACB MSR. 3CAMK@GL?RGML ?LB QNJGAGLE K?W PCQSJR GL MTCP 106 NMQQG@JC 7+ AMK@GL?RGMLQ. "Q ? PCQSJR, C?AF BGDDCPCLRG?RCB # ACJJ GL RFC FSK?L @MBW RWNGA?JJW F?Q ? SLGOSC T?PG?@JC AF?GL. 5FC AMLQR?LR BMK?GL, UFGAF BMCQ LMR @GLB ?LRG@MBW, GQ RFC Q?KC DMP ?JJ ?LRG@MBGCQ.
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q*E., *E(, *E", *E%, *E&q@?QCB ML RFCGP NFWQGMAFCKGA?J, QRPSARSP?J, ?LB GKKSLMJMEGA?J NPMNCPRGCQ. *E(Q, UFGAF K?IC SN ?@MSR 80 NCPACLR MD ?JJ ?LRG@MBGCQ, F?TC FC?TW AF?GLQ RF?R AMLQGQR MD MLC T?PG?@JC BMK?GL ?LB RFPCC GBCLRGA?J AMLQR?LR BMK?GLQ. *E" ?LB *E% ?JQM F?TC RFPCC AMLQR?LR BMK?GLQ NCP FC?TW AF?GL, UFCPC?Q *E. ?LB *E& C?AF F?TC DMSP AMLQR?LR BMK?GLQ NCP FC?TW AF?GL. 5FC T?PG?@JC BMK?GL BCRCPKGLCQ @GLBGLE QNCAGDGAGRW ?LB RFC AMLQR?LR BMK?GL MD RFC FC?TW AF?GL BCRCPKGLCQ RFC GKKSLMJMEGA?J KCAF?LGQK MD ?ARGML MD RFC AMPPCQNMLBGLE ?LRG@MBW AJ?QQ. *R GQ NMQQG@JC DMP RUM ?LRG@MBGCQ RM F?TC RFC Q?KC @GLBGLE QNCAGDGAGRGCQ @SR @C GL BGDDCPCLR AJ?QQCQ ?LB, RFCPCDMPC, RM @C GLTMJTCB GL BGDDCPCLR DSLARGMLQ. "DRCP ?L ?B?NRGTC BCDCLQC GQ NPMBSACB ?E?GLQR ? N?RFMECL, RWNGA?JJW NJ?QK? ACJJQ DGPQR QCAPCRC *E. GLRM RFC @JMMB. #$3Q ML L?bTC # ACJJQ ?PC MD RFC *E. AJ?QQ ?LB MAA?QGML?JJW *E% AJ?QQ. *E. KMJCASJCQ K?IC SN ?NNPMVGK?RCJW RCL NCPACLR MD ?JJ ?LRG@MBGCQ. 1PGMP RM ?LRG@MBW QCAPCRGML, NJ?QK? ACJJQ ?QQCK@JC
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*E. KMJCASJCQ GLRM NCLR?KCPQ (DGTC GLBGTGBS?J ?LRG@MBGCQ) JGLICB @W ? HMGLGLE (+) AF?GL, ?Q QFMUL GL Figure 42.23. 5FC NCLR?KCP ?PP?LECKCLR KC?LQ RF?R RFCQC K?APMKMJCASJCQ A?L @GLB RCL GBCLRGA?J ?LRGECLQ. )MUCTCP, *E. KMJCASJCQ PCJC?QCB C?PJW GL RFC ?B?NRGTC GKKSLC PCQNMLQC BM LMR @GLB RM ?LRGECLQ ?Q QR?@JW ?Q *E(Q, UFGAF ?PC MLC MD RFC NMQQG@JC RWNCQ MD ?LRG@MBGCQ QCAPCRCB GL J?PEC OS?LRGRGCQ SNML PC-CVNMQSPC RM RFC Q?KC N?RFMECL. Figure 42.23 QSKK?PGXCQ RFC NPMNCPRGCQ MD GKKSLMEJM@SJGLQ ?LB GJJSQRP?RCQ RFCGP @?QGA QRPSARSPCQ.
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*E"Q NMNSJ?RC RFC Q?JGT?, RC?PQ, @PC?QR KGJI, ?LB KSASQ QCAPCRGMLQ MD RFC E?QRPMGLRCQRGL?J, PCQNGP?RMPW, ?LB ECLGRMSPGL?PW RP?ARQ. $MJJCARGTCJW, RFCQC @MBGJW DJSGBQ AM?R ?LB NPMRCAR RFC CVRCLQGTC KSAMQ? (4000 QOS?PC DCCR GL FSK?LQ). 5FC RMR?J LSK@CP MD *E" KMJCASJCQ GL RFCQC @MBGJW QCAPCRGMLQ GQ EPC?RCP RF?L RFC LSK@CP MD *E( KMJCASJCQ GL RFC @JMMB QCPSK. " QK?JJ ?KMSLR MD *E" GQ ?JQM QCAPCRCB GLRM RFC QCPSK GL KMLMKCPGA DMPK. $MLTCPQCJW, QMKC *E. GQ QCAPCRCB GLRM @MBGJW DJSGBQ MD RFC KSAMQ?. 4GKGJ?P RM *E., *E" KMJCASJCQ ?PC QCAPCRCB ?Q NMJWKCPGA QRPSARSPCQ JGLICB UGRF ? + AF?GL. )MUCTCP, *E"Q ?PC QCAPCRCB KMQRJW ?Q BGKCPGA KMJCASJCQ, LMR NCLR?KCPQ. *E& GQ NPCQCLR GL RFC QCPSK GL QK?JJ OS?LRGRGCQ ?LB GQ @CQR AF?P?ARCPGXCB GL GRQ PMJC ?Q ?L ?JJCPEW KCBG?RMP. *E% GQ ?JQM NPCQCLR GL QK?JJ OS?LRGRGCQ. 4GKGJ?P RM *E., #$3Q MD RFC *E% AJ?QQ ?PC DMSLB ML RFC QSPD?AC MD L?bTC # ACJJQ. 5FGQ AJ?QQ QSNNMPRQ ?LRGECL PCAMELGRGML ?LB K?RSP?RGML MD # ACJJQ RM NJ?QK? ACJJQ.
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%GDDCPCLRG?RCB NJ?QK? ACJJQ ?PC APSAG?J NJ?WCPQ GL RFC FSKMP?J PCQNMLQC, ?LB RFC ?LRG@MBGCQ RFCW QCAPCRC ?PC N?PRGASJ?PJW QGELGDGA?LR ?E?GLQR CVRP?ACJJSJ?P N?RFMECLQ ?LB RMVGLQ. "LRG@MBGCQ AGPASJ?RC DPCCJW ?LB ?AR GLBCNCLBCLRJW MD NJ?QK? ACJJQ. "LRG@MBGCQ A?L @C RP?LQDCPPCB DPMK MLC GLBGTGBS?J RM ?LMRFCP RM RCKNMP?PGJW NPMRCAR ?E?GLQR GLDCARGMSQ BGQC?QC. 'MP GLQR?LAC, ? NCPQML UFM F?Q PCACLRJW NPMBSACB ? QSAACQQDSJ GKKSLC PCQNMLQC ?E?GLQR ? N?PRGASJ?P BGQC?QC ?ECLR A?L BML?RC @JMMB RM ? LMLGKKSLC PCAGNGCLR ?LB AMLDCP RCKNMP?PW GKKSLGRW RFPMSEF ?LRG@MBGCQ GL RFC BMLMPsQ @JMMB QCPSK. 5FGQ NFCLMKCLML GQ A?JJCB passive immunity; GR ?JQM MAASPQ L?RSP?JJW BSPGLE @PC?QRDCCBGLE, UFGAF K?ICQ @PC?QRDCB GLD?LRQ FGEFJW PCQGQR?LR RM GLDCARGMLQ BSPGLE RFC DGPQR DCU KMLRFQ MD JGDC. "LRG@MBGCQ AM?R CVRP?ACJJSJ?P N?RFMECLQ ?LB LCSRP?JGXC RFCK, ?Q GJJSQRP?RCB GL Figure 42.24, @W @JMAIGLE ICW QGRCQ ML RFC N?RFMECL RF?R CLF?LAC RFCGP GLDCARGTGRW (QSAF ?Q PCACNRMPQ RF?R tBMAIu N?RFMECLQ ML FMQR ACJJQ). "LRG@MBW LCSRP?JGX?RGML A?L NPCTCLR N?RFMECLQ DPMK CLRCPGLE ?LB GLDCARGLE FMQR ACJJQ, ?Q MNNMQCB RM RFC $5--KCBG?RCB ?NNPM?AF MD IGJJGLE ACJJQ RF?R ?PC ?JPC?BW GLDCARCB RM NPCTCLR NPMEPCQQGML MD ?L CQR?@JGQFCB GLDCARGML. 5FC LCSRP?JGXCB ?LRG@MBW-AM?RCB N?RFMECLQ A?L RFCL @C DGJRCPCB @W RFC QNJCCL ?LB CJGKGL?RCB GL SPGLC MP DCACQ.
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affinities (?RRP?ARGML) BCNCLBGLE ML RFC KMJCASJ?P AMKNJCKCLR?PGRW @CRUCCL ?LRGECL ?LB ?LRG@MBW KMJCASJCQ, ?Q GJJSQRP?RCB GL Figure 42.25. "L ?LRG@MBW UGRF ? FGEFCP ?DDGLGRW DMP ? N?PRGASJ?P ?LRGECL UMSJB @GLB KMPC QRPMLEJW ?LB QR?@JW, ?LB RFSQ UMSJB @C CVNCARCB RM NPCQCLR ? KMPC AF?JJCLEGLE BCDCLQC ?E?GLQR RFC N?RFMECL AMPPCQNMLBGLE RM RFC QNCAGDGA ?LRGECL.
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!@>LI< 42.25 (>) !CCFKFQV OBCBOP QL QEB PQOBKDQE LC PFKDIB FKQBO>@QFLK ?BQTBBK >KQFDBK >KA >KQF?LAV, TEFIB >SFAFQV OBCBOP QL QEB PQOBKDQE LC >II FKQBO>@QFLKP @LJ?FKBA. (?) !K >KQF?LAV J>V @OLPP OB>@Q TFQE AFCCBOBKQ BMFQLMBP.
5FC RCPK avidity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ross reactivity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
AEK@9F;@<J F= K?< (L:FJ8C $DDLE< .PJK<D

"LRG@MBGCQ QWLRFCQGXCB @W RFC KSAMQ?J GKKSLC QWQRCK GLAJSBC *E" ?LB *E.. "ARGT?RCB # ACJJQ BGDDCPCLRG?RC GLRM KSAMQ?J NJ?QK? ACJJQ RF?R QWLRFCQGXC ?LB QCAPCRC BGKCPGA *E", ?LB RM ? JCQQCP CVRCLR, NCLR?KCPGA *E.. 4CAPCRCB *E" GQ ?@SLB?LR GL RC?PQ, Q?JGT?, @PC?QR KGJI, ?LB GL QCAPCRGMLQ MD RFC
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#(!04%2 42 X 4(% )--5.% 3934%-
E?QRPMGLRCQRGL?J ?LB PCQNGP?RMPW RP?ARQ. "LRG@MBW QCAPCRGML PCQSJRQ GL ? JMA?J FSKMP?J PCQNMLQC ?R CNGRFCJG?J QSPD?ACQ ?LB NPCTCLRQ GLDCARGML MD RFC KSAMQ? @W @GLBGLE ?LB LCSRP?JGXGLE N?RFMECLQ.
42.4 R D@JILGK@FEJ @E K?< $DDLE< .PJK<D

#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CQAPG@C FWNCPQCLQGRGTGRW z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sQ A?N?AGRW RM ECLCP?RC QSDDGAGCLR PCQNMLQCQ RM GLDCARGML MP RSKMPQ. "Q ? PCQSJR, )*7-GLDCARCB GLBGTGBS?JQ MDRCL QSDDCP DPMK GLDCARGMLQ RF?R UMSJB LMR A?SQC GJJLCQQ GL NCMNJC UGRF FC?JRFW GKKSLC QWQRCKQ @SR UFGAF A?L A?SQC BCT?QR?RGLE GJJLCQQ RM GKKSLC-AMKNPMKGQCB GLBGTGBS?JQ. .?J?B?NRGTC PCQNMLQCQ MD GKKSLC ACJJQ ?LB KMJCASJCQ RFCKQCJTCQ A?L ?JQM BGQPSNR RFC NPMNCP DSLARGMLGLE MD RFC CLRGPC QWQRCK, JC?BGLE RM FMQR ACJJ B?K?EC RF?R AMSJB @CAMKC D?R?J.
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'?GJSPCQ, GLQSDDGAGCLAGCQ, MP BCJ?WQ ?R ?LW JCTCJ MD RFC GKKSLC PCQNMLQC A?L ?JJMU N?RFMECLQ MP RSKMP ACJJQ RM E?GL ? DMMRFMJB ?LB PCNJGA?RC MP NPMJGDCP?RC RM FGEF CLMSEF JCTCJQ RF?R RFC GKKSLC QWQRCK @CAMKCQ MTCPUFCJKCB. Immunodeficiency GQ RFC D?GJSPC, GLQSDDGAGCLAW, MP BCJ?W GL RFC PCQNMLQC MD RFC GKKSLC QWQRCK, UFGAF K?W @C ?AOSGPCB MP GLFCPGRCB. *KKSLMBCDGAGCLAW A?L @C ?AOSGPCB ?Q ? PCQSJR MD GLDCARGML UGRF ACPR?GL N?RFMECLQ (QSAF ?Q )*7), AFCKGA?J CVNMQSPC (GLAJSBGLE ACPR?GL KCBGA?J RPC?RKCLRQ), K?JLSRPGRGML, MP NMQQG@JW @W CVRPCKC QRPCQQ. 'MP GLQR?LAC, P?BG?RGML CVNMQSPC A?L BCQRPMW NMNSJ?RGMLQ MD JWKNFMAWRCQ ?LB CJCT?RC ?L GLBGTGBS?JsQ QSQACNRG@GJGRW RM GLDCARGMLQ ?LB A?LACP. %MXCLQ MD ECLCRGA BGQMPBCPQ PCQSJR GL GKKSLMBCDGAGCLAGCQ, GLAJSBGLE 4CTCPC $MK@GLCB *KKSLMBCDGAGCLAW (4$*%), #?PC JWKNFMAWRC QWLBPMKC, ?LB .)$ ** BCDGAGCLAGCQ. 3?PCJW, NPGK?PW GKKSLMBCDGAGCLAGCQ RF?R ?PC NPCQCLR DPMK @GPRF K?W MAASP. /CSRPMNCLG? GQ MLC DMPK GL UFGAF RFC GKKSLC QWQRCK NPMBSACQ ? @CJMU?TCP?EC LSK@CP MD LCSRPMNFGJQ, RFC @MBWsQ KMQR ?@SLB?LR NF?EMAWRCQ. "Q ? PCQSJR, @?ARCPG?J GLDCARGMLQ K?W EM SLPCQRPGARCB GL RFC @JMMB, A?SQGLE QCPGMSQ AMKNJGA?RGMLQ.
#PG<IJ<EJ@K@M@K@<J
.?J?B?NRGTC GKKSLC PCQNMLQCQ RMU?PB F?PKJCQQ DMPCGEL QS@QR?LACQ MP QCJD ?LRGECLQ RF?R MAASP ?DRCP RGQQSC QCLQGRGX?RGML ?PC RCPKCB hypersensitivities. 5FC RWNCQ MD FWNCPQCLQGRGTGRGCQ GLAJSBC GKKCBG?RC, BCJ?WCB, ?LB ?SRMGKKSLGRW. " J?PEC NPMNMPRGML MD RFC NMNSJ?RGML GQ ?DDCARCB @W MLC MP KMPC RWNCQ MD FWNCPQCLQGRGTGRW. A11+7-/+8 5FC GKKSLC PC?ARGML RF?R PCQSJRQ DPMK GKKCBG?RC FWNCPQCLQGRGTGRGCQ GL UFGAF ?L ?LRG@MBW-KCBG?RCB GKKSLC PCQNMLQC MAASPQ UGRFGL KGLSRCQ MD CVNMQSPC RM ? F?PKJCQQ ?LRGECL GQ A?JJCB ?L allergy. *L RFC 6LGRCB 4R?RCQ, 20 NCPACLR MD RFC NMNSJ?RGML CVFG@GRQ QWKNRMKQ MD ?JJCPEW MP ?QRFK?, UFCPC?Q 55 NCPACLR RCQR NMQGRGTC ?E?GLQR MLC MP KMPC ?JJCPECLQ. 6NML GLGRG?J CVNMQSPC RM ? NMRCLRG?J ?JJCPECL, ?L ?JJCPEGA GLBGTGBS?J QWLRFCQGXCQ ?LRG@MBGCQ MD RFC *E& AJ?QQ TG? RFC RWNGA?J NPMACQQ MD "1$Q NPCQCLRGLE NPMACQQCB ?LRGECL RM 5) ACJJQ RF?R QRGKSJ?RC # ACJJQ RM NPMBSAC *E&. 5FGQ AJ?QQ MD ?LRG@MBGCQ ?JQM KCBG?RCQ RFC GKKSLC PCQNMLQC RM N?P?QGRGA UMPKQ. 5FC AMLQR?LR BMK?GL MD RFC *E& KMJCASJCQ GLRCP?AR UGRF K?QR ACJJQ CK@CBBCB GL AMLLCARGTC RGQQSCQ. 5FGQ NPMACQQ NPGKCQ, MP QCLQGRGXCQ, RFC RGQQSC. 6NML QS@QCOSCLR CVNMQSPC RM RFC Q?KC ?JJCPECL, *E& KMJCASJCQ ML K?QR ACJJQ @GLB RFC ?LRGECL TG? RFCGP T?PG?@JC BMK?GLQ ?LB QRGKSJ?RC RFC K?QR ACJJ RM PCJC?QC RFC KMBGDGCB ?KGLM ?AGBQ FGQR?KGLC ?LB QCPMRMLGL; RFCQC AFCKGA?J KCBG?RMPQ RFCL PCAPSGR CMQGLMNFGJQ UFGAF KCBG?RC ?JJCPEGA PCQNMLQCQ. Figure 42.26 QFMUQ ?L CV?KNJC MD ?L ?JJCPEGA PCQNMLQC RM P?EUCCB NMJJCL. 5FC CDDCARQ MD ?L ?JJCPEGA PC?ARGML P?LEC DPMK KGJB QWKNRMKQ JGIC QLCCXGLE ?LB GRAFW, U?RCPW CWCQ RM KMPC QCTCPC MP CTCL JGDC-RFPC?RCLGLE PC?ARGMLQ GLTMJTGLE GLRCLQCJW
#(!04%2 42 X 4(% )--5.% 3934%-
1239
GRAFW UCJRQ MP FGTCQ, ?GPU?W AMLRP?ARGML UGRF QCTCPC PCQNGP?RMPW BGQRPCQQ, ?LB NJSKKCRGLE @JMMB NPCQQSPC. 5FGQ CVRPCKC PC?ARGML GQ ILMUL ?Q ?L?NFWJ?ARGA QFMAI. *D LMR RPC?RCB UGRF CNGLCNFPGLC RM AMSLRCP RFC @JMMB NPCQQSPC ?LB @PC?RFGLE CDDCARQ, RFGQ AMLBGRGML A?L @C D?R?J.
!@>LI< 42.26 /K CFOPQ BUMLPROB QL >K >IIBODBK, >K )D% >KQF?LAV FP PVKQEBPFWBA ?V MI>PJ> @BIIP FK OBPMLKPB QL > E>OJIBPP >KQFDBK. 4EB )D% JLIB@RIBP ?FKA QL J>PQ @BIIP, >KA LK PB@LKA>OV BUMLPROB, QEB J>PQ @BIIP OBIB>PB EFPQ>JFKBP >KA LQEBO JLARI>QLOP QE>Q >CCB@Q QEB PVJMQLJP LC >IIBODV. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V .)()
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utoimmunity GQ ? RWNC MD FWNCPQCLQGRGTGRW RM QCJD ?LRGECLQ RF?R ?DDCARQ ?NNPMVGK?RCJW DGTC NCPACLR MD RFC NMNSJ?RGML. .MQR RWNCQ MD ?SRMGKKSLGRW GLTMJTC RFC FSKMP?J GKKSLC PCQNMLQC. "LRG@MBGCQ RF?R GL?NNPMNPG?RCJW K?PI QCJD AMKNMLCLRQ ?Q DMPCGEL ?PC RCPKCB autoantibodies. *L N?RGCLRQ UGRF RFC ?SRMGKKSLC BGQC?QC KW?QRFCLG? EP?TGQ, KSQAJC ACJJ PCACNRMPQ RF?R GLBSAC AMLRP?ARGML GL PCQNMLQC RM ?ACRWJAFMJGLC ?PC R?PECRCB @W ?LRG@MBGCQ. 5FC PCQSJR GQ KSQAJC UC?ILCQQ RF?R K?W GLAJSBC K?PICB BGDDGASJRJW UGRF DGLC ?LB/MP EPMQQ KMRMP DSLARGMLQ. *L QWQRCKGA JSNSQ CPWRFCK?RMQSQ, ? BGDDSQC ?SRM?LRG@MBW PCQNMLQC RM RFC GLBGTGBS?JsQ MUL %/" ?LB NPMRCGLQ PCQSJRQ GL T?PGMSQ QWQRCKGA BGQC?QCQ. "Q GJJSQRP?RCB GL Figure 42.27, QWQRCKGA JSNSQ CPWRFCK?RMQSQ K?W ?DDCAR RFC FC?PR, HMGLRQ, JSLEQ, QIGL,
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#(!04%2 42 X 4(% )--5.% 3934%-
IGBLCWQ, ACLRP?J LCPTMSQ QWQRCK, MP MRFCP RGQQSCQ, A?SQGLE RGQQSC B?K?EC TG? ?LRG@MBW @GLBGLE, AMKNJCKCLR PCAPSGRKCLR, JWQGQ, ?LB GLDJ?KK?RGML.
!@>LI< 42.27 3VPQBJF@ IRMRP BOVQEBJ>QLPRP FP @E>O>@QBOFWBA ?V >RQLFJJRKFQV QL QEB FKAFSFAR>IwP LTK $.! >KA/LO MOLQBFKP, TEF@E IB>AP QL S>OFBA AVPCRK@QFLK LC QEB LOD>KP. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V -FH>BI (aDDPQOhJ)
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
#(!04%2 42 X 4(% )--5.% 3934%-
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& 4 / -(.
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tLML-QCJDu NPMRCGL RF?R RPGEECPQ RFC GKKSLC PCQNMLQC 8LKF8EK@9F;P ?LRG@MBW RF?R GLAMPPCARJW K?PIQ tQCJDu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#(!04%2 42 X 4(% )--5.% 3934%-
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"QCVS?J PCNPMBSARGML MAASPQ GL NPMI?PWMRGA KGAPMMPE?LGQKQ (@?ARCPG?) ?LB GL QMKC CSI?PWMRGA QGLEJCACJJCB ?LB KSJRG-ACJJCB MPE?LGQKQ. 5FCPC ?PC ? LSK@CP MD U?WQ RF?R ?LGK?JQ PCNPMBSAC ?QCVS?JJW. F/88/43 Fission, ?JQM A?JJCB @GL?PW DGQQGML, MAASPQ GL NPMI?PWMRGA KGAPMMPE?LGQKQ ?LB GL QMKC GLTCPRC@P?RC, KSJRG-ACJJCB MPE?LGQKQ. "DRCP ? NCPGMB MD EPMURF, ?L MPE?LGQK QNJGRQ GLRM RUM QCN?P?RC MPE?LGQKQ. 4MKC SLGACJJSJ?P CSI?PWMRGA MPE?LGQKQ SLBCPEM @GL?PW DGQQGML @W KGRMQGQ. *L MRFCP MPE?LGQKQ, N?PR MD RFC GLBGTGBS?J QCN?P?RCQ ?LB DMPKQ ? QCAMLB GLBGTGBS?J. 5FGQ NPMACQQ MAASPQ, DMP CV?KNJC, GL K?LW ?QRCPMGB CAFGLMBCPKQ RFPMSEF QNJGRRGLE MD RFC ACLRP?J BGQI. 4MKC QC? ?LCKMLCQ ?LB QMKC AMP?J NMJWNQ (Figure 43.2) ?JQM PCNPMBSAC RFPMSEF DGQQGML.
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B:**/3Budding GQ ? DMPK MD ?QCVS?J PCNPMBSARGML RF?R PCQSJRQ DPMK RFC MSREPMURF MD ? N?PR MD ? ACJJ MP @MBW PCEGML JC?BGLE RM ? QCN?P?RGML DPMK RFC MPGEGL?J MPE?LGQK GLRM RUM GLBGTGBS?JQ. #SBBGLE MAASPQ AMKKMLJW GL QMKC GLTCPRC@P?RC ?LGK?JQ QSAF ?Q AMP?JQ ?LB FWBP?Q. *L FWBP?Q, ? @SB DMPKQ RF?R BCTCJMNQ GLRM ?L ?BSJR ?LB @PC?IQ ?U?W DPMK RFC K?GL @MBW, ?Q GJJSQRP?RCB GL Figure 43.3, UFCPC?Q GL AMP?J @SBBGLE, RFC @SB BMCQ LMR BCR?AF ?LB KSJRGNJGCQ ?Q N?PR MD ? LCU AMJMLW.
!@>LI< 43.3 (VAO> OBMOLAR@B >PBUR>IIV QEOLRDE ?RAAFKD.
8?RAF ? video (http://openstaxcollege.org/l/budding_hydra) MD ? FWBP? @SBBGLE. F7'-2+39'9/43 Fragmentation GQ RFC @PC?IGLE MD RFC @MBW GLRM RUM N?PRQ UGRF QS@QCOSCLR PCECLCP?RGML. *D RFC ?LGK?J GQ A?N?@JC MD DP?EKCLR?RGML, ?LB RFC N?PR GQ @GE CLMSEF, ? QCN?P?RC GLBGTGBS?J UGJJ PCEPMU. 'MP CV?KNJC, GL K?LW QC? QR?PQ, ?QCVS?J PCNPMBSARGML GQ ?AAMKNJGQFCB @W DP?EKCLR?RGML. Figure 43.4 GJJSQRP?RCQ ? QC? QR?P DMP UFGAF ?L ?PK MD RFC GLBGTGBS?J GQ @PMICL MDD ?LB PCECLCP?RCQ ? LCU QC? QR?P. 'GQFCPGCQ UMPICPQ F?TC @CCL ILMUL RM RPW RM IGJJ RFC QC? QR?PQ C?RGLE RFCGP AJ?K MP MWQRCP @CBQ @W ASRRGLE RFCK GL F?JD ?LB RFPMUGLE RFCK @?AI GLRM RFC MAC?L. 6LDMPRSL?RCJW DMP RFC UMPICPQ, RFC RUM N?PRQ A?L C?AF PCECLCP?RC ? LCU F?JD, PCQSJRGLE GL RUGAC ?Q K?LW QC? QR?PQ RM NPCW SNML RFC MWQRCPQ ?LB AJ?KQ. 'P?EKCLR?RGML ?JQM MAASPQ GL ?LLCJGB UMPKQ, RSP@CJJ?PG?LQ, ?LB NMPGDCP?LQ.
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1250
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/MRC RF?R GL DP?EKCLR?RGML, RFCPC GQ ECLCP?JJW ? LMRGAC?@JC BGDDCPCLAC GL RFC QGXC MD RFC GLBGTGBS?JQ, UFCPC?Q GL DGQQGML, RUM GLBGTGBS?JQ MD ?NNPMVGK?RC QGXC ?PC DMPKCB. P'79.+34-+3+8/8 Parthenogenesis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qQSAF ?Q ACPR?GL PCNRGJCQ, ?KNFG@G?LQ, ?LB DGQFq?JQM PCNPMBSAC RFPMSEF N?PRFCLMECLCQGQ. "JRFMSEF KMPC AMKKML GL NJ?LRQ, N?PRFCLMECLCQGQ F?Q @CCL M@QCPTCB GL ?LGK?J QNCAGCQ RF?R UCPC QCEPCE?RCB @W QCV GL RCPPCQRPG?J MP K?PGLC XMMQ. 5UM DCK?JC ,MKMBM BP?EMLQ, ? F?KKCPFC?B QF?PI, ?LB ? @J?AIRMN QF?PI F?TC NPMBSACB N?PRFCLMECLGA WMSLE UFCL RFC DCK?JCQ F?TC @CCL GQMJ?RCB DPMK K?JCQ.
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4CVS?J PCNPMBSARGML GQ RFC AMK@GL?RGML MD (SQS?JJW F?NJMGB) PCNPMBSARGTC ACJJQ DPMK RUM GLBGTGBS?JQ RM DMPK ? RFGPB (SQS?JJW BGNJMGB) SLGOSC MDDQNPGLE. 4CVS?J PCNPMBSARGML NPMBSACQ MDDQNPGLE UGRF LMTCJ AMK@GL?RGMLQ MD ECLCQ. 5FGQ A?L @C ?L ?B?NRGTC ?BT?LR?EC GL SLQR?@JC MP SLNPCBGAR?@JC CLTGPMLKCLRQ. "Q FSK?LQ, UC ?PC SQCB RM RFGLIGLE MD ?LGK?JQ ?Q F?TGLE RUM QCN?P?RC QCVCQqK?JC ?LB DCK?JCqBCRCPKGLCB ?R AMLACNRGML. )MUCTCP, GL RFC ?LGK?J IGLEBMK, RFCPC ?PC K?LW T?PG?RGMLQ ML RFGQ RFCKC. H+72'5.74*/9/82 Hermaphroditism MAASPQ GL ?LGK?JQ UFCPC MLC GLBGTGBS?J F?Q @MRF K?JC ?LB DCK?JC PCNPMBSARGTC N?PRQ. *LTCPRC@P?RCQ QSAF ?Q C?PRFUMPKQ, QJSEQ, R?NCUMPKQ ?LB QL?GJQ, QFMUL GL Figure 43.5, ?PC MDRCL FCPK?NFPMBGRGA. )CPK?NFPMBGRCQ K?W QCJD-DCPRGJGXC MP K?W K?RC UGRF ?LMRFCP MD RFCGP QNCAGCQ, DCPRGJGXGLE C?AF MRFCP ?LB @MRF NPMBSAGLE MDDQNPGLE. 4CJD DCPRGJGX?RGML GQ AMKKML GL ?LGK?JQ RF?R F?TC JGKGRCB KM@GJGRW MP ?PC LMR KMRGJC, QSAF ?Q @?PL?AJCQ ?LB AJ?KQ.
!@>LI< 43.5 ->KV PK>FIP >OB EBOJ>MEOLAFQBP. 7EBK QTL FKAFSFAR>IP J>QB, QEBV @>K MOLAR@B RM QL LKB ERKAOBA BDDP B>@E. (@OBAFQ: !PP>C 3EQFIJ>K)
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.?KK?JG?L QCV BCRCPKGL?RGML GQ BCRCPKGLCB ECLCRGA?JJW @W RFC NPCQCLAC MD 9 ?LB : AFPMKMQMKCQ. *LBGTGBS?JQ FMKMXWEMSQ DMP 9 (99) ?PC DCK?JC ?LB FCRCPMXWEMSQ GLBGTGBS?JQ (9:) ?PC K?JC. 5FC NPCQCLAC MD ? : AFPMKMQMKC A?SQCQ RFC BCTCJMNKCLR MD K?JC AF?P?ARCPGQRGAQ ?LB GRQ ?@QCLAC PCQSJRQ GL DCK?JC AF?P?ARCPGQRGAQ. 5FC 9: QWQRCK GQ ?JQM DMSLB GL QMKC GLQCARQ ?LB NJ?LRQ. "TG?L QCV BCRCPKGL?RGML GQ BCNCLBCLR ML RFC NPCQCLAC MD ; ?LB 8 AFPMKMQMKCQ. )MKMXWEMSQ DMP ; (;;) PCQSJRQ GL ? K?JC ?LB FCRCPMXWEMSQ (;8) PCQSJRQ GL ? DCK?JC. 5FC 8 ?NNC?PQ RM @C CQQCLRG?J GL BCRCPKGLGLE RFC QCV MD RFC GLBGTGBS?J, QGKGJ?P RM RFC : AFPMKMQMKC GL K?KK?JQ. 4MKC DGQF, APSQR?AC?LQ, GLQCARQ (QSAF ?Q @SRRCPDJGCQ ?LB KMRFQ), ?LB PCNRGJCQ SQC RFGQ QWQRCK. 5FC QCV MD QMKC QNCAGCQ GQ LMR BCRCPKGLCB @W ECLCRGAQ @SR @W QMKC ?QNCAR MD RFC CLTGPMLKCLR. 4CV BCRCPKGL?RGML GL QMKC APMAMBGJCQ ?LB RSPRJCQ, DMP CV?KNJC, GQ MDRCL BCNCLBCLR ML RFC RCKNCP?RSPC BSPGLE APGRGA?J NCPGMBQ MD CEE BCTCJMNKCLR. 5FGQ GQ PCDCPPCB RM ?Q CLTGPMLKCLR?J QCV BCRCPKGL?RGML, MP KMPC QNCAGDGA?JJW ?Q RCKNCP?RSPC-BCNCLBCLR QCV BCRCPKGL?RGML. *L K?LW RSPRJCQ, AMMJCP RCKNCP?RSPCQ BSPGLE
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CEE GLAS@?RGML NPMBSAC K?JCQ ?LB U?PK RCKNCP?RSPCQ NPMBSAC DCK?JCQ. *L QMKC APMAMBGJCQ, KMBCP?RC RCKNCP?RSPCQ NPMBSAC K?JCQ ?LB @MRF U?PK ?LB AMMJ RCKNCP?RSPCQ NPMBSAC DCK?JCQ. *L QMKC QNCAGCQ, QCV GQ @MRF ECLCRGA- ?LB RCKNCP?RSPC-BCNCLBCLR. *LBGTGBS?JQ MD QMKC QNCAGCQ AF?LEC RFCGP QCV BSPGLE RFCGP JGTCQ, ?JRCPL?RGLE @CRUCCL K?JC ?LB DCK?JC. *D RFC GLBGTGBS?J GQ DCK?JC DGPQR, GR GQ RCPKCB NPMRMEWLW MP tDGPQR DCK?JC,u GD GR GQ K?JC DGPQR, GRQ RCPKCB NPMR?LBPW MP tDGPQR K?JC.u 0WQRCPQ, DMP CV?KNJC, ?PC @MPL K?JC, EPMU, ?LB @CAMKC DCK?JC ?LB J?W CEEQ; QMKC MWQRCP QNCAGCQ AF?LEC QCV KSJRGNJC RGKCQ.
43.2 R !<IK@C@Q8K@FE
#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %GQASQQ GLRCPL?J ?LB CVRCPL?J KCRFMBQ MD DCPRGJGX?RGML z %CQAPG@C RFC KCRFMBQ SQCB @W ?LGK?JQ DMP BCTCJMNKCLR MD MDDQNPGLE BSPGLE ECQR?RGML z %CQAPG@C RFC ?L?RMKGA?J ?B?NRGMLQ RF?R MAASPPCB GL ?LGK?JQ RM D?AGJGR?RC PCNPMBSARGML 4CVS?J PCNPMBSARGML QR?PRQ UGRF RFC AMK@GL?RGML MD ? QNCPK ?LB ?L CEE GL ? NPMACQQ A?JJCB DCPRGJGX?RGML. 5FGQ A?L MAASP CGRFCP GLQGBC ( internal fertilization) MP MSRQGBC ( external fertilization) RFC @MBW MD RFC DCK?JC. )SK?LQ NPMTGBC ?L CV?KNJC MD RFC DMPKCP UFCPC?Q QC?FMPQC PCNPMBSARGML GQ ?L CV?KNJC MD RFC J?RRCP.
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&VRCPL?J DCPRGJGX?RGML SQS?JJW MAASPQ GL ?OS?RGA CLTGPMLKCLRQ UFCPC @MRF CEEQ ?LB QNCPK ?PC PCJC?QCB GLRM RFC U?RCP. "DRCP RFC QNCPK PC?AFCQ RFC CEE, DCPRGJGX?RGML R?ICQ NJ?AC. .MQR CVRCPL?J DCPRGJGX?RGML F?NNCLQ BSPGLE RFC NPMACQQ MD QN?ULGLE UFCPC MLC MP QCTCP?J DCK?JCQ PCJC?QC RFCGP CEEQ ?LB RFC K?JC(Q) PCJC?QC QNCPK GL RFC Q?KC ?PC?, ?R RFC Q?KC RGKC. 5FC PCJC?QC MD RFC PCNPMBSARGTC K?RCPG?J K?W @C RPGEECPCB @W U?RCP RCKNCP?RSPC MP RFC JCLERF MD B?WJGEFR. /C?PJW ?JJ DGQF QN?UL, ?Q BM APSQR?AC?LQ (QSAF ?Q AP?@Q ?LB QFPGKN), KMJJSQIQ (QSAF ?Q MWQRCPQ), QOSGB, ?LB CAFGLMBCPKQ (QSAF ?Q QC? SPAFGLQ ?LB QC? ASASK@CPQ). Figure 43.6 QFMUQ Q?JKML QN?ULGLE GL ? QF?JJMU QRPC?K. 'PMEQ, JGIC RFMQC QFMUL GL Figure 43.7, AMP?JQ, K?WDJGCQ, ?LB KMQOSGRMCQ ?JQM QN?UL.
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1252
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oviparity, DCPRGJGXCB CEEQ ?PC J?GB MSRQGBC RFC DCK?JCsQ @MBW ?LB BCTCJMN RFCPC, PCACGTGLE LMSPGQFKCLR DPMK RFC WMJI RF?R GQ ? N?PR MD RFC CEE. 5FGQ MAASPQ GL KMQR @MLW DGQF, K?LW PCNRGJCQ, QMKC A?PRGJ?EGLMSQ DGQF, KMQR ?KNFG@G?LQ, RUM K?KK?JQ, ?LB ?JJ @GPBQ. 3CNRGJCQ ?LB GLQCARQ NPMBSAC JC?RFCPW CEEQ, UFGJC @GPBQ ?LB RSPRJCQ NPMBSAC CEEQ UGRF FGEF AMLACLRP?RGMLQ MD A?JAGSK A?P@ML?RC GL RFC QFCJJ, K?IGLE RFCK F?PB. $FGAICL CEEQ ?PC ?L CV?KNJC MD RFGQ QCAMLB RWNC. *L ovoviparity, DCPRGJGXCB CEEQ ?PC PCR?GLCB GL RFC DCK?JC, @SR RFC CK@PWM M@R?GLQ GRQ LMSPGQFKCLR DPMK RFC CEEsQ WMJI ?LB RFC WMSLE ?PC DSJJW BCTCJMNCB UFCL RFCW ?PC F?RAFCB. 5FGQ MAASPQ GL QMKC @MLW DGQF (JGIC RFC ESNNW L,)0:;,: 9,;0*<3(;<:), QMKC QF?PIQ, QMKC JGX?PBQ, QMKC QL?ICQ (QSAF ?Q RFC E?PRCP QL?IC !/(4567/0: :09;(30:), QMKC TGNCPQ, ?LB QMKC GLTCPRC@P?RC ?LGK?JQ (JGIC RFC .?B?E?QA?P FGQQGLE AMAIPM?AF G9647/(+69/05( 769;,5;6:(). *L viviparity RFC WMSLE BCTCJMN UGRFGL RFC DCK?JC, PCACGTGLE LMSPGQFKCLR DPMK RFC KMRFCPsQ @JMMB RFPMSEF ? NJ?ACLR?. 5FC MDDQNPGLE BCTCJMNQ GL RFC DCK?JC ?LB GQ @MPL ?JGTC. 5FGQ MAASPQ GL KMQR K?KK?JQ, QMKC A?PRGJ?EGLMSQ DGQF, ?LB ? DCU PCNRGJCQ. *LRCPL?J DCPRGJGX?RGML F?Q RFC ?BT?LR?EC MD NPMRCARGLE RFC DCPRGJGXCB CEE DPMK BCFWBP?RGML ML J?LB. 5FC CK@PWM GQ GQMJ?RCB UGRFGL RFC DCK?JC, UFGAF JGKGRQ NPCB?RGML ML RFC WMSLE. *LRCPL?J DCPRGJGX?RGML CLF?LACQ RFC DCPRGJGX?RGML MD CEEQ @W ? QNCAGDGA K?JC. 'CUCP MDDQNPGLE ?PC NPMBSACB RFPMSEF RFGQ KCRFMB, @SR RFCGP QSPTGT?J P?RC GQ FGEFCP RF?L RF?R DMP CVRCPL?J DCPRGJGX?RGML.
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0LAC KSJRGACJJSJ?P MPE?LGQKQ CTMJTCB ?LB BCTCJMNCB QNCAG?JGXCB ACJJQ, QMKC ?JQM BCTCJMNCB RGQQSCQ ?LB MPE?LQ UGRF QNCAG?JGXCB DSLARGMLQ. "L C?PJW BCTCJMNKCLR GL PCNPMBSARGML MAASPPCB GL RFC "LLCJGBQ. 5FCQC MPE?LGQKQ NPMBSAC QNCPK ?LB CEEQ DPMK SLBGDDCPCLRG?RCB ACJJQ GL RFCGP AMCJMK ?LB QRMPC RFCK GL RF?R A?TGRW. 8FCL RFC AMCJMK @CAMKCQ DGJJCB, RFC ACJJQ ?PC PCJC?QCB RFPMSEF ?L CVAPCRMPW MNCLGLE MP @W RFC @MBW QNJGRRGLE MNCL. 3CNPMBSARGTC MPE?LQ CTMJTCB UGRF RFC BCTCJMNKCLR MD EML?BQ RF?R NPMBSAC
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(?KCRMECLCQGQ, RFC NPMBSARGML MD QNCPK ?LB CEEQ, R?ICQ NJ?AC RFPMSEF RFC NPMACQQ MD KCGMQGQ. %SPGLE KCGMQGQ, RUM ACJJ BGTGQGMLQ QCN?P?RC RFC N?GPCB AFPMKMQMKCQ GL RFC LSAJCSQ ?LB RFCL QCN?P?RC RFC AFPMK?RGBQ RF?R UCPC K?BC BSPGLE ?L C?PJGCP QR?EC MD RFC ACJJsQ JGDC AWAJC. .CGMQGQ NPMBSACQ F?NJMGB ACJJQ UGRF F?JD MD C?AF N?GP MD AFPMKMQMKCQ LMPK?JJW DMSLB GL BGNJMGB ACJJQ. 5FC NPMBSARGML MD QNCPK GQ A?JJCB spermatogenesis ?LB RFC NPMBSARGML MD CEEQ GQ A?JJCB oogenesis. !5+72'94-+3+8/8
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4NCPK?RMECLCQGQ, GJJSQRP?RCB GL Figure 43.12, MAASPQ GL RFC U?JJ MD RFC QCKGLGDCPMSQ RS@SJCQ (Figure 43.8), UGRF QRCK ACJJQ ?R RFC NCPGNFCPW MD RFC RS@C ?LB RFC QNCPK?RMXM? ?R RFC JSKCL MD RFC RS@C. *KKCBG?RCJW SLBCP RFC A?NQSJC MD RFC RS@SJC ?PC BGNJMGB, SLBGDDCPCLRG?RCB ACJJQ. 5FCQC QRCK ACJJQ, A?JJCB QNCPK?RMEMLG? (QGLESJ?P: QNCPK?R?EMLGSK), EM RFPMSEF KGRMQGQ UGRF MLC MDDQNPGLE EMGLE ML RM BGDDCPCLRG?RC GLRM ? QNCPK ACJJ ?LB RFC MRFCP EGTGLE PGQC RM RFC LCVR ECLCP?RGML MD QNCPK. .CGMQGQ QR?PRQ UGRF ? ACJJ A?JJCB ? NPGK?PW QNCPK?RMAWRC. "R RFC CLB MD RFC DGPQR KCGMRGA BGTGQGML, ? F?NJMGB ACJJ GQ NPMBSACB A?JJCB ? QCAMLB?PW QNCPK?RMAWRC. 5FGQ ACJJ GQ F?NJMGB ?LB KSQR EM RFPMSEF ?LMRFCP KCGMRGA ACJJ BGTGQGML. 5FC ACJJ NPMBSACB ?R RFC CLB MD KCGMQGQ GQ A?JJCB ? QNCPK?RGB ?LB UFCL GR PC?AFCQ RFC JSKCL MD RFC RS@SJC ?LB EPMUQ ? DJ?ECJJSK, GR GQ A?JJCB ? QNCPK ACJJ. 'MSP QNCPK PCQSJR DPMK C?AF NPGK?PW QNCPK?RMAWRC RF?R EMCQ RFPMSEF KCGMQGQ. 4RCK ACJJQ ?PC BCNMQGRCB BSPGLE ECQR?RGML ?LB ?PC NPCQCLR ?R @GPRF RFPMSEF RFC @CEGLLGLE MD ?BMJCQACLAC, @SR GL ?L GL?ARGTC QR?RC. %SPGLE ?BMJCQACLAC, EML?BMRPMNGA FMPKMLCQ DPMK RFC ?LRCPGMP NGRSGR?PW A?SQC RFC ?ARGT?RGML MD RFCQC ACJJQ ?LB RFC NPMBSARGML MD TG?@JC QNCPK. 5FGQ AMLRGLSCQ GLRM MJB ?EC.
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7GQGR this site (http://openstaxcollege.org/l/spermatogenesis) RM QCC RFC NPMACQQ MD QNCPK?RMECLCQGQ. O4-+3+8/8 0MECLCQGQ, GJJSQRP?RCB GL Figure 43.13, MAASPQ GL RFC MSRCPKMQR J?WCPQ MD RFC MT?PGCQ. "Q UGRF QNCPK NPMBSARGML, MMECLCQGQ QR?PRQ UGRF ? ECPK ACJJ, A?JJCB ?L MMEMLGSK (NJSP?J: MMEMLG?), @SR RFGQ ACJJ SLBCPEMCQ KGRMQGQ RM GLAPC?QC GL LSK@CP, CTCLRS?JJW PCQSJRGLE GL SN RM ?@MSR MLC RM RUM KGJJGML ACJJQ GL RFC CK@PWM.
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*L DCK?JCQ, '4) QRGKSJ?RCQ BCTCJMNKCLR MD CEE ACJJQ, A?JJCB MT?, UFGAF BCTCJMN GL QRPSARSPCQ A?JJCB DMJJGAJCQ. 'MJJGAJC ACJJQ NPMBSAC RFC FMPKMLC GLFG@GL, UFGAF GLFG@GRQ '4) NPMBSARGML. -) ?JQM NJ?WQ ? PMJC GL RFC BCTCJMNKCLR MD MT?, GLBSARGML MD MTSJ?RGML, ?LB QRGKSJ?RGML MD CQRP?BGMJ ?LB NPMECQRCPMLC NPMBSARGML @W RFC MT?PGCQ. &QRP?BGMJ ?LB NPMECQRCPMLC ?PC QRCPMGB FMPKMLCQ RF?R NPCN?PC RFC @MBW DMP NPCEL?LAW. &QRP?BGMJ NPMBSACQ QCAMLB?PW QCV AF?P?ARCPGQRGAQ GL DCK?JCQ, UFGJC @MRF CQRP?BGMJ ?LB NPMECQRCPMLC PCESJ?RC RFC KCLQRPS?J AWAJC. ".+ O;'7/'3 C>)1+ '3* 9.+ M+3897:'1 C>)1+ 5FC ovarian cycle EMTCPLQ RFC NPCN?P?RGML MD CLBMAPGLC RGQQSCQ ?LB PCJC?QC MD CEEQ, UFGJC RFC menstrual cycle EMTCPLQ RFC NPCN?P?RGML ?LB K?GLRCL?LAC MD RFC SRCPGLC JGLGLE. 5FCQC AWAJCQ MAASP AMLASPPCLRJW ?LB ?PC AMMPBGL?RCB MTCP ? 22p32 B?W AWAJC, UGRF ?L ?TCP?EC JCLERF MD 28 B?WQ. 5FC DGPQR F?JD MD RFC MT?PG?L AWAJC GQ RFC DMJJGASJ?P NF?QC QFMUL GL Figure 43.15. 4JMUJW PGQGLE JCTCJQ MD '4) ?LB -) A?SQC RFC EPMURF MD DMJJGAJCQ ML RFC QSPD?AC MD RFC MT?PW. 5FGQ NPMACQQ NPCN?PCQ RFC CEE DMP MTSJ?RGML. "Q RFC DMJJGAJCQ EPMU, RFCW @CEGL PCJC?QGLE CQRPMECLQ ?LB ? JMU JCTCJ MD NPMECQRCPMLC. 1PMECQRCPMLC K?GLR?GLQ RFC CLBMKCRPGSK RM FCJN CLQSPC NPCEL?LAW. 5FC RPGN RFPMSEF RFC D?JJMNG?L RS@C R?ICQ ?@MSR QCTCL B?WQ. "R RFGQ QR?EC MD BCTCJMNKCLR, A?JJCB RFC KMPSJ?, RFCPC ?PC 30-60 ACJJQ. *D NPCEL?LAW GKNJ?LR?RGML BMCQ LMR MAASP, RFC JGLGLE GQ QJMSEFCB MDD. "DRCP ?@MSR DGTC B?WQ, CQRPMECL JCTCJQ PGQC ?LB RFC KCLQRPS?J AWAJC CLRCPQ RFC NPMJGDCP?RGTC NF?QC. 5FC CLBMKCRPGSK @CEGLQ RM PCEPMU, PCNJ?AGLE RFC @JMMB TCQQCJQ ?LB EJ?LBQ RF?R BCRCPGMP?RCB BSPGLE RFC CLB MD RFC J?QR AWAJC.
1262
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1264
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5UCLRW-DMSP FMSPQ @CDMPC DCPRGJGX?RGML, RFC CEE F?Q DGLGQFCB KCGMQGQ ?LB @CAMKCQ ? K?RSPC MMAWRC. 8FCL DCPRGJGXCB (?R AMLACNRGML) RFC CEE @CAMKCQ ILMUL ?Q ? XWEMRC. 5FC XWEMRC RP?TCJQ RFPMSEF RFC MTGBSAR RM RFC SRCPSQ (Figure 43.18). 5FC BCTCJMNGLE CK@PWM KSQR GKNJ?LR GLRM RFC U?JJ MD RFC SRCPSQ UGRFGL QCTCL B?WQ, MP GR UGJJ BCRCPGMP?RC ?LB BGC. 5FC MSRCP J?WCPQ MD RFC XWEMRC (@J?QRMAWQR) EPMU GLRM RFC CLBMKCRPGSK @W BGECQRGLE RFC CLBMKCRPG?J ACJJQ, ?LB UMSLB FC?JGLE MD RFC CLBMKCRPGSK AJMQCQ SN RFC @J?QRMAWQR GLRM RFC RGQQSC. "LMRFCP J?WCP MD RFC @J?QRMAWQR, RFC AFMPGML, @CEGLQ PCJC?QGLE ? FMPKMLC A?JJCB human beta chorionic gonadotropin (-HCG) UFGAF K?ICQ GRQ U?W RM RFC AMPNSQ JSRCSK ?LB ICCNQ RF?R QRPSARSPC ?ARGTC. 5FGQ CLQSPCQ ?BCOS?RC JCTCJQ MD NPMECQRCPMLC RF?R UGJJ K?GLR?GL RFC CLBMKCRPGSK MD RFC SRCPSQ DMP RFC QSNNMPR MD RFC BCTCJMNGLE CK@PWM. 1PCEL?LAW RCQRQ BCRCPKGLC RFC JCTCJ MD D-)$( GL SPGLC MP QCPSK. *D RFC FMPKMLC GQ NPCQCLR, RFC RCQR GQ NMQGRGTC.
1266
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5FC ECQR?RGML NCPGMB GQ BGTGBCB GLRM RFPCC COS?J NCPGMBQ MP RPGKCQRCPQ. %SPGLE RFC DGPQR RUM RM DMSP UCCIQ MD RFC DGPQR RPGKCQRCP, LSRPGRGML ?LB U?QRC ?PC F?LBJCB @W RFC CLBMKCRPG?J JGLGLE RFPMSEF BGDDSQGML. "Q RFC RPGKCQRCP NPMEPCQQCQ, RFC MSRCP J?WCP MD RFC CK@PWM @CEGLQ RM KCPEC UGRF RFC CLBMKCRPGSK, ?LB RFC placenta DMPKQ. 5FGQ MPE?L R?ICQ MTCP RFC LSRPGCLR ?LB U?QRC PCOSGPCKCLRQ MD RFC CK@PWM ?LB DCRSQ, UGRF RFC KMRFCPsQ @JMMB N?QQGLE LSRPGCLRQ RM RFC NJ?ACLR? ?LB PCKMTGLE U?QRC DPMK GR. $FCKGA?JQ DPMK RFC DCRSQ, QSAF ?Q @GJGPS@GL, ?PC NPMACQQCB @W RFC KMRFCPsQ JGTCP DMP CJGKGL?RGML. 4MKC MD RFC KMRFCPsQ GKKSLMEJM@SJGLQ UGJJ N?QQ RFPMSEF RFC NJ?ACLR?, NPMTGBGLE N?QQGTC GKKSLGRW ?E?GLQR QMKC NMRCLRG?J GLDCARGMLQ. *LRCPL?J MPE?LQ ?LB @MBW QRPSARSPCQ @CEGL RM BCTCJMN BSPGLE RFC DGPQR RPGKCQRCP. #W DGTC UCCIQ, JGK@ @SBQ, CWCQ, RFC FC?PR, ?LB JGTCP F?TC @CCL @?QGA?JJW DMPKCB. #W CGEFR UCCIQ, RFC RCPK DCRSQ ?NNJGCQ, ?LB RFC @MBW GQ CQQCLRG?JJW DMPKCB, ?Q QFMUL GL Figure 43.19. 5FC GLBGTGBS?J GQ ?@MSR DGTC ACLRGKCRCPQ (RUM GLAFCQ) GL JCLERF ?LB K?LW MD RFC MPE?LQ, QSAF ?Q RFC JSLEQ ?LB JGTCP, ?PC LMR WCR DSLARGMLGLE. &VNMQSPC RM ?LW RMVGLQ GQ CQNCAG?JJW B?LECPMSQ BSPGLE RFC DGPQR RPGKCQRCP, ?Q ?JJ MD RFC @MBWsQ MPE?LQ ?LB QRPSARSPCQ ?PC EMGLE RFPMSEF GLGRG?J BCTCJMNKCLR. "LWRFGLE RF?R ?DDCARQ RF?R BCTCJMNKCLR A?L F?TC ? QCTCPC CDDCAR ML RFC DCRSQs QSPTGT?J.
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!@>LI< 43.19 &BQ>I ABSBILMJBKQ FP PELTK >Q KFKB TBBHP DBPQ>QFLK. (@OBAFQ: %A 5QEJ>K)
%SPGLE RFC QCAMLB RPGKCQRCP, RFC DCRSQ EPMUQ RM ?@MSR 30 AK (12 GLAFCQ), ?Q QFMUL GL Figure 43.20. *R @CAMKCQ ?ARGTC ?LB RFC KMRFCP SQS?JJW DCCJQ RFC DGPQR KMTCKCLRQ. "JJ MPE?LQ ?LB QRPSARSPCQ AMLRGLSC RM BCTCJMN. 5FC NJ?ACLR? F?Q R?ICL MTCP RFC DSLARGMLQ MD LSRPGRGML ?LB U?QRC ?LB RFC NPMBSARGML MD CQRPMECL ?LB NPMECQRCPMLC DPMK RFC AMPNSQ JSRCSK, UFGAF F?Q BCECLCP?RCB. 5FC NJ?ACLR? UGJJ AMLRGLSC DSLARGMLGLE SN RFPMSEF RFC BCJGTCPW MD RFC @?@W.
!@>LI< 43.20 4EFP CBQRP FP GRPQ BKQBOFKD QEB PB@LKA QOFJBPQBO, TEBK QEB MI>@BKQ> Q>HBP LSBO JLOB LC QEB CRK@QFLKP MBOCLOJBA >P QEB ?>?V ABSBILMP. (@OBAFQ: .>QFLK>I -RPBRJ LC (B>IQE >KA -BAF@FKB)
%SPGLE RFC RFGPB RPGKCQRCP, RFC DCRSQ EPMUQ RM 3 RM 4 IE (6 _ -8 _ J@Q.) ?LB ?@MSR 50 AK (19-20 GLAFCQ) JMLE, ?Q GJJSQRP?RCB GL Figure 43.21. 5FGQ GQ RFC NCPGMB MD RFC KMQR P?NGB EPMURF BSPGLE RFC NPCEL?LAW. 0PE?L BCTCJMNKCLR AMLRGLSCQ RM @GPRF (?LB QMKC QWQRCKQ, QSAF ?Q RFC LCPTMSQ QWQRCK ?LB JGTCP, AMLRGLSC RM BCTCJMN ?DRCP @GPRF). 5FC KMRFCP UGJJ @C ?R FCP KMQR SLAMKDMPR?@JC BSPGLE RFGQ RPGKCQRCP. 4FC K?W SPGL?RC DPCOSCLRJW BSC RM NPCQQSPC ML RFC @J?BBCP DPMK RFC DCRSQ. 5FCPC K?W ?JQM @C GLRCQRGL?J @JMAI?EC ?LB AGPASJ?RMPW NPM@JCKQ, CQNCAG?JJW GL FCP JCEQ. $JMRQ K?W DMPK GL FCP JCEQ BSC RM NPCQQSPC DPMK RFC DCRSQ ML PCRSPLGLE TCGLQ ?Q RFCW CLRCP RFC ?@BMKGL?J A?TGRW.
1268
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Table 43.3 JGQRQ AMKKML KCRFMBQ MD AMLRP?ACNRGML. 5FC D?GJSPC P?RCQ JGQRCB ?PC LMR RFC GBC?J P?RCQ RF?R AMSJB @C PC?JGXCB, @SR RFC RWNGA?J P?RCQ RF?R MAASP. " D?GJSPC P?RC GQ RFC LSK@CP MD NPCEL?LAGCQ PCQSJRGLE DPMK RFC KCRFMBsQ SQC MTCP ? RUCJTC-KMLRF NCPGMB. #?PPGCP KCRFMBQ, QSAF ?Q AMLBMKQ, ACPTGA?J A?NQ, ?LB BG?NFP?EKQ, @JMAI QNCPK DPMK CLRCPGLE RFC SRCPSQ, NPCTCLRGLE DCPRGJGX?RGML. 4NCPKGAGBCQ ?PC AFCKGA?JQ RF?R ?PC NJ?ACB GL RFC T?EGL? RF?R IGJJ QNCPK. 4NMLECQ, UFGAF ?PC Q?RSP?RCB UGRF QNCPKGAGBCQ, ?PC NJ?ACB GL RFC T?EGL? ?R RFC ACPTGA?J MNCLGLE. $MK@GL?RGMLQ MD QNCPKGAGB?J AFCKGA?JQ ?LB @?PPGCP KCRFMBQ ?AFGCTC JMUCP D?GJSPC P?RCQ RF?L BM RFC KCRFMBQ UFCL SQCB QCN?P?RCJW. /C?PJW ? OS?PRCP MD RFC AMSNJCQ SQGLE @?PPGCP KCRFMBQ, L?RSP?J D?KGJW NJ?LLGLE, MP UGRFBP?U?J A?L CVNCAR ? D?GJSPC MD RFC KCRFMB. /?RSP?J D?KGJW NJ?LLGLE GQ @?QCB ML RFC KMLGRMPGLE MD RFC KCLQRPS?J AWAJC ?LB F?TGLE GLRCPAMSPQC MLJW BSPGLE RGKCQ UFCL RFC CEE GQ LMR ?T?GJ?@JC. " UMK?LsQ @MBW RCKNCP?RSPC K?W PGQC ? BCEPCC $CJQGSQ ?R MTSJ?RGML ?LB RFC ACPTGA?J KSASQ K?W GLAPC?QC GL TMJSKC ?LB @CAMKC KMPC NJG?@JC. 5FCQC AF?LECQ EGTC ? ECLCP?J GLBGA?RGML MD UFCL GLRCPAMSPQC GQ KMPC MP JCQQ JGICJW RM PCQSJR GL DCPRGJGX?RGML. 8GRFBP?U?J GLTMJTCQ RFC PCKMT?J MD RFC NCLGQ DPMK RFC T?EGL? BSPGLE GLRCPAMSPQC, @CDMPC CH?ASJ?RGML MAASPQ. 5FGQ GQ ? PGQIW KCRFMB UGRF ? FGEF D?GJSPC P?RC BSC RM RFC NMQQG@JC NPCQCLAC MD QNCPK GL RFC @SJ@MSPCRFP?J EJ?LBsQ QCAPCRGML, UFGAF K?W CLRCP RFC T?EGL? NPGMP RM PCKMTGLE RFC NCLGQ. )MPKML?J KCRFMBQ SQC QWLRFCRGA NPMECQRCPMLC (QMKCRGKCQ GL AMK@GL?RGML UGRF CQRPMECL), RM GLFG@GR RFC FWNMRF?J?KSQ DPMK PCJC?QGLE '4) MP -), ?LB RFSQ NPCTCLR ?L CEE DPMK @CGLE ?T?GJ?@JC DMP DCPRGJGX?RGML. 5FC KCRFMB MD ?BKGLGQRCPGLE RFC FMPKMLC ?DDCARQ D?GJSPC P?RC. 5FC KMQR PCJG?@JC KCRFMB, UGRF ? D?GJSPC P?RC MD JCQQ RF?L 1 NCPACLR, GQ RFC GKNJ?LR?RGML MD RFC FMPKMLC SLBCP RFC QIGL. 5FC Q?KC P?RC A?L @C ?AFGCTCB RFPMSEF RFC QRCPGJGX?RGML NPMACBSPCQ MD T?QCARMKW GL RFC K?L MP MD RS@?J JGE?RGML GL RFC UMK?L, MP @W SQGLE ?L GLRP?SRCPGLC BCTGAC (*6%). *6%Q ?PC GLQCPRCB GLRM RFC SRCPSQ ?LB CQR?@JGQF ?L GLDJ?KK?RMPW AMLBGRGML RF?R NPCTCLRQ DCPRGJGXCB CEEQ DPMK GKNJ?LRGLE GLRM RFC SRCPGLC U?JJ. $MKNJG?LAC UGRF RFC AMLRP?ACNRGTC KCRFMB GQ ? QRPMLE AMLRPG@SRMP RM RFC QSAACQQ MP D?GJSPC P?RC MD ?LW N?PRGASJ?P KCRFMB. 5FC MLJW KCRFMB RF?R GQ AMKNJCRCJW CDDCARGTC ?R NPCTCLRGLE AMLACNRGML GQ ?@QRGLCLAC. 5FC AFMGAC MD AMLRP?ACNRGTC KCRFMB BCNCLBQ ML RFC EM?JQ MD RFC UMK?L MP AMSNJC. 5S@?J JGE?RGML ?LB T?QCARMKW ?PC AMLQGBCPCB NCPK?LCLR NPCTCLRGML, UFGJC MRFCP KCRFMBQ ?PC PCTCPQG@JC ?LB NPMTGBC QFMPR-RCPK AMLRP?ACNRGML. 5CPKGL?RGML MD ?L CVGQRGLE NPCEL?LAW A?L @C QNMLR?LCMSQ MP TMJSLR?PW. 4NMLR?LCMSQ RCPKGL?RGML GQ ? KGQA?PPG?EC ?LB SQS?JJW MAASPQ TCPW C?PJW GL RFC NPCEL?LAW, SQS?JJW UGRFGL RFC DGPQR DCU UCCIQ. 5FGQ MAASPQ UFCL RFC DCRSQ A?LLMR BCTCJMN NPMNCPJW ?LB RFC ECQR?RGML GQ L?RSP?JJW RCPKGL?RCB. 7MJSLR?PW RCPKGL?RGML MD ? NPCEL?LAW GQ ?L ?@MPRGML. -?UQ PCESJ?RGLE ?@MPRGML T?PW @CRUCCL QR?RCQ ?LB RCLB RM TGCU DCR?J TG?@GJGRW ?Q RFC APGRCPG? DMP ?JJMUGLE MP NPCTCLRGLE RFC NPMACBSPC.
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Infertility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sQ SRCPSQ. *D DCPRGJGX?RGML GQ LMR ?AAMKNJGQFCB @W QGKNJC *7', ? NPMACBSPC RF?R GLHCARQ RFC QNCPK GLRM ?L CEE A?L @C SQCB. 5FGQ GQ A?JJCB GLRP?AWRMNJ?QKGA QNCPK GLHCARGML (*$4*) ?LB GQ QFMUL GL Figure 43.22. *7' NPMACBSPCQ NPMBSAC ? QSPNJSQ MD DCPRGJGXCB CEEQ ?LB CK@PWMQ RF?R A?L @C DPMXCL ?LB QRMPCB DMP DSRSPC SQC. 5FC NPMACBSPCQ A?L ?JQM PCQSJR GL KSJRGNJC @GPRFQ.
!@>LI< 43.22 ! PMBOJ FP FKPBOQBA FKQL >K BDD CLO CBOQFIFW>QFLK AROFKD FKQO>@VQLMI>PJF@ PMBOJ FKGB@QFLK ()#3)). (@OBAFQ: P@>IB-?>O A>Q> COLJ ->QQ 2RPPBII)
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %GQASQQ FMU DCPRGJGX?RGML MAASPQ z &VNJ?GL FMU RFC CK@PWM DMPKQ DPMK RFC XWEMRC z %GQASQQ RFC PMJC MD AJC?T?EC ?LB E?QRPSJ?RGML GL ?LGK?J BCTCJMNKCLR 5FC NPMACQQ GL UFGAF ?L MPE?LGQK BCTCJMNQ DPMK ? QGLEJC-ACJJCB XWEMRC RM ? KSJRG-ACJJSJ?P MPE?LGQK GQ AMKNJCV ?LB UCJJ-PCESJ?RCB. 5FC C?PJW QR?ECQ MD CK@PWMLGA BCTCJMNKCLR ?PC ?JQM APSAG?J DMP CLQSPGLE RFC DGRLCQQ MD RFC MPE?LGQK.
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'CPRGJGX?RGML, NGARSPCB GL Figure 43.23a GQ RFC NPMACQQ GL UFGAF E?KCRCQ (?L CEE ?LB QNCPK) DSQC RM DMPK ? XWEMRC. 5FC CEE ?LB QNCPK C?AF AMLR?GL MLC QCR MD AFPMKMQMKCQ. 5M CLQSPC RF?R RFC MDDQNPGLE F?Q MLJW MLC AMKNJCRC BGNJMGB QCR MD AFPMKMQMKCQ, MLJW MLC QNCPK KSQR DSQC UGRF MLC CEE. *L K?KK?JQ, RFC CEE GQ NPMRCARCB @W ? J?WCP MD CVRP?ACJJSJ?P K?RPGV AMLQGQRGLE K?GLJW MD EJWAMNPMRCGLQ A?JJCB RFC zona pellucida. 8FCL ? QNCPK @GLBQ RM RFC XML? NCJJSAGB?, ? QCPGCQ MD @GMAFCKGA?J CTCLRQ, A?JJCB RFC
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acrosomal reactions, R?IC NJ?AC. *L NJ?ACLR?J K?KK?JQ, RFC ?APMQMKC AMLR?GLQ BGECQRGTC CLXWKCQ RF?R GLGRG?RC RFC BCEP?B?RGML MD RFC EJWAMNPMRCGL K?RPGV NPMRCARGLE RFC CEE ?LB ?JJMUGLE RFC QNCPK NJ?QK? KCK@P?LC RM DSQC UGRF RFC CEE NJ?QK? KCK@P?LC, ?Q GJJSQRP?RCB GL Figure 43.23b. 5FC DSQGML MD RFCQC RUM KCK@P?LCQ APC?RCQ ?L MNCLGLE RFPMSEF UFGAF RFC QNCPK LSAJCSQ GQ RP?LQDCPPCB GLRM RFC MTSK. 5FC LSAJC?P KCK@P?LCQ MD RFC CEE ?LB QNCPK @PC?I BMUL ?LB RFC RUM F?NJMGB ECLMKCQ AMLBCLQC RM DMPK ? BGNJMGB ECLMKC.
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5M CLQSPC RF?R LM KMPC RF?L MLC QNCPK DCPRGJGXCQ RFC CEE, MLAC RFC ?APMQMK?J PC?ARGMLQ R?IC NJ?AC ?R MLC JMA?RGML MD RFC CEE KCK@P?LC, RFC CEE PCJC?QCQ NPMRCGLQ GL MRFCP JMA?RGMLQ RM NPCTCLR MRFCP QNCPK DPMK DSQGLE UGRF RFC CEE. *D RFGQ KCAF?LGQK D?GJQ, KSJRGNJC QNCPK A?L DSQC UGRF RFC CEE, PCQSJRGLE GL polyspermy. 5FC PCQSJRGLE CK@PWM GQ LMR ECLCRGA?JJW TG?@JC ?LB BGCQ UGRFGL ? DCU B?WQ.
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5FC BCTCJMNKCLR MD KSJRG-ACJJSJ?P MPE?LGQKQ @CEGLQ DPMK ? QGLEJC-ACJJCB XWEMRC, UFGAF SLBCPEMCQ P?NGB ACJJ BGTGQGML RM DMPK RFC @J?QRSJ?. 5FC P?NGB, KSJRGNJC PMSLBQ MD ACJJ BGTGQGML ?PC RCPKCB AJC?T?EC. $JC?T?EC GQ GJJSQRP?RCB GL (Figure 43.24a). "DRCP RFC AJC?T?EC F?Q NPMBSACB MTCP 100 ACJJQ, RFC CK@PWM GQ A?JJCB ? @J?QRSJ?. 5FC @J?QRSJ? GQ SQS?JJW ? QNFCPGA?J J?WCP MD ACJJQ (RFC @J?QRMBCPK) QSPPMSLBGLE ? DJSGBDGJJCB MP WMJI-DGJJCB A?TGRW (RFC @J?QRMAMCJ). .?KK?JQ ?R RFGQ QR?EC DMPK ? QRPSARSPC A?JJCB RFC @J?QRMAWQR, AF?P?ARCPGXCB @W ?L GLLCP ACJJ K?QQ RF?R GQ BGQRGLAR DPMK RFC QSPPMSLBGLE @J?QRSJ?, QFMUL GL Figure 43.24b. %SPGLE AJC?T?EC, RFC ACJJQ BGTGBC UGRFMSR ?L GLAPC?QC GL K?QQ; RF?R GQ, MLC J?PEC QGLEJC-ACJJCB XWEMRC BGTGBCQ GLRM KSJRGNJC QK?JJCP ACJJQ. &?AF ACJJ UGRFGL RFC @J?QRSJ? GQ A?JJCB ? @J?QRMKCPC.
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$JC?T?EC A?L R?IC NJ?AC GL RUM U?WQ: holoblastic (RMR?J) AJC?T?EC MP meroblastic (N?PRG?J) AJC?T?EC. 5FC RWNC MD AJC?T?EC BCNCLBQ ML RFC ?KMSLR MD WMJI GL RFC CEEQ. *L NJ?ACLR?J K?KK?JQ (GLAJSBGLE FSK?LQ) UFCPC LMSPGQFKCLR GQ NPMTGBCB @W RFC KMRFCPsQ @MBW, RFC CEEQ F?TC ? TCPW QK?JJ ?KMSLR MD WMJI ?LB SLBCPEM FMJM@J?QRGA AJC?T?EC. 0RFCP QNCAGCQ, QSAF ?Q @GPBQ, UGRF ? JMR MD WMJI GL RFC CEE RM LMSPGQF RFC CK@PWM BSPGLE BCTCJMNKCLR, SLBCPEM KCPM@J?QRGA AJC?T?EC. *L K?KK?JQ, RFC @J?QRSJ? DMPKQ RFC blastocyst GL RFC LCVR QR?EC MD BCTCJMNKCLR. )CPC RFC ACJJQ GL RFC @J?QRSJ? ?PP?LEC RFCKQCJTCQ GL RUM J?WCPQ: RFC inner cell mass, ?LB ?L MSRCP J?WCP A?JJCB RFC trophoblast. 5FC GLLCP ACJJ K?QQ GQ ?JQM ILMUL ?Q RFC CK@PWM@J?QR ?LB RFGQ K?QQ MD ACJJQ UGJJ EM ML RM DMPK RFC CK@PWM. "R RFGQ QR?EC MD BCTCJMNKCLR, GJJSQRP?RCB GL Figure 43.25 RFC GLLCP ACJJ K?QQ AMLQGQRQ MD CK@PWMLGA QRCK ACJJQ RF?R UGJJ BGDDCPCLRG?RC GLRM RFC BGDDCPCLR ACJJ RWNCQ LCCBCB @W RFC MPE?LGQK. 5FC RPMNFM@J?QR UGJJ AMLRPG@SRC RM RFC NJ?ACLR? ?LB LMSPGQF RFC CK@PWM.
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43.7 R *I>8EF><E<J@J 8E; 1<IK<9I8K< !FID8K@FE

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*L TCPRC@P?RCQ, MLC MD RFC NPGK?PW QRCNQ BSPGLE MPE?LMECLCQGQ GQ RFC DMPK?RGML MD RFC LCSP?J QWQRCK. 5FC CARMBCPK DMPKQ CNGRFCJG?J ACJJQ ?LB RGQQSCQ, ?LB LCSPML?J RGQQSCQ. %SPGLE RFC DMPK?RGML MD RFC LCSP?J QWQRCK, QNCAG?J QGEL?JGLE KMJCASJCQ A?JJCB EPMURF D?ARMPQ QGEL?J QMKC ACJJQ ?R RFC CBEC MD RFC CARMBCPK RM @CAMKC CNGBCPKGQ ACJJQ. 5FC PCK?GLGLE ACJJQ GL RFC ACLRCP DMPK RFC LCSP?J NJ?RC. *D RFC QGEL?JGLE @W EPMURF D?ARMPQ UCPC BGQPSNRCB, RFCL RFC CLRGPC CARMBCPK UMSJB BGDDCPCLRG?RC GLRM LCSP?J RGQQSC. 5FC LCSP?J NJ?RC SLBCPEMCQ ? QCPGCQ MD ACJJ KMTCKCLRQ UFCPC GR PMJJQ SN ?LB DMPKQ ? RS@C A?JJCB RFC neural tube, ?Q GJJSQRP?RCB GL Figure 43.28. *L DSPRFCP BCTCJMNKCLR, RFC LCSP?J RS@C UGJJ EGTC PGQC RM RFC @P?GL ?LB RFC QNGL?J AMPB.
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1276
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1278
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K<JK<J N?GP MD PCNPMBSARGTC MPE?LQ GL K?JCQ K<JKFJK<IFE< PCNPMBSARGTC FMPKMLC GL KCL RF?R ?QQGQRQ GL QNCPK NPMBSARGML ?LB NPMKMRGLE QCAMLB?PW QCVS?J AF?P?ARCPGQRGAQ KIFG?F9C8JK MSRCP J?WCP MD ACJJQ GL RFC @J?QRMAWQR LK<ILJ CLTGPMLKCLR DMP BCTCJMNGLE CK@PWM ?LB DCRSQ M8>@E8 KSQASJ?P RS@C DMP RFC N?QQ?EC MD KCLQRPS?J DJMU, AMNSJ?RGML, ?LB @GPRF MD MDDQNPGLE M@M@G8I@KP NPMACQQ GL UFGAF RFC WMSLE BCTCJMN UGRFGL RFC DCK?JC, PCACGTGLE LMSPGQFKCLR DPMK RFC KMRFCPsQ @JMMB RFPMSEF ? NJ?ACLR? QFE8 G<CCL:@;8 NPMRCARGTC J?WCP MD EJWAMNPMRCGLQ ML RFC K?KK?JG?L CEE
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3CNPMBSARGML K?W @C ?QCVS?J UFCL MLC GLBGTGBS?J NPMBSACQ ECLCRGA?JJW GBCLRGA?J MDDQNPGLE, MP QCVS?J UFCL RFC ECLCRGA K?RCPG?J DPMK RUM GLBGTGBS?JQ GQ AMK@GLCB RM NPMBSAC ECLCRGA?JJW BGTCPQC MDDQNPGLE. "QCVS?J PCNPMBSARGML MAASPQ RFPMSEF DGQQGML, @SBBGLE, ?LB DP?EKCLR?RGML. 4CVS?J PCNPMBSARGML K?W KC?L RFC HMGLGLE MD QNCPK ?LB CEEQ UGRFGL ?LGK?JQs @MBGCQ MP GR K?W KC?L RFC PCJC?QC MD QNCPK ?LB CEEQ GLRM RFC CLTGPMLKCLR. "L GLBGTGBS?J K?W @C MLC QCV, MP @MRF; GR K?W QR?PR MSR ?Q MLC QCV ?LB QUGRAF BSPGLE GRQ JGDC, MP GR K?W QR?W K?JC MP DCK?JC.
43.2 !<IK@C@Q8K@FE
4CVS?J PCNPMBSARGML QR?PRQ UGRF RFC AMK@GL?RGML MD ? QNCPK ?LB ?L CEE GL ? NPMACQQ A?JJCB DCPRGJGX?RGML. 5FGQ A?L MAASP CGRFCP MSRQGBC RFC @MBGCQ MP GLQGBC RFC DCK?JC. #MRF KCRFMBQ F?TC ?BT?LR?ECQ ?LB BGQ?BT?LR?ECQ. 0LAC DCPRGJGXCB, RFC CEEQ A?L BCTCJMN GLQGBC RFC DCK?JC MP MSRQGBC. *D RFC CEE BCTCJMNQ MSRQGBC RFC @MBW, GR SQS?JJW F?Q ? NPMRCARGTC AMTCPGLE MTCP GR. "LGK?J ?L?RMKW CTMJTCB T?PGMSQ U?WQ RM DCPRGJGXC, FMJB, MP CVNCJ RFC CEE. 5FC KCRFMB MD DCPRGJGX?RGML T?PGCQ ?KMLE ?LGK?JQ. 4MKC QNCAGCQ PCJC?QC RFC CEE ?LB QNCPK GLRM RFC CLTGPMLKCLR, QMKC QNCAGCQ PCR?GL RFC CEE ?LB PCACGTC RFC QNCPK GLRM RFC DCK?JC @MBW ?LB RFCL CVNCJ RFC BCTCJMNGLE CK@PWM AMTCPCB UGRF QFCJJ, UFGJC QRGJJ MRFCP QNCAGCQ PCR?GL RFC BCTCJMNGLE MDDQNPGLE RFPMSEF RFC ECQR?RGML NCPGMB.
43.3 #LD8E -<GIF;L:K@M< AE8KFDP 8E; "8D<KF><E<J@J

"Q ?LGK?JQ @CA?KC KMPC AMKNJCV, QNCAGDGA MPE?LQ ?LB MPE?L QWQRCKQ BCTCJMNCB RM QSNNMPR QNCAGDGA DSLARGMLQ DMP RFC MPE?LGQK. 5FC PCNPMBSARGTC QRPSARSPCQ RF?R CTMJTCB GL J?LB ?LGK?JQ ?JJMU K?JCQ ?LB DCK?JCQ RM K?RC, DCPRGJGXC GLRCPL?JJW, ?LB QSNNMPR RFC EPMURF ?LB BCTCJMNKCLR MD MDDQNPGLE. 1PMACQQCQ BCTCJMNCB RM NPMBSAC PCNPMBSARGTC ACJJQ RF?R F?B CV?ARJW F?JD RFC LSK@CP MD AFPMKMQMKCQ MD C?AF N?PCLR QM RF?R LCU AMK@GL?RGMLQ UMSJB F?TC RFC ?NNPMNPG?RC ?KMSLR MD ECLCRGA K?RCPG?J. (?KCRMECLCQGQ, RFC NPMBSARGML MD QNCPK (QNCPK?RMECLCQGQ) ?LB CEEQ (MMECLCQGQ), R?ICQ NJ?AC RFPMSEF RFC NPMACQQ MD KCGMQGQ.
43.4 #FIDFE8C CFEKIFC F= #LD8E -<GIF;L:K@FE

5FC K?JC ?LB DCK?JC PCNPMBSARGTC AWAJCQ ?PC AMLRPMJJCB @W FMPKMLCQ PCJC?QCB DPMK RFC FWNMRF?J?KSQ ?LB ?LRCPGMP NGRSGR?PW ?Q UCJJ ?Q FMPKMLCQ DPMK PCNPMBSARGTC RGQQSCQ ?LB MPE?LQ. 5FC FWNMRF?J?KSQ KMLGRMPQ RFC LCCB DMP RFC '4) ?LB -) FMPKMLCQ K?BC ?LB PCJC?QCB DPMK RFC ?LRCPGMP NGRSGR?PW. '4) ?LB -) ?DDCAR PCNPMBSARGTC QRPSARSPCQ RM A?SQC RFC DMPK?RGML MD QNCPK ?LB RFC NPCN?P?RGML MD CEEQ DMP PCJC?QC ?LB NMQQG@JC DCPRGJGX?RGML. *L RFC K?JC, '4) ?LB -) QRGKSJ?RC 4CPRMJG ACJJQ ?LB GLRCPQRGRG?J ACJJQ MD -CWBGE GL RFC RCQRCQ RM D?AGJGR?RC QNCPK NPMBSARGML. 5FC -CWBGE ACJJQ NPMBSAC RCQRMQRCPMLC, UFGAF ?JQM GQ PCQNMLQG@JC DMP RFC QCAMLB?PW QCVS?J AF?P?ARCPGQRGAQ MD K?JCQ. *L DCK?JCQ, '4) ?LB -) A?SQC CQRPMECL ?LB NPMECQRCPMLC RM @C NPMBSACB. 5FCW PCESJ?RC RFC DCK?JC PCNPMBSARGTC QWQRCK UFGAF GQ BGTGBCB GLRM RFC MT?PG?L AWAJC ?LB RFC KCLQRPS?J AWAJC. .CLMN?SQC MAASPQ UFCL RFC MT?PGCQ JMQC RFCGP QCLQGRGTGRW RM '4) ?LB -) ?LB RFC DCK?JC PCNPMBSARGTC AWAJCQ QJMU RM ? QRMN.
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)SK?L NPCEL?LAW @CEGLQ UGRF DCPRGJGX?RGML MD ?L CEE ?LB NPMACCBQ RFPMSEF RFC RFPCC RPGKCQRCPQ MD ECQR?RGML. 5FC J?@MP NPMACQQ F?Q RFPCC QR?ECQ (AMLRP?ARGMLQ, BCJGTCPW MD RFC DCRSQ, CVNSJQGML MD RFC NJ?ACLR?), C?AF NPMNCJJCB @W FMPKMLCQ. 5FC DGPQR RPGKCQRCP J?WQ BMUL RFC @?QGA QRPSARSPCQ MD RFC @MBW, GLAJSBGLE RFC JGK@ @SBQ, FC?PR, CWCQ, ?LB RFC JGTCP. 5FC QCAMLB RPGKCQRCP AMLRGLSCQ RFC BCTCJMNKCLR MD ?JJ MD RFC MPE?LQ ?LB QWQRCKQ. 5FC RFGPB RPGKCQRCP CVFG@GRQ RFC EPC?RCQR EPMURF MD RFC DCRSQ ?LB ASJKGL?RCQ GL J?@MP ?LB BCJGTCPW. 1PCTCLRGML MD ? NPCEL?LAW A?L @C ?AAMKNJGQFCB RFPMSEF ? T?PGCRW MD KCRFMBQ GLAJSBGLE @?PPGCPQ, FMPKMLCQ, MP MRFCP KC?LQ. "QQGQRCB PCNPMBSARGTC RCAFLMJMEGCQ K?W FCJN GLBGTGBS?JQ UFM F?TC GLDCPRGJGRW NPM@JCKQ.
43.6 !<IK@C@Q8K@FE 8E; 8ICP D9IPFE@: D<M<CFGD<EK

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
43.7 *I>8EF><E<J@J 8E; 1<IK<9I8K< !FID8K@FE

0PE?LMECLCQGQ GQ RFC DMPK?RGML MD MPE?LQ DPMK RFC ECPK J?WCPQ. &?AF ECPK J?WCP EGTCQ PGQC RM QNCAGDGA RGQQSC RWNCQ. 5FC DGPQR QR?EC GQ RFC DMPK?RGML MD RFC LCSP?J QWQRCK GL RFC CARMBCPK. 5FC KCQMBCPK EGTCQ PGQC RM QMKGRCQ ?LB RFC LMRMAFMPB. 'MPK?RGML MD TCPRC@P?RC ?VGQ GQ ?LMRFCP GKNMPR?LR BCTCJMNKCLR?J QR?EC.
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1. Figure 43.8 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR RFC K?JC PCNPMBSARGTC QWQRCK GQ D?JQC >. 5FC T?Q BCDCPCLQ A?PPGCQ QNCPK DPMK RFC RCQRCQ RM RFC NCLGQ. ?. 4NCPK K?RSPC GL QCKGLGDCPMSQ RS@SJCQ GL RFC RCQRCQ. @. #MRF RFC NPMQR?RC ?LB RFC @SJ@MSPCRFP?J EJ?LBQ NPMBSAC AMKNMLCLRQ MD RFC QCKCL. A. 5FC NPMQR?RC EJ?LB GQ JMA?RCB GL RFC RCQRCQ. 2. Figure 43.15 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR FMPKMLC PCESJ?RGML MD RFC DCK?JC PCNPMBSARGTC AWAJC GQ D?JQC >. -) ?LB '4) ?PC NPMBSACB GL RFC NGRSGR?PW, ?LB CQRP?BGMJ ?LB NPMECQRCPMLC ?PC NPMBSACB GL RFC MT?PGCQ. ?. &QRP?BGMJ ?LB NPMECQRCPMLC QCAPCRCB DPMK RFC AMPNSQ JSRCSK A?SQC RFC CLBMKCRPGSK RM RFGAICL. @. #MRF NPMECQRCPMLC ?LB CQRP?BGMJ ?PC NPMBSACB @W RFC DMJJGAJCQ. A. 4CAPCRGML MD (L3) @W RFC FWNMRF?J?KSQ GQ GLFG@GRCB @W JMU JCTCJQ MD CQRP?BGMJ @SR QRGKSJ?RCB @W FGEF JCTCJQ MD CQRP?BGMJ. 3. Figure 43.17 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR RFC KCLQRPS?J AWAJC GQ D?JQC >. 1PMECQRCPMLC JCTCJQ PGQC BSPGLE RFC JSRC?J NF?QC MD RFC MT?PG?L AWAJC ?LB RFC QCAPCRMPW NF?QC MD RFC SRCPGLC AWAJC. ?. .CLQRPS?RGML MAASPQ HSQR ?DRCP -) ?LB '4) JCTCJQ NC?I. @. .CLQRPS?RGML MAASPQ ?DRCP NPMECQRCPMLC JCTCJQ BPMN. A. &QRPMECL JCTCJQ PGQC @CDMPC MTSJ?RGML, UFGJC NPMECQRCPMLC JCTCJQ PGQC ?DRCP.
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4. 8FGAF DMPK MD PCNPMBSARGML GQ RFMSEFR RM @C @CQR GL ? QR?@JC CLTGPMLKCLR >. ?QCVS?J ?. QCVS?J @. @SBBGLE A. N?PRFCLMECLCQGQ 5. 8FGAF DMPK MD PCNPMBSARGML A?L PCQSJR DPMK B?K?EC RM RFC MPGEGL?J ?LGK?J >. ?. @. A. ?QCVS?J DP?EKCLR?RGML @SBBGLE N?PRFCLMECLCQGQ
6. 8FGAF DMPK MD PCNPMBSARGML GQ SQCDSJ RM ?L ?LGK?J UGRF JGRRJC KM@GJGRW RF?R PCNPMBSACQ QCVS?JJW >. DGQQGML
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A. SRCPSQ 16. )MU K?LW CEEQ ?PC NPMBSACB ?Q ? PCQSJR MD MLC KCGMRGA QCPGCQ MD ACJJ BGTGQGMLQ >. MLC ?. RUM @. RFPCC A. DMSP 17. 8FGAF FMPKMLC A?SQCQ -CWBGE ACJJQ RM K?IC RCQRMQRCPMLC >. '4) ?. -) @. GLFG@GL A. CQRPMECL 18. 8FGAF FMPKMLC A?SQCQ '4) ?LB -) RM @C PCJC?QCB >. RCQRMQRCPMLC ?. CQRPMECL @. (L3) A. NPMECQRCPMLC 19. 8FGAF FMPKMLC QGEL?JQ MTSJ?RGML >. '4) ?. -) @. GLFG@GL A. CQRPMECL 20. 8FGAF FMPKMLC A?SQCQ RFC PC-EPMURF MD RFC CLBMKCRPG?J JGLGLE MD RFC SRCPSQ >. RCQRMQRCPMLC ?. CQRPMECL @. (L3) A. NPMECQRCPMLC 21. /SRPGCLR ?LB U?QRC PCOSGPCKCLRQ DMP RFC BCTCJMNGLE DCRSQ ?PC F?LBJCB BSPGLE RFC DGPQR DCU UCCIQ @W: >. RFC NJ?ACLR? ?. BGDDSQGML RFPMSEF RFC CLBMKCRPGSK @. RFC AFMPGML A. RFC @J?QRMAWQR 22. 1PMECQRCPMLC GQ K?BC BSPGLE RFC RFGPB RPGKCQRCP @W RFC: >. NJ?ACLR? ?. CLBMKCRPG?J JGLGLE @. AFMPGML A. AMPNSQ JSRCSK 23. 8FGAF AMLRP?ACNRGTC KCRFMB GQ 100 NCPACLR CDDCARGTC ?R NPCTCLRGLE NPCEL?LAW >. AMLBMK ?. MP?J FMPKML?J KCRFMBQ @. QRCPGJGX?RGML A. ?@QRGLCLAC 24. 8FGAF RWNC MD QFMPR RCPK AMLRP?ACNRGTC KCRFMB GQ ECLCP?JJW KMPC CDDCARGTC RF?L MRFCPQ >. ?. @. A. @?PPGCP FMPKML?J L?RSP?J D?KGJW NJ?LLGLE UGRFBP?U?J
10. 8FGAF RCPK ?NNJGCQ RM CEE BCTCJMNKCLR MSRQGBC RFC DCK?JC UGRF LMSPGQFKCLR BCPGTCB DPMK ? WMJI >. MTGN?PGRW ?. TGTGN?PGRW @. MTMTGN?PGRW A. MTMTMN?PGRW 11. 4NCPK ?PC NPMBSACB GL RFC >>>>>>>>. >. ?. @. A. QAPMRSK QCKGL?J TCQGAJCQ QCKGLGDCPMSQ RS@SJCQ NPMQR?RC EJ?LB
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
25. 8FGAF FMPKMLC GQ NPGK?PGJW PCQNMLQG@JC DMP RFC AMLRP?ARGMLQ BSPGLE J?@MP
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28. %SPGLE AJC?T?EC, RFC K?QQ MD ACJJQ: >. GLAPC?QCQ ?. BCAPC?QCQ @. BMS@JCQ UGRF CTCPW ACJJ BGTGQGML A. BMCQ LMR AF?LEC QGELGDGA?LRJW 29. 8FGAF MD RFC DMJJMUGLE EGTCQ PGQC RM RFC QIGL ACJJQ >. CARMBCPK ?. CLBMBCPK @. KCQMBCPK A. LMLC MD RFC ?@MTC 30. 5FC PG@Q DMPK DPMK RFC >>>>>>>>. >. LMRMAFMPB ?. LCSP?J NJ?RC @. LCSP?J RS@C A. QMKGRCQ
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1300
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$JGK?RC AF?LEC A?L @C SLBCPQRMMB @W ?NNPM?AFGLE RFPCC ?PC?Q MD QRSBW: z ASPPCLR ?LB N?QR EJM@?J AJGK?RC AF?LEC z A?SQCQ MD N?QR ?LB NPCQCLR-B?W EJM@?J AJGK?RC AF?LEC z ?LAGCLR ?LB ASPPCLR PCQSJRQ MD AJGK?RC AF?LEC *R GQ FCJNDSJ RM ICCN RFCQC RFPCC BGDDCPCLR ?QNCARQ MD AJGK?RC AF?LEC AJC?PJW QCN?P?RCB UFCL AMLQSKGLE KCBG? PCNMPRQ ?@MSR EJM@?J AJGK?RC AF?LEC. *R GQ AMKKML DMP PCNMPRQ ?LB BGQASQQGMLQ ?@MSR EJM@?J AJGK?RC AF?LEC RM AMLDSQC RFC B?R? QFMUGLE RF?R &?PRFsQ AJGK?RC GQ AF?LEGLE UGRF RFC D?ARMPQ RF?R BPGTC RFGQ AJGK?RC AF?LEC. E;/*+3)+ ,47 G14('1 C1/2'9+ C.'3-+ 4GLAC QAGCLRGQRQ A?LLMR EM @?AI GL RGKC RM BGPCARJW KC?QSPC AJGK?RGA T?PG?@JCQ, QSAF ?Q ?TCP?EC RCKNCP?RSPC ?LB NPCAGNGR?RGML, RFCW KSQR GLQRC?B GLBGPCARJW KC?QSPC RCKNCP?RSPC. 5M BM RFGQ, QAGCLRGQRQ PCJW ML FGQRMPGA?J CTGBCLAC MD &?PRFsQ N?QR AJGK?RC. "LR?PARGA GAC AMPCQ ?PC ? ICW CV?KNJC MD QSAF CTGBCLAC. 5FCQC GAC AMPCQ ?PC Q?KNJCQ MD NMJ?P GAC M@R?GLCB @W KC?LQ MD BPGJJQ RF?R PC?AF RFMSQ?LBQ MD KCRCPQ GLRM GAC QFCCRQ MP FGEF KMSLR?GL EJ?AGCPQ. 7GCUGLE RFC GAC AMPCQ GQ JGIC RP?TCJGLE @?AIU?PBQ RFPMSEF RGKC; RFC BCCNCP RFC Q?KNJC, RFC C?PJGCP RFC RGKC NCPGMB. 5P?NNCB UGRFGL RFC GAC ?PC @S@@JCQ MD ?GP ?LB MRFCP @GMJMEGA?J CTGBCLAC RF?R A?L PCTC?J RCKNCP?RSPC ?LB A?P@ML BGMVGBC B?R?. "LR?PARGA GAC AMPCQ F?TC @CCL AMJJCARCB ?LB ?L?JWXCB RM GLBGPCARJW CQRGK?RC RFC RCKNCP?RSPC MD RFC &?PRF MTCP RFC N?QR 400,000 WC?PQ (Figure 44.26a). 5FC 0 \$ ML RFGQ EP?NF PCDCPQ RM RFC JMLE-RCPK ?TCP?EC. 5CKNCP?RSPCQ RF?R ?PC EPC?RCP RF?L 0 \$ CVACCB &?PRFsQ JMLE-RCPK ?TCP?EC RCKNCP?RSPC. $MLTCPQCJW, RCKNCP?RSPCQ RF?R ?PC JCQQ RF?L 0 \$ ?PC JCQQ RF?L &?PRFsQ ?TCP?EC RCKNCP?RSPC. 5FGQ DGESPC QFMUQ RF?R RFCPC F?TC @CCL NCPGMBGA AWAJCQ MD GLAPC?QGLE ?LB BCAPC?QGLE RCKNCP?RSPC. #CDMPC RFC J?RC 1800Q, RFC &?PRF F?Q @CCL ?Q KSAF ?Q 9 \$ AMMJCP ?LB ?@MSR 3 \$ U?PKCP. /MRC RF?R RFC EP?NF GL Figure 44.26b QFMUQ RF?R RFC ?RKMQNFCPGA AMLACLRP?RGML MD A?P@ML BGMVGBC F?Q ?JQM PGQCL ?LB D?JJCL GL NCPGMBGA AWAJCQ; LMRC RFC PCJ?RGMLQFGN @CRUCCL A?P@ML BGMVGBC AMLACLRP?RGML ?LB RCKNCP?RSPC. Figure 44.26b QFMUQ RF?R A?P@ML BGMVGBC JCTCJQ GL RFC ?RKMQNFCPC F?TC FGQRMPGA?JJW AWAJCB @CRUCCL 180 ?LB 300 N?PRQ NCP KGJJGML (NNK) @W TMJSKC.
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Figure 44.26a BMCQ LMR QFMU RFC J?QR 2,000 WC?PQ UGRF CLMSEF BCR?GJ RM AMKN?PC RFC AF?LECQ MD &?PRFs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igure 44.27).
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C:77+39 '3* P'89 D7/;+78 4, G14('1 C1/2'9+ C.'3-+ 4GLAC GR GQ LMR NMQQG@JC RM EM @?AI GL RGKC RM BGPCARJW M@QCPTC ?LB KC?QSPC AJGK?RC, QAGCLRGQRQ SQC GLBGPCAR CTGBCLAC RM BCRCPKGLC RFC BPGTCPQ, MP D?ARMPQ, RF?R K?W @C PCQNMLQG@JC DMP AJGK?RC AF?LEC. 5FC GLBGPCAR CTGBCLAC GLAJSBCQ B?R? AMJJCARCB SQGLE GAC AMPCQ, @MPCFMJCQ (? L?PPMU QF?DR @MPCB GLRM RFC EPMSLB), RPCC PGLEQ, EJ?AGCP JCLERFQ, NMJJCL PCK?GLQ, ?LB MAC?L QCBGKCLRQ. 5FC B?R? QFMUQ ? AMPPCJ?RGML @CRUCCL RFC RGKGLE MD RCKNCP?RSPC AF?LECQ ?LB BPGTCPQ MD AJGK?RC AF?LEC: @CDMPC RFC *LBSQRPG?J &P? (NPC-1780), RFCPC UCPC RFPCC BPGTCPQ MD AJGK?RC AF?LEC RF?R UCPC LMR PCJ?RCB RM FSK?L ?ARGTGRW MP ?RKMQNFCPGA E?QCQ. 5FC DGPQR MD RFCQC GQ RFC .GJ?LIMTGRAF AWAJCQ. 5FC Milankovitch cycles BCQAPG@C RFC CDDCARQ MD QJGEFR AF?LECQ GL RFC &?PRFsQ MP@GR ML &?PRFsQ AJGK?RC. 5FC JCLERF MD RFC .GJ?LIMTGRAF AWAJCQ P?LECQ @CRUCCL 19,000 ?LB 100,000 WC?PQ. *L MRFCP UMPBQ, MLC AMSJB CVNCAR RM QCC QMKC NPCBGAR?@JC AF?LECQ GL RFC &?PRFsQ AJGK?RC ?QQMAG?RCB UGRF AF?LECQ GL RFC &?PRFsQ MP@GR ?R ? KGLGKSK MD CTCPW 19,000 WC?PQ. 5FC T?PG?RGML GL RFC QSLsQ GLRCLQGRW GQ RFC QCAMLB L?RSP?J D?ARMP PCQNMLQG@JC DMP AJGK?RC AF?LEC. Solar intensity GQ RFC ?KMSLR MD QMJ?P NMUCP MP CLCPEW RFC QSL CKGRQ GL ? EGTCL ?KMSLR MD RGKC. 5FCPC GQ ? BGPCAR PCJ?RGMLQFGN @CRUCCL QMJ?P GLRCLQGRW ?LB RCKNCP?RSPC. "Q QMJ?P GLRCLQGRW GLAPC?QCQ (MP BCAPC?QCQ), RFC &?PRFsQ RCKNCP?RSPC AMPPCQNMLBGLEJW GLAPC?QCQ (MP BCAPC?QCQ). $F?LECQ GL QMJ?P GLRCLQGRW F?TC @CCL NPMNMQCB ?Q MLC MD QCTCP?J NMQQG@JC CVNJ?L?RGMLQ DMP RFC -GRRJC *AC "EC. 'GL?JJW, TMJA?LGA CPSNRGMLQ ?PC ? RFGPB L?RSP?J BPGTCP MD AJGK?RC AF?LEC. 7MJA?LGA CPSNRGMLQ A?L J?QR ? DCU B?WQ, @SR RFC QMJGBQ ?LB E?QCQ PCJC?QCB BSPGLE ?L CPSNRGML A?L GLDJSCLAC RFC AJGK?RC MTCP ? NCPGMB MD ? DCU WC?PQ, A?SQGLE QFMPR-RCPK AJGK?RC AF?LECQ. 5FC E?QCQ ?LB QMJGBQ PCJC?QCB @W TMJA?LGA CPSNRGMLQ A?L GLAJSBC A?P@ML BGMVGBC, U?RCP T?NMP, QSJDSP BGMVGBC, FWBPMECL QSJDGBC, FWBPMECL, ?LB A?P@ML KMLMVGBC. (CLCP?JJW, TMJA?LGA CPSNRGMLQ AMMJ RFC AJGK?RC. 5FGQ MAASPPCB GL 1783 UFCL TMJA?LMQ GL *ACJ?LB CPSNRCB ?LB A?SQCB RFC PCJC?QC MD J?PEC TMJSKCQ MD QSJDSPGA MVGBC. 5FGQ JCB RM haze-effect cooling, ? EJM@?J NFCLMKCLML RF?R MAASPQ UFCL BSQR, ?QF, MP MRFCP QSQNCLBCB N?PRGAJCQ @JMAI MSR QSLJGEFR ?LB RPGEECP JMUCP EJM@?J RCKNCP?RSPCQ ?Q ? PCQSJR; F?XC-CDDCAR AMMJGLE SQS?JJW CVRCLBQ DMP MLC MP KMPC WC?PQ. *L &SPMNC ?LB /MPRF "KCPGA?, F?XC-CDDCAR AMMJGLE NPMBSACB QMKC MD RFC JMUCQR ?TCP?EC UGLRCP RCKNCP?RSPCQ ML PCAMPB GL 1783 ?LB 1784. (PCCLFMSQC E?QCQ ?PC NPM@?@JW RFC KMQR QGELGDGA?LR BPGTCPQ MD RFC AJGK?RC. 8FCL FC?R CLCPEW DPMK RFC QSL QRPGICQ RFC &?PRF, E?QCQ ILMUL ?Q greenhouse gases RP?N RFC FC?R GL RFC ?RKMQNFCPC, ?Q BM RFC EJ?QQ N?LCQ MD ? EPCCLFMSQC ICCN FC?R DPMK CQA?NGLE. 5FC EPCCLFMSQC E?QCQ RF?R ?DDCAR &?PRF GLAJSBC A?P@ML BGMVGBC, KCRF?LC, U?RCP T?NMP, LGRPMSQ MVGBC, ?LB MXMLC. "NNPMVGK?RCJW F?JD MD RFC P?BG?RGML DPMK RFC QSL N?QQCQ RFPMSEF RFCQC E?QCQ GL RFC ?RKMQNFCPC ?LB QRPGICQ RFC &?PRF. 5FGQ P?BG?RGML GQ AMLTCPRCB GLRM RFCPK?J P?BG?RGML ML RFC &?PRFsQ QSPD?AC, ?LB RFCL ? NMPRGML MD RF?R CLCPEW GQ PC-P?BG?RCB @?AI GLRM RFC ?RKMQNFCPC. (PCCLFMSQC E?QCQ, FMUCTCP, PCDJCAR KSAF MD RFC RFCPK?J CLCPEW @?AI RM RFC &?PRFsQ QSPD?AC. 5FC KMPC EPCCLFMSQC E?QCQ RFCPC ?PC GL RFC ?RKMQNFCPC, RFC KMPC RFCPK?J CLCPEW GQ PCDJCARCB @?AI RM RFC &?PRFsQ QSPD?AC. (PCCLFMSQC E?QCQ ?@QMP@ ?LB CKGR P?BG?RGML ?LB ?PC ?L GKNMPR?LR D?ARMP GL RFC greenhouse effect: RFC U?PKGLE MD &?PRF BSC RM A?P@ML BGMVGBC ?LB MRFCP EPCCLFMSQC E?QCQ GL RFC ?RKMQNFCPC. &TGBCLAC QSNNMPRQ RFC PCJ?RGMLQFGN @CRUCCL ?RKMQNFCPGA AMLACLRP?RGMLQ MD A?P@ML BGMVGBC ?LB RCKNCP?RSPC: ?Q A?P@ML BGMVGBC PGQCQ, EJM@?J RCKNCP?RSPC PGQCQ. 4GLAC 1950, RFC AMLACLRP?RGML MD ?RKMQNFCPGA A?P@ML BGMVGBC F?Q GLAPC?QCB DPMK ?@MSR 280 NNK RM 382 NNK GL 2006. *L 2011, RFC ?RKMQNFCPGA A?P@ML BGMVGBC AMLACLRP?RGML U?Q 392 NNK. )MUCTCP, RFC NJ?LCR UMSJB LMR @C GLF?@GR?@JC @W ASPPCLR JGDC DMPKQ GD U?RCP T?NMP BGB LMR NPMBSAC GRQ BP?QRGA EPCCLFMSQC U?PKGLE CDDCAR.
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qGL ? K?RRCP MD FSLBPCBQ MD WC?PQ P?RFCP RF?L RFMSQ?LBQ MD WC?PQ. 8F?R GQ RFC PC?QML DMP RFGQ BGDDCPCLAC GL RFC P?RC MD AF?LEC ?LB RFC ?KMSLR MD GLAPC?QC GL A?P@ML BGMVGBC " ICW D?ARMP RF?R KSQR @C PCAMELGXCB UFCL AMKN?PGLE RFC FGQRMPGA?J B?R? ?LB RFC ASPPCLR B?R? GQ RFC NPCQCLAC MD KMBCPL FSK?L QMAGCRW; LM MRFCP BPGTCP MD AJGK?RC AF?LEC F?Q WGCJBCB AF?LECQ GL ?RKMQNFCPGA A?P@ML BGMVGBC JCTCJQ ?R RFGQ P?RC MP RM RFGQ K?ELGRSBC. )SK?L ?ARGTGRW PCJC?QCQ A?P@ML BGMVGBC ?LB KCRF?LC, RUM MD RFC KMQR GKNMPR?LR EPCCLFMSQC E?QCQ, GLRM RFC ?RKMQNFCPC GL QCTCP?J U?WQ. 5FC NPGK?PW KCAF?LGQK RF?R PCJC?QCQ A?P@ML BGMVGBC GQ RFC @SPLGLE MD DMQQGJ DSCJQ, QSAF ?Q E?QMJGLC, AM?J, ?LB L?RSP?J E?Q (Figure 44.28). %CDMPCQR?RGML, ACKCLR K?LSD?ARSPC, ?LGK?J ?EPGASJRSPC, RFC AJC?PGLE MD J?LB, ?LB RFC @SPLGLE MD DMPCQRQ ?PC MRFCP FSK?L ?ARGTGRGCQ RF?R PCJC?QC A?P@ML BGMVGBC. .CRF?LC ($)4) GQ NPMBSACB UFCL @?ARCPG? @PC?I BMUL MPE?LGA K?RRCP SLBCP ?L?CPM@GA AMLBGRGMLQ. "L?CPM@GA AMLBGRGMLQ A?L F?NNCL UFCL MPE?LGA K?RRCP GQ RP?NNCB SLBCPU?RCP (QSAF ?Q GL PGAC N?BBGCQ) MP GL RFC GLRCQRGLCQ MD FCP@GTMPCQ. .CRF?LC A?L ?JQM @C PCJC?QCB DPMK L?RSP?J E?Q DGCJBQ ?LB RFC BCAMKNMQGRGML RF?R MAASPQ GL J?LBDGJJQ. "LMRFCP QMSPAC MD KCRF?LC GQ RFC KCJRGLE MD AJ?RFP?RCQ. Clathrates ?PC DPMXCL AFSLIQ MD GAC ?LB KCRF?LC DMSLB ?R RFC @MRRMK MD RFC MAC?L. 8FCL U?RCP U?PKQ, RFCQC AFSLIQ MD GAC KCJR ?LB KCRF?LC GQ PCJC?QCB. "Q RFC MAC?LsQ U?RCP RCKNCP?RSPC GLAPC?QCQ, RFC P?RC ?R UFGAF AJ?RFP?RCQ KCJR GQ GLAPC?QGLE, PCJC?QGLE CTCL KMPC KCRF?LC. 5FGQ JC?BQ RM GLAPC?QCB JCTCJQ MD KCRF?LC GL RFC ?RKMQNFCPC, UFGAF DSPRFCP ?AACJCP?RCQ RFC P?RC MD EJM@?J U?PKGLE. 5FGQ GQ ?L CV?KNJC MD RFC NMQGRGTC DCCB@?AI JMMN RF?R GQ JC?BGLE RM RFC P?NGB P?RC MD GLAPC?QC MD EJM@?J RCKNCP?RSPCQ.
!@>LI< 44.28 4EB ?ROKFKD LC CLPPFI CRBIP FK FKARPQOV >KA ?V SBEF@IBP OBIB>PBP @>O?LK AFLUFAB >KA LQEBO DOBBKELRPB D>PBP FKQL QEB >QJLPMEBOB. (@OBAFQ: x0hIIhy/7FHFJBAF> #LJJLKP)
DF:LD<EK<; -<JLCKJ F= CC@D8K< C?8E><: +8JK 8E; +I<J<EK

4AGCLRGQRQ F?TC ECMJMEGA?J CTGBCLAC MD RFC AMLQCOSCLACQ MD JMLE-?EM AJGK?RC AF?LEC. .MBCPL-B?W NFCLMKCL? QSAF ?Q PCRPC?RGLE EJ?AGCPQ ?LB KCJRGLE NMJ?P GAC A?SQC ? AMLRGLS?J PGQC GL QC? JCTCJ. .C?LUFGJC, AF?LECQ GL AJGK?RC A?L LCE?RGTCJW ?DDCAR MPE?LGQKQ. G+414-/)'1 C1/2'9+ C.'3-+ (JM@?J U?PKGLE F?Q @CCL ?QQMAG?RCB UGRF ?R JC?QR MLC NJ?LCR-UGBC CVRGLARGML CTCLR BSPGLE RFC ECMJMEGA?J N?QR. 5FC 1CPKG?L CVRGLARGML CTCLR MAASPPCB ?@MSR 251 KGJJGML WC?PQ ?EM RMU?PB RFC CLB MD RFC PMSEFJW 50-KGJJGML-WC?P-JMLE ECMJMEGA?J RGKC QN?L ILMUL ?Q RFC 1CPKG?L NCPGMB. 5FGQ ECMJMEGA RGKC NCPGMB U?Q MLC MD RFC RFPCC U?PKCQR NCPGMBQ GL &?PRFsQ ECMJMEGA FGQRMPW. 4AGCLRGQRQ CQRGK?RC RF?R ?NNPMVGK?RCJW 70 NCPACLR MD RFC RCPPCQRPG?J NJ?LR ?LB ?LGK?J QNCAGCQ ?LB 84 NCPACLR MD K?PGLC QNCAGCQ @CA?KC CVRGLAR, T?LGQFGLE DMPCTCP LC?P RFC CLB MD RFC 1CPKG?L NCPGMB. 0PE?LGQKQ RF?R F?B ?B?NRCB RM UCR ?LB U?PK
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1320
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1321
DCK?JC-@G?QCB QCV P?RGM (RFC P?RGM MD K?JCQ RM DCK?JCQ) MP ?EC QRPSARSPC (RFC NPMNMPRGML MD NMNSJ?RGML KCK@CPQ ?R QNCAGDGA ?EC P?LECQ) AMKNMQCB MD K?LW GLBGTGBS?JQ MD PCNPMBSARGTC ?EC A?L GLAPC?QC @GPRF P?RCQ. *L ?BBGRGML, RFC BCKMEP?NFGA AF?P?ARCPGQRGAQ MD ? NMNSJ?RGML A?L GLDJSCLAC FMU RFC NMNSJ?RGML EPMUQ MP BCAJGLCQ MTCP RGKC. *D @GPRF ?LB BC?RF P?RCQ ?PC COS?J, RFC NMNSJ?RGML PCK?GLQ QR?@JC. )MUCTCP, RFC NMNSJ?RGML QGXC UGJJ GLAPC?QC GD @GPRF P?RCQ CVACCB BC?RF P?RCQ; RFC NMNSJ?RGML UGJJ BCAPC?QC GD @GPRF P?RCQ ?PC JCQQ RF?L BC?RF P?RCQ. -GDC CVNCAR?LAW GQ ?LMRFCP GKNMPR?LR D?ARMP; RFC JCLERF MD RGKC GLBGTGBS?JQ PCK?GL GL RFC NMNSJ?RGML GKN?ARQ JMA?J PCQMSPACQ, PCNPMBSARGML, ?LB RFC MTCP?JJ FC?JRF MD RFC NMNSJ?RGML. 5FCQC BCKMEP?NFGA AF?P?ARCPGQRGAQ ?PC MDRCL BGQNJ?WCB GL RFC DMPK MD ? JGDC R?@JC. L/,+ "'(1+8 -GDC R?@JCQ NPMTGBC GKNMPR?LR GLDMPK?RGML ?@MSR RFC JGDC FGQRMPW MD ?L MPE?LGQK. -GDC R?@JCQ BGTGBC RFC NMNSJ?RGML GLRM ?EC EPMSNQ ?LB MDRCL QCVCQ, ?LB QFMU FMU JMLE ? KCK@CP MD RF?R EPMSN GQ JGICJW RM JGTC. 5FCW ?PC KMBCJCB ?DRCP ?ARS?PG?J R?@JCQ SQCB @W RFC GLQSP?LAC GLBSQRPW DMP CQRGK?RGLE FSK?L JGDC CVNCAR?LAW. -GDC R?@JCQ K?W GLAJSBC RFC NPM@?@GJGRW MD GLBGTGBS?JQ BWGLE @CDMPC RFCGP LCVR @GPRFB?W (G.C., RFCGP mortality rate), RFC NCPACLR?EC MD QSPTGTGLE GLBGTGBS?JQ BWGLE ?R ? N?PRGASJ?P ?EC GLRCPT?J, ?LB RFCGP JGDC CVNCAR?LAW ?R C?AF GLRCPT?J. "L CV?KNJC MD ? JGDC R?@JC GQ QFMUL GL Table 45.1 DPMK ? QRSBW MD %?JJ KMSLR?GL QFCCN, ? QNCAGCQ L?RGTC RM LMPRFUCQRCPL /MPRF "KCPGA?. /MRGAC RF?R RFC NMNSJ?RGML GQ BGTGBCB GLRM ?EC GLRCPT?JQ (AMJSKL "). 5FC KMPR?JGRW P?RC (NCP 1000), QFMUL GL AMJSKL %, GQ @?QCB ML RFC LSK@CP MD GLBGTGBS?JQ BWGLE BSPGLE RFC ?EC GLRCPT?J (AMJSKL #) BGTGBCB @W RFC LSK@CP MD GLBGTGBS?JQ QSPTGTGLE ?R RFC @CEGLLGLE MD RFC GLRCPT?J ($MJSKL $), KSJRGNJGCB @W 1000.
mortality rate =
number of individuals dying x 1000 number of individuals surviving
'MP CV?KNJC, @CRUCCL ?ECQ RFPCC ?LB DMSP, 12 GLBGTGBS?JQ BGC MSR MD RFC 776 RF?R UCPC PCK?GLGLE DPMK RFC MPGEGL?J 1000 QFCCN. 5FGQ LSK@CP GQ RFCL KSJRGNJGCB @W 1000 RM ECR RFC KMPR?JGRW P?RC NCP RFMSQ?LB.
mortality rate = 12 x 1000 15.5 776
"Q A?L @C QCCL DPMK RFC KMPR?JGRW P?RC B?R? (AMJSKL %), ? FGEF BC?RF P?RC MAASPPCB UFCL RFC QFCCN UCPC @CRUCCL 6 ?LB 12 KMLRFQ MJB, ?LB RFCL GLAPC?QCB CTCL KMPC DPMK 8 RM 12 WC?PQ MJB, ?DRCP UFGAF RFCPC UCPC DCU QSPTGTMPQ. 5FC B?R? GLBGA?RC RF?R GD ? QFCCN GL RFGQ NMNSJ?RGML UCPC RM QSPTGTC RM ?EC MLC, GR AMSJB @C CVNCARCB RM JGTC ?LMRFCP 7.7 WC?PQ ML ?TCP?EC, ?Q QFMUL @W RFC JGDC CVNCAR?LAW LSK@CPQ GL AMJSKL &.
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54 145 12 13 12 30 46
617
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1000 946 801 789 776 764 734
(FIK8C@KP I8K< G@EE@E> F= 8>< @EK<IM8C

54.0 153.3 15.0 16.5 15.5 39.3 62.7
'@=< <OG<:K8E:P FI D<8E C@=<K@D< I<D8@E@E> KF K?FJ< 8KK8@E@E> 8>< @EK<IM8C

7.06 -7.7 6.8 5.9 5.0 4.2
0-0.5 0.5-1 1-2 2-3 3-4 4-5 5-6
/89C< 45.1 /?@J C@=< K89C< F= O;/8 *'11/ J?FNJ K?< ELD9< @EK<IM8C =FI K?< D8CC DFLEK8@E J?<<G.
1. %?R? "B?NRCB DPMK &BU?PB 4. %CCTCW, +P., t-GDC 5?@JCQ DMP /?RSP?J 1MNSJ?RGMLQ MD "LGK?JQ,u !/, Q<(9;,93@ R,=0,> 6- B0636.@ 22, LM. 4 (%CACK@CP 1947): 283-314.
1322
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48 69 132 187 156 90 3 3
627
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)LD9 8K 9<>@EE@E> F= 8>< @EK<IM8C FLK F= 1000 9FIE

688 640 571 439 252 96 6 3
(FIK8C@KP I8K< G@EE@E> F= 8>< @EK<IM8C

69.8 107.8 231.2 426.0 619.0 937.5 500.0 1000
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3.4 2.6 1.9 1.3 0.9 0.6 1.2 0.7
6-7 7-8 8-9 9-10 10-11 11-12 12-13 13-14
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2. %?R? "B?NRCB DPMK &BU?PB 4. %CCTCW, +P., t-GDC 5?@JCQ DMP /?RSP?J 1MNSJ?RGMLQ MD "LGK?JQ,u !/, Q<(9;,93@ R,=0,> 6- B0636.@ 22, LM. 4 (%CACK@CP 1947): 283-314.
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1323
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1324
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$F?PJCQ %?PUGL, GL FGQ RFCMPW MD L?RSP?J QCJCARGML, U?Q EPC?RJW GLDJSCLACB @W RFC &LEJGQF AJCPEWK?L 5FMK?Q .?JRFSQ. .?JRFSQ NS@JGQFCB ? @MMI GL 1798 QR?RGLE RF?R NMNSJ?RGMLQ UGRF SLJGKGRCB L?RSP?J PCQMSPACQ EPMU TCPW P?NGBJW, ?LB RFCL NMNSJ?RGML EPMURF BCAPC?QCQ ?Q PCQMSPACQ @CAMKC BCNJCRCB. 5FGQ ?AACJCP?RGLE N?RRCPL MD GLAPC?QGLE NMNSJ?RGML QGXC GQ A?JJCB exponential growth. 5FC @CQR CV?KNJC MD CVNMLCLRG?J EPMURF GQ QCCL GL @?ARCPG?. #?ARCPG? ?PC NPMI?PWMRCQ RF?R PCNPMBSAC @W NPMI?PWMRGA DGQQGML. 5FGQ BGTGQGML R?ICQ ?@MSR ?L FMSP DMP K?LW @?ARCPG?J QNCAGCQ. *D 1000 @?ARCPG? ?PC NJ?ACB GL ? J?PEC DJ?QI UGRF ?L SLJGKGRCB QSNNJW MD LSRPGCLRQ (QM RFC LSRPGCLRQ UGJJ LMR @CAMKC BCNJCRCB), ?DRCP ?L FMSP, RFCPC GQ MLC PMSLB MD BGTGQGML ?LB C?AF MPE?LGQK BGTGBCQ, PCQSJRGLE GL 2000 MPE?LGQKQq?L GLAPC?QC MD 1000. *L ?LMRFCP FMSP, C?AF MD RFC 2000 MPE?LGQKQ UGJJ BMS@JC, NPMBSAGLE 4000, ?L GLAPC?QC MD 2000 MPE?LGQKQ. "DRCP RFC RFGPB FMSP, RFCPC QFMSJB @C 8000 @?ARCPG? GL RFC DJ?QI, ?L GLAPC?QC MD 4000 MPE?LGQKQ. 5FC GKNMPR?LR AMLACNR MD CVNMLCLRG?J EPMURF GQ RF?R RFC population growth rateqRFC LSK@CP MD MPE?LGQKQ ?BBCB GL C?AF PCNPMBSARGTC ECLCP?RGMLqGQ ?AACJCP?RGLE; RF?R GQ, GR GQ GLAPC?QGLE ?R ? EPC?RCP ?LB EPC?RCP P?RC. "DRCP 1 B?W ?LB 24 MD RFCQC AWAJCQ, RFC NMNSJ?RGML UMSJB F?TC GLAPC?QCB DPMK 1000 RM KMPC RF?L 16 @GJJGML. 8FCL RFC NMNSJ?RGML QGXC, N, GQ NJMRRCB MTCP RGKC, ? J-shaped growth curve GQ NPMBSACB (Figure 45.9). 5FC @?ARCPG? CV?KNJC GQ LMR PCNPCQCLR?RGTC MD RFC PC?J UMPJB UFCPC PCQMSPACQ ?PC JGKGRCB. 'SPRFCPKMPC, QMKC @?ARCPG? UGJJ BGC BSPGLE RFC CVNCPGKCLR ?LB RFSQ LMR PCNPMBSAC, JMUCPGLE RFC EPMURF P?RC. 5FCPCDMPC, UFCL A?JASJ?RGLE RFC EPMURF P?RC MD ? NMNSJ?RGML, RFC death rate (D) (LSK@CP MPE?LGQKQ RF?R BGC BSPGLE ? N?PRGASJ?P RGKC GLRCPT?J) GQ QS@RP?ARCB DPMK RFC birth rate (B) (LSK@CP MPE?LGQKQ RF?R ?PC @MPL BSPGLE RF?R GLRCPT?J). 5FGQ GQ QFMUL GL RFC DMJJMUGLE DMPKSJ?:
N (change in number) = B (birth rate) - D (death rate) T (change in time)
5FC @GPRF P?RC GQ SQS?JJW CVNPCQQCB ML ? NCP A?NGR? (DMP C?AF GLBGTGBS?J) @?QGQ. 5FSQ, B (@GPRF P?RC) = )N (RFC NCP A?NGR? @GPRF P?RC t)u KSJRGNJGCB @W RFC LSK@CP MD GLBGTGBS?JQ tNu) ?LB D (BC?RF P?RC) =+N (RFC NCP A?NGR? BC?RF P?RC tBu KSJRGNJGCB @W RFC LSK@CP MD GLBGTGBS?JQ tNu). "BBGRGML?JJW, CAMJMEGQRQ ?PC GLRCPCQRCB GL RFC NMNSJ?RGML ?R ? N?PRGASJ?P NMGLR GL RGKC, ?L GLDGLGRCJW QK?JJ RGKC GLRCPT?J. 'MP RFGQ PC?QML, RFC RCPKGLMJMEW MD BGDDCPCLRG?J A?JASJSQ GQ SQCB RM M@R?GL RFC tGLQR?LR?LCMSQu EPMURF P?RC, PCNJ?AGLE RFC */(5., GL LSK@CP ?LB RGKC UGRF ?L GLQR?LR-QNCAGDGA KC?QSPCKCLR MD LSK@CP ?LB RGKC.
dN = bN dN = (b - d)N dT
/MRGAC RF?R RFC t+u ?QQMAG?RCB UGRF RFC DGPQR RCPK PCDCPQ RM RFC BCPGT?RGTC (?Q RFC RCPK GQ SQCB GL A?JASJSQ) ?LB GQ BGDDCPCLR DPMK RFC BC?RF P?RC, ?JQM A?JJCB t+.u 5FC BGDDCPCLAC @CRUCCL @GPRF ?LB BC?RF P?RCQ GQ DSPRFCP QGKNJGDGCB @W QS@QRGRSRGLE RFC RCPK t9u (GLRPGLQGA P?RC MD GLAPC?QC) DMP RFC PCJ?RGMLQFGN @CRUCCL @GPRF ?LB BC?RF P?RCQ:
dN = rN dT
5FC T?JSC t9K A?L @C NMQGRGTC, KC?LGLE RFC NMNSJ?RGML GQ GLAPC?QGLE GL QGXC; MP LCE?RGTC, KC?LGLE RFC NMNSJ?RGML GQ BCAPC?QGLE GL QGXC; MP XCPM, UFCPC RFC NMNSJ?RGMLsQ QGXC GQ SLAF?LEGLE, ? AMLBGRGML ILMUL
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?Q zero population growth. " DSPRFCP PCDGLCKCLR MD RFC DMPKSJ? PCAMELGXCQ RF?R BGDDCPCLR QNCAGCQ F?TC GLFCPCLR BGDDCPCLACQ GL RFCGP GLRPGLQGA P?RC MD GLAPC?QC (MDRCL RFMSEFR MD ?Q RFC NMRCLRG?J DMP PCNPMBSARGML), CTCL SLBCP GBC?J AMLBGRGMLQ. 0@TGMSQJW, ? @?ARCPGSK A?L PCNPMBSAC KMPC P?NGBJW ?LB F?TC ? FGEFCP GLRPGLQGA P?RC MD EPMURF RF?L ? FSK?L. 5FC K?VGK?J EPMURF P?RC DMP ? QNCAGCQ GQ GRQ biotic potential, or rmax, RFSQ AF?LEGLE RFC COS?RGML RM:
dN = r max N dT
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s F?@GR?R ?LB @CF?TGMP, CQNCAG?JJW GL RFC U?W RF?R RFCW M@R?GL PCQMSPACQ ?LB A?PC DMP RFCGP WMSLE. *R GLAJSBCQ JCLERF MD JGDC ?LB QSPTGTMPQFGN D?ARMPQ ?Q UCJJ. 'MP RFGQ ?L?JWQGQ, NMNSJ?RGML @GMJMEGQRQ F?TC EPMSNCB QNCAGCQ GLRM RFC RUM J?PEC A?RCEMPGCQqK-QCJCARCB ?LB 9-QCJCARCBq?JRFMSEF RFCW ?PC PC?JJW RUM CLBQ MD ? AMLRGLSSK.
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1333
K-selected species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igure 45.13a). 0?I RPCCQ EPMU TCPW QJMUJW ?LB R?IC, ML ?TCP?EC, 20 WC?PQ RM NPMBSAC RFCGP DGPQR QCCBQ, ILMUL ?Q ?AMPLQ. "Q K?LW ?Q 50,000 ?AMPLQ A?L @C NPMBSACB @W ?L GLBGTGBS?J RPCC, @SR RFC ECPKGL?RGML P?RC GQ JMU ?Q K?LW MD RFCQC PMR MP ?PC C?RCL @W ?LGK?JQ QSAF ?Q QOSGPPCJQ. *L QMKC WC?PQ, M?IQ K?W NPMBSAC ?L CVACNRGML?JJW J?PEC LSK@CP MD ?AMPLQ, ?LB RFCQC WC?PQ K?W @C ML ? RUM- MP RFPCC-WC?P AWAJC BCNCLBGLE ML RFC QNCAGCQ MD M?I (9-QCJCARGML). "Q M?I RPCCQ EPMU RM ? J?PEC QGXC ?LB DMP K?LW WC?PQ @CDMPC RFCW @CEGL RM NPMBSAC ?AMPLQ, RFCW BCTMRC ? J?PEC NCPACLR?EC MD RFCGP CLCPEW @SBECR RM EPMURF ?LB K?GLRCL?LAC. 5FC RPCCsQ FCGEFR ?LB QGXC ?JJMU GR RM BMKGL?RC MRFCP NJ?LRQ GL RFC AMKNCRGRGML DMP QSLJGEFR, RFC M?IsQ NPGK?PW CLCPEW PCQMSPAC. 'SPRFCPKMPC, UFCL GR BMCQ PCNPMBSAC, RFC M?I NPMBSACQ J?PEC, CLCPEW-PGAF QCCBQ RF?R SQC RFCGP CLCPEW PCQCPTC RM @CAMKC OSGAIJW CQR?@JGQFCB (K-QCJCARGML). *L AMLRP?QR, r-selected species F?TC ? J?PEC LSK@CP MD QK?JJ MDDQNPGLE (FCLAC RFCGP 9 BCQGEL?RGML (Table 45.2). 5FGQ QRP?RCEW GQ MDRCL CKNJMWCB GL SLNPCBGAR?@JC MP AF?LEGLE CLTGPMLKCLRQ. "LGK?JQ RF?R ?PC 9-QCJCARCB BM LMR EGTC JMLE-RCPK N?PCLR?J A?PC ?LB RFC MDDQNPGLE ?PC PCJ?RGTCJW K?RSPC ?LB QCJD-QSDDGAGCLR ?R @GPRF. &V?KNJCQ MD 9-QCJCARCB QNCAGCQ ?PC K?PGLC GLTCPRC@P?RCQ, QSAF ?Q HCJJWDGQF, ?LB NJ?LRQ, QSAF ?Q RFC B?LBCJGML (Figure 45.13b). %?LBCJGMLQ F?TC QK?JJ QCCBQ RF?R ?PC UGLB BGQNCPQCB JMLE BGQR?LACQ. .?LW QCCBQ ?PC NPMBSACB QGKSJR?LCMSQJW RM CLQSPC RF?R ?R JC?QR QMKC MD RFCK PC?AF ? FMQNGR?@JC CLTGPMLKCLR. 4CCBQ RF?R J?LB GL GLFMQNGR?@JC CLTGPMLKCLRQ F?TC JGRRJC AF?LAC DMP QSPTGT?J QGLAC RFCGP QCCBQ ?PC JMU GL CLCPEW AMLRCLR. /MRC RF?R QSPTGT?J GQ LMR LCACQQ?PGJW ? DSLARGML MD CLCPEW QRMPCB GL RFC QCCB GRQCJD.
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1335
"JRFMSEF FSK?LQ F?TC GLAPC?QCB RFC A?PPWGLE A?N?AGRW MD RFCGP CLTGPMLKCLR, RFC RCAFLMJMEGCQ SQCB RM ?AFGCTC RFGQ RP?LQDMPK?RGML F?TC A?SQCB SLNPCACBCLRCB AF?LECQ RM &?PRFsQ CLTGPMLKCLR, ?JRCPGLE CAMQWQRCKQ RM RFC NMGLR UFCPC QMKC K?W @C GL B?LECP MD AMJJ?NQC. 5FC BCNJCRGML MD RFC MXMLC J?WCP, CPMQGML BSC RM ?AGB P?GL, ?LB B?K?EC DPMK EJM@?J AJGK?RC AF?LEC ?PC A?SQCB @W FSK?L ?ARGTGRGCQ. 5FC SJRGK?RC CDDCAR MD RFCQC AF?LECQ ML MSP A?PPWGLE A?N?AGRW GQ SLILMUL. "Q QMKC NMGLR MSR, GR GQ JGICJW RF?R RFC LCE?RGTC CDDCARQ MD GLAPC?QGLE A?PPWGLE A?N?AGRW UGJJ MSRUCGEF RFC NMQGRGTC MLCQqRFC A?PPWGLE A?N?AGRW MD RFC UMPJB DMP FSK?L @CGLEQ KGEFR ?ARS?JJW BCAPC?QC. 5FC UMPJBsQ FSK?L NMNSJ?RGML GQ ASPPCLRJW CVNCPGCLAGLE CVNMLCLRG?J EPMURF CTCL RFMSEF FSK?L PCNPMBSARGML GQ D?P @CJMU GRQ @GMRGA NMRCLRG?J (Figure 45.14). 5M PC?AF GRQ @GMRGA NMRCLRG?J, ?JJ DCK?JCQ UMSJB F?TC RM @CAMKC NPCEL?LR CTCPW LGLC KMLRFQ MP QM BSPGLE RFCGP PCNPMBSARGTC WC?PQ. "JQM, PCQMSPACQ UMSJB F?TC RM @C QSAF RF?R RFC CLTGPMLKCLR UMSJB QSNNMPR QSAF EPMURF. /CGRFCP MD RFCQC RUM AMLBGRGMLQ CVGQRQ. *L QNGRC MD RFGQ D?AR, FSK?L NMNSJ?RGML GQ QRGJJ EPMUGLE CVNMLCLRG?JJW.
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" AMLQCOSCLAC MD CVNMLCLRG?J FSK?L NMNSJ?RGML EPMURF GQ RFC RGKC RF?R GR R?ICQ RM ?BB ? N?PRGASJ?P LSK@CP MD FSK?LQ RM RFC &?PRF GQ @CAMKGLE QFMPRCP. Figure 45.15 QFMUQ RF?R 123 WC?PQ UCPC LCACQQ?PW RM ?BB 1 @GJJGML FSK?LQ GL 1930, @SR GR MLJW RMMI 24 WC?PQ RM ?BB RUM @GJJGML NCMNJC @CRUCCL 1975 ?LB 1999. "Q ?JPC?BW BGQASQQCB, ?R QMKC NMGLR GR UMSJB ?NNC?P RF?R MSP ?@GJGRW RM GLAPC?QC MSP A?PPWGLE A?N?AGRW GLBCDGLGRCJW ML ? DGLGRC UMPJB GQ SLACPR?GL. 8GRFMSR LCU RCAFLMJMEGA?J ?BT?LACQ, RFC FSK?L EPMURF P?RC F?Q @CCL NPCBGARCB RM QJMU GL RFC AMKGLE BCA?BCQ. )MUCTCP, RFC NMNSJ?RGML UGJJ QRGJJ @C GLAPC?QGLE ?LB RFC RFPC?R MD MTCPNMNSJ?RGML PCK?GLQ.
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7GQGR RFGQ website (http://openstaxcollege.org/l/populations) ?LB QCJCAR t-?SLAF KMTGCu DMP ?L ?LGK?RGML BGQASQQGLE RFC EJM@?J GKN?ARQ MD FSK?L NMNSJ?RGML EPMURF.
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1CPF?NQ RFC AJ?QQGA?J CV?KNJC MD QNCAGCQ GLRCP?ARGML GQ NPCB?RGML: RFC FSLRGLE MD NPCW @W GRQ NPCB?RMP. /?RSPC QFMUQ ML RCJCTGQGML FGEFJGEFR RFC BP?K? MD MLC JGTGLE MPE?LGQK IGJJGLE ?LMRFCP. 1MNSJ?RGMLQ MD NPCB?RMPQ ?LB NPCW GL ? AMKKSLGRW ?PC LMR AMLQR?LR MTCP RGKC: GL KMQR A?QCQ, RFCW T?PW GL AWAJCQ RF?R ?NNC?P RM @C PCJ?RCB. 5FC KMQR MDRCL AGRCB CV?KNJC MD NPCB?RMP-NPCW BWL?KGAQ GQ QCCL GL RFC AWAJGLE MD RFC JWLV (NPCB?RMP) ?LB RFC QLMUQFMC F?PC (NPCW), SQGLE LC?PJW 200 WC?P-MJB RP?NNGLE B?R? DPMK /MPRF "KCPGA?L DMPCQRQ (Figure 45.18). 5FGQ AWAJC MD NPCB?RMP ?LB NPCW J?QRQ ?NNPMVGK?RCJW 10 WC?PQ, UGRF RFC NPCB?RMP NMNSJ?RGML J?EEGLE 1p2 WC?PQ @CFGLB RF?R MD RFC NPCW NMNSJ?RGML. "Q RFC F?PC LSK@CPQ GLAPC?QC, RFCPC GQ KMPC DMMB ?T?GJ?@JC DMP RFC JWLV, ?JJMUGLE RFC JWLV NMNSJ?RGML RM GLAPC?QC ?Q UCJJ. 8FCL RFC JWLV NMNSJ?RGML EPMUQ RM ? RFPCQFMJB JCTCJ, FMUCTCP, RFCW IGJJ QM K?LW F?PCQ RF?R F?PC NMNSJ?RGML @CEGLQ RM BCAJGLC, DMJJMUCB @W ? BCAJGLC GL RFC JWLV NMNSJ?RGML @CA?SQC MD QA?PAGRW MD DMMB. 8FCL RFC JWLV NMNSJ?RGML GQ JMU, RFC F?PC NMNSJ?RGML QGXC @CEGLQ RM GLAPC?QC BSC, ?R JC?QR GL N?PR, RM JMU NPCB?RGML NPCQQSPC, QR?PRGLE RFC AWAJC ?LCU.
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R?QRC MP NPCQCLAC MD RMVGA AFCKGA?JQ ?LB JC?PL LMR RM C?R RFCK GL RFC DSRSPC. 5FGQ RWNC MD BCDCLQGTC KCAF?LGQK GQ A?JJCB aposematic coloration, MP U?PLGLE AMJMP?RGML (Figure 45.21).
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!@>LI< 45.32 $ROFKD MOFJ>OV PR@@BPPFLK FK I>S> LK ->RF, (>T>FF, PR@@RIBKQ MI>KQP >OB QEB MFLKBBO PMB@FBP. (@OBAFQ: &LOBPQ >KA +FJ 3Q>OO)
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1352
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Conditioned behaviors ?PC RWNCQ MD ?QQMAG?RGTC JC?PLGLE, UFCPC ? QRGKSJSQ @CAMKCQ ?QQMAG?RCB UGRF ? AMLQCOSCLAC. %SPGLE MNCP?LR AMLBGRGMLGLE, RFC @CF?TGMP?J PCQNMLQC GQ KMBGDGCB @W GRQ AMLQCOSCLACQ, UGRF PCE?PBQ RM GRQ DMPK, QRPCLERF, MP DPCOSCLAW. C1'88/)'1 C43*/9/43/3*L classical conditioning, ? PCQNMLQC A?JJCB RFC AMLBGRGMLCB PCQNMLQC GQ ?QQMAG?RCB UGRF ? QRGKSJSQ RF?R GR F?B NPCTGMSQJW LMR @CCL ?QQMAG?RCB UGRF, RFC AMLBGRGMLCB QRGKSJSQ. 5FC PCQNMLQC RM RFC MPGEGL?J, SLAMLBGRGMLCB QRGKSJSQ GQ A?JJCB RFC SLAMLBGRGMLCB PCQNMLQC. 5FC KMQR AGRCB CV?KNJC MD AJ?QQGA?J AMLBGRGMLGLE GQ *T?L 1?TJMTsQ CVNCPGKCLRQ UGRF BMEQ (Figure 45.40). *L 1?TJMTsQ CVNCPGKCLRQ, RFC SLAMLBGRGMLCB PCQNMLQC U?Q RFC Q?JGT?RGML MD BMEQ GL PCQNMLQC RM RFC SLAMLBGRGMLCB QRGKSJSQ MD QCCGLE
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MP QKCJJGLE RFCGP DMMB. 5FC AMLBGRGMLGLE QRGKSJSQ RF?R PCQC?PAFCPQ ?QQMAG?RCB UGRF RFC SLAMLBGRGMLCB PCQNMLQC U?Q RFC PGLEGLE MD ? @CJJ. %SPGLE AMLBGRGMLGLE, CTCPW RGKC RFC ?LGK?J U?Q EGTCL DMMB, RFC @CJJ U?Q PSLE. 5FGQ U?Q PCNC?RCB BSPGLE QCTCP?J RPG?JQ. "DRCP QMKC RGKC, RFC BME JC?PLCB RM ?QQMAG?RC RFC PGLEGLE MD RFC @CJJ UGRF DMMB ?LB RM PCQNMLB @W Q?JGT?RGLE. "DRCP RFC AMLBGRGMLGLE NCPGMB U?Q DGLGQFCB, RFC BME UMSJB PCQNMLB @W Q?JGT?RGLE UFCL RFC @CJJ U?Q PSLE, CTCL UFCL RFC SLAMLBGRGMLCB QRGKSJSQ, RFC DMMB, U?Q ?@QCLR. 5FSQ, RFC PGLEGLE MD RFC @CJJ @CA?KC RFC AMLBGRGMLCB QRGKSJSQ ?LB RFC Q?JGT?RGML @CA?KC RFC AMLBGRGMLCB PCQNMLQC. "JRFMSEF GR GQ RFMSEFR @W QMKC QAGCLRGQRQ RF?R RFC SLAMLBGRGMLCB ?LB AMLBGRGMLCB PCQNMLQCQ ?PC GBCLRGA?J, CTCL 1?TJMT BGQAMTCPCB RF?R RFC Q?JGT? GL RFC AMLBGRGMLCB BMEQ F?B AF?P?ARCPGQRGA BGDDCPCLACQ UFCL AMKN?PCB RM RFC SLAMLBGRGMLCB BME.
!@>LI< 45.40 )K QEB @I>PPF@ 0>SILSF>K OBPMLKPB, QEB ALD ?B@LJBP @LKAFQFLKBA QL >PPL@F>QB QEB OFKDFKD LC QEB ?BII TFQE CLLA.
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operant conditioning, RFC AMLBGRGMLCB @CF?TGMP GQ EP?BS?JJW KMBGDGCB @W GRQ AMLQCOSCLACQ ?Q RFC ?LGK?J PCQNMLBQ RM RFC QRGKSJSQ. " K?HMP NPMNMLCLR MD QSAF AMLBGRGMLGLE U?Q NQWAFMJMEGQR #.'. 4IGLLCP, RFC GLTCLRMP MD RFC 4IGLLCP @MV. 4IGLLCP NSR P?RQ GL FGQ @MVCQ RF?R AMLR?GLCB ? JCTCP RF?R UMSJB BGQNCLQC DMMB RM RFC P?R UFCL BCNPCQQCB. 8FGJC GLGRG?JJW RFC P?R UMSJB NSQF RFC JCTCP ? DCU RGKCQ @W ?AAGBCLR, GR CTCLRS?JJW ?QQMAG?RCB NSQFGLE RFC JCTCP UGRF ECRRGLE RFC DMMB. 5FGQ RWNC MD JC?PLGLE GQ ?L CV?KNJC MD MNCP?LR AMLBGRGMLGLE. 0NCP?LR JC?PLGLE GQ RFC @?QGQ MD KMQR ?LGK?J RP?GLGLE. 5FC AMLBGRGMLCB @CF?TGMP GQ AMLRGLS?JJW KMBGDGCB @W NMQGRGTC MP LCE?RGTC PCGLDMPACKCLR, MDRCL ? PCU?PB QSAF ?Q DMMB MP QMKC RWNC MD NSLGQFKCLR, PCQNCARGTCJW. *L RFGQ U?W, RFC ?LGK?J GQ AMLBGRGMLCB RM ?QQMAG?RC ? RWNC MD @CF?TGMP UGRF RFC NSLGQFKCLR MP PCU?PB, ?LB, MTCP RGKC, A?L @C GLBSACB RM NCPDMPK @CF?TGMPQ RF?R RFCW UMSJB LMR F?TC BMLC GL RFC UGJB, QSAF ?Q RFC tRPGAIQu BMJNFGLQ NCPDMPK ?R K?PGLC ?KSQCKCLR N?PI QFMUQ (Figure 45.41).
!@>LI< 45.41 4EB QO>FKFKD LC ALIMEFKP ?V OBT>OAFKD QEBJ TFQE CLLA FP >K BU>JMIB LC MLPFQFSB OBFKCLO@BJBKQ LMBO>KQ @LKAFQFLKFKD. (@OBAFQ: 2LI>KA 4>KDI>L)
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$J?QQGA?J ?LB MNCP?LR AMLBGRGMLGLE ?PC GLCDDGAGCLR U?WQ DMP FSK?LQ ?LB MRFCP GLRCJJGECLR ?LGK?JQ RM JC?PL. 4MKC NPGK?RCQ, GLAJSBGLE FSK?LQ, ?PC ?@JC RM JC?PL @W GKGR?RGLE RFC @CF?TGMP MD MRFCPQ ?LB @W R?IGLE GLQRPSARGMLQ. 5FC BCTCJMNKCLR MD AMKNJCV J?LES?EC @W FSK?LQ F?Q K?BC cognitive learning, RFC
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K?LGNSJ?RGML MD GLDMPK?RGML SQGLE RFC KGLB, RFC KMQR NPMKGLCLR KCRFMB MD FSK?L JC?PLGLE. *L D?AR, RF?R GQ FMU QRSBCLRQ ?PC JC?PLGLE PGEFR LMU @W PC?BGLE RFGQ @MMI. "Q QRSBCLRQ PC?B, RFCW A?L K?IC KCLR?J GK?ECQ MD M@HCARQ MP MPE?LGQKQ ?LB GK?EGLC AF?LECQ RM RFCK, MP @CF?TGMPQ @W RFCK, ?LB ?LRGAGN?RC RFC AMLQCOSCLACQ. *L ?BBGRGML RM TGQS?J NPMACQQGLE, AMELGRGTC JC?PLGLE GQ ?JQM CLF?LACB @W PCKCK@CPGLE N?QR CVNCPGCLACQ, RMSAFGLE NFWQGA?J M@HCARQ, FC?PGLE QMSLBQ, R?QRGLE DMMB, ?LB ? T?PGCRW MD MRFCP QCLQMPW-@?QCB GLNSRQ. $MELGRGTC JC?PLGLE GQ QM NMUCPDSJ RF?R GR A?L @C SQCB RM SLBCPQR?LB AMLBGRGMLGLE GL BCR?GJ. *L RFC PCTCPQC QACL?PGM, AMLBGRGMLGLE A?LLMR FCJN QMKCMLC JC?PL ?@MSR AMELGRGML. $J?QQGA UMPI ML AMELGRGTC JC?PLGLE U?Q BMLC @W 8MJDE?LE ,dFJCP UGRF AFGKN?LXCCQ. )C BCKMLQRP?RCB RF?R RFCQC ?LGK?JQ UCPC A?N?@JC MD ?@QRP?AR RFMSEFR @W QFMUGLE RF?R RFCW AMSJB JC?PL FMU RM QMJTC ? NSXXJC. 8FCL ? @?L?L? U?Q FSLE GL RFCGP A?EC RMM FGEF DMP RFCK RM PC?AF, ?LB QCTCP?J @MVCQ UCPC NJ?ACB P?LBMKJW ML RFC DJMMP, QMKC MD RFC AFGKNQ UCPC ?@JC RM QR?AI RFC @MVCQ MLC ML RMN MD RFC MRFCP, AJGK@ ML RMN MD RFCK, ?LB ECR RFC @?L?L?. 5FGQ GKNJGCQ RF?R RFCW AMSJB TGQS?JGXC RFC PCQSJR MD QR?AIGLE RFC @MVCQ CTCL @CDMPC RFCW F?B NCPDMPKCB RFC ?ARGML. 5FGQ RWNC MD JC?PLGLE GQ KSAF KMPC NMUCPDSJ ?LB TCPQ?RGJC RF?L AMLBGRGMLGLE. $MELGRGTC JC?PLGLE GQ LMR JGKGRCB RM NPGK?RCQ, ?JRFMSEF RFCW ?PC RFC KMQR CDDGAGCLR GL SQGLE GR. .?XC PSLLGLE CVNCPGKCLRQ BMLC UGRF P?RQ @W ).$. #JMBECRR GL RFC 1920Q UCPC RFC DGPQR RM QFMU AMELGRGTC QIGJJQ GL ? QGKNJC K?KK?J. 5FC KMRGT?RGML DMP RFC ?LGK?JQ RM UMPI RFCGP U?W RFPMSEF RFC K?XC U?Q ? NGCAC MD DMMB ?R GRQ CLB. *L RFCQC QRSBGCQ, RFC ?LGK?JQ GL (PMSN * UCPC PSL GL MLC RPG?J NCP B?W ?LB F?B DMMB ?T?GJ?@JC RM RFCK C?AF B?W ML AMKNJCRGML MD RFC PSL (Figure 45.42). (PMSN ** P?RQ UCPC LMR DCB GL RFC K?XC DMP RFC DGPQR QGV B?WQ ?LB RFCL QS@QCOSCLR PSLQ UCPC BMLC UGRF DMMB DMP QCTCP?J B?WQ ?DRCP. (PMSN *** P?RQ F?B DMMB ?T?GJ?@JC ML RFC RFGPB B?W ?LB CTCPW B?W RFCPC?DRCP. 5FC PCQSJRQ UCPC RF?R RFC AMLRPMJ P?RQ, (PMSN *, JC?PLCB OSGAIJW, ?LB DGESPCB MSR FMU RM PSL RFC K?XC GL QCTCL B?WQ. (PMSN *** BGB LMR JC?PL KSAF BSPGLE RFC RFPCC B?WQ UGRFMSR DMMB, @SR P?NGBJW A?SEFR SN RM RFC AMLRPMJ EPMSN UFCL EGTCL RFC DMMB PCU?PB. (PMSN ** JC?PLCB TCPW QJMUJW DMP RFC QGV B?WQ UGRF LM PCU?PB RM KMRGT?RC RFCK, ?LB RFCW BGB LMR @CEGL RM A?RAF SN RM RFC AMLRPMJ EPMSN SLRGJ RFC B?W DMMB U?Q EGTCL, ?LB RFCL GR RMMI RUM B?WQ JMLECP RM JC?PL RFC K?XC.
!@>LI< 45.42 'OLRM ) (QEB DOBBK PLIFA IFKB) CLRKA CLLA >Q QEB BKA LC B>@E QOF>I, DOLRM )) (QEB ?IRB A>PEBA IFKB) AFA KLQ CFKA CLLA CLO QEB CFOPQ 6 A>VP, >KA DOLRM ))) (QEB OBA ALQQBA IFKB) AFA KLQ CFKA CLLA AROFKD ORKP LK QEB CFOPQ QEOBB A>VP. .LQF@B QE>Q O>QP DFSBK CLLA B>OIFBO IB>OKBA C>PQBO >KA BSBKQR>IIV @>RDEQ RM QL QEB @LKQOLI DOLRM. 4EB LO>KDB ALQP LK QEB DOLRM )) >KA ))) IFKBP PELT QEB A>VP TEBK CLLA OBT>OAP TBOB >AABA QL QEB J>WBP.
*R K?W LMR @C GKKCBG?RCJW M@TGMSQ RF?R RFGQ RWNC MD JC?PLGLE GQ BGDDCPCLR RF?L AMLBGRGMLGLE. "JRFMSEF MLC KGEFR @C RCKNRCB RM @CJGCTC RF?R RFC P?RQ QGKNJW JC?PLCB FMU RM DGLB RFCGP U?W RFPMSEF ? AMLBGRGMLCB QCPGCQ MD PGEFR ?LB JCDR RSPLQ, &.$. 5MJK?L NPMTCB ? BCA?BC J?RCP RF?R RFC P?RQ UCPC K?IGLE ? PCNPCQCLR?RGML MD RFC K?XC GL RFCGP KGLBQ, UFGAF FC A?JJCB ? tAMELGRGTC K?N.u 5FGQ U?Q ?L C?PJW BCKMLQRP?RGML MD RFC NMUCP MD AMELGRGTC JC?PLGLE ?LB FMU RFCQC ?@GJGRGCQ UCPC LMR HSQR JGKGRCB RM FSK?LQ.
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GLAJSBGLE ?EEPCQQGTCLCQQ ?LB MRFCP QMAG?J GLRCP?ARGMLQ, A?L @C CVNJ?GLCB ?JKMQR QMJCJW GL RCPKQ MD ECLCRGAQ ?LB L?RSP?J QCJCARGML. 5FGQ QAGCLAC GQ AMLRPMTCPQG?J; LMRCB QAGCLRGQR QSAF ?Q RFC J?RC 4RCNFCL +?W (MSJB APGRGAGXCB RFC ?NNPM?AF DMP GELMPGLE RFC CLTGPMLKCLR?J CDDCARQ ML @CF?TGMP. 5FGQ GQ ?LMRFCP CV?KNJC MD RFC tL?RSPC TCPQSQ LSPRSPCu BC@?RC MD RFC PMJC MD ECLCRGAQ TCPQSQ RFC PMJC MD CLTGPMLKCLR GL BCRCPKGLGLE ?L MPE?LGQKsQ AF?P?ARCPGQRGAQ. 4MAGM@GMJMEW ?JQM JGLIQ ECLCQ UGRF @CF?TGMPQ ?LB F?Q @CCL ?QQMAG?RCB UGRF t@GMJMEGA?J BCRCPKGLGQK,u RFC @CJGCD RF?R ?JJ @CF?TGMPQ ?PC F?PBUGPCB GLRM MSP ECLCQ. /M MLC BGQNSRCQ RF?R ACPR?GL @CF?TGMPQ A?L @C GLFCPGRCB ?LB RF?R L?RSP?J QCJCARGML NJ?WQ ? PMJC PCR?GLGLE RFCK. *R GQ RFC ?NNJGA?RGML MD QSAF NPGLAGNJCQ RM FSK?L @CF?TGMP RF?R QN?PIQ RFGQ AMLRPMTCPQW, UFGAF PCK?GLQ ?ARGTC RMB?W.
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sQ QRPSARSPC B@E J<C<:K@FE Q?APGDGAGLE MLCsQ MUL JGDC QM RF?R MLCs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#(!04%2 46 X %#/3934%-3
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1370
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PCQC?PAF KCRFMBMJMEGCQ. 4MKC CAMJMEGQRQ QRSBW CAMQWQRCKQ SQGLE AMLRPMJJCB CVNCPGKCLR?J QWQRCKQ, UFGJC QMKC QRSBW CLRGPC CAMQWQRCKQ GL RFCGP L?RSP?J QR?RC, ?LB MRFCPQ SQC @MRF ?NNPM?AFCQ. " holistic ecosystem model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sQ QSPD?AC. 'MP RFCQC PC?QMLQ, QAGCLRGQRQ QRSBW CAMQWQRCKQ SLBCP KMPC AMLRPMJJCB AMLBGRGMLQ. &VNCPGKCLR?J QWQRCKQ SQS?JJW GLTMJTC CGRFCP N?PRGRGMLGLE ? N?PR MD ? L?RSP?J CAMQWQRCK RF?R A?L @C SQCB DMP CVNCPGKCLRQ, RCPKCB ? mesocosm, MP @W PC-APC?RGLE ?L CAMQWQRCK CLRGPCJW GL ?L GLBMMP MP MSRBMMP J?@MP?RMPW CLTGPMLKCLR, UFGAF GQ PCDCPPCB RM ?Q ? microcosm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conceptual model GQ ?L CAMQWQRCK KMBCJ RF?R AMLQGQRQ MD DJMU AF?PRQ RM QFMU GLRCP?ARGMLQ MD BGDDCPCLR AMKN?PRKCLRQ MD RFC JGTGLE ?LB LMLJGTGLE AMKNMLCLRQ MD RFC CAMQWQRCK. " AMLACNRS?J KMBCJ BCQAPG@CQ CAMQWQRCK QRPSARSPC ?LB BWL?KGAQ ?LB QFMUQ FMU CLTGPMLKCLR?J BGQRSP@?LACQ ?DDCAR RFC CAMQWQRCK; FMUCTCP, GRQ ?@GJGRW RM NPCBGAR RFC CDDCARQ MD RFCQC BGQRSP@?LACQ GQ JGKGRCB. "L?JWRGA?J ?LB QGKSJ?RGML KMBCJQ, GL AMLRP?QR, ?PC K?RFCK?RGA?J KCRFMBQ MD BCQAPG@GLE CAMQWQRCKQ RF?R ?PC GLBCCB A?N?@JC MD NPCBGARGLE RFC CDDCARQ MD NMRCLRG?J CLTGPMLKCLR?J AF?LECQ UGRFMSR BGPCAR CVNCPGKCLR?RGML, ?JRFMSEF UGRF QMKC JGKGR?RGMLQ ?Q RM ?AASP?AW. "L analytical model GQ ?L CAMQWQRCK KMBCJ RF?R GQ APC?RCB SQGLE QGKNJC K?RFCK?RGA?J DMPKSJ?Q RM NPCBGAR RFC CDDCARQ MD CLTGPMLKCLR?J BGQRSP@?LACQ ML CAMQWQRCK QRPSARSPC ?LB BWL?KGAQ. " simulation model GQ ?L CAMQWQRCK KMBCJ RF?R GQ APC?RCB SQGLE AMKNJCV AMKNSRCP ?JEMPGRFKQ RM FMJGQRGA?JJW KMBCJ CAMQWQRCKQ ?LB RM NPCBGAR RFC CDDCARQ MD CLTGPMLKCLR?J BGQRSP@?LACQ ML CAMQWQRCK QRPSARSPC ?LB BWL?KGAQ. *BC?JJW, RFCQC KMBCJQ ?PC ?AASP?RC CLMSEF RM BCRCPKGLC UFGAF AMKNMLCLRQ MD RFC CAMQWQRCK ?PC N?PRGASJ?PJW QCLQGRGTC RM BGQRSP@?LACQ, ?LB RFCW A?L QCPTC ?Q ? ESGBC RM CAMQWQRCK K?L?ECPQ (QSAF ?Q AMLQCPT?RGML CAMJMEGQRQ MP DGQFCPGCQ @GMJMEGQRQ) GL RFC NP?ARGA?J K?GLRCL?LAC MD CAMQWQRCK FC?JRF. C43)+59:'1 M4*+18 $MLACNRS?J KMBCJQ ?PC SQCDSJ DMP BCQAPG@GLE CAMQWQRCK QRPSARSPC ?LB BWL?KGAQ ?LB DMP BCKMLQRP?RGLE RFC PCJ?RGMLQFGNQ @CRUCCL BGDDCPCLR MPE?LGQKQ GL ? AMKKSLGRW ?LB RFCGP CLTGPMLKCLR. $MLACNRS?J KMBCJQ ?PC SQS?JJW BCNGARCB EP?NFGA?JJW ?Q DJMU AF?PRQ. 5FC MPE?LGQKQ ?LB RFCGP PCQMSPACQ ?PC EPMSNCB GLRM QNCAGDGA AMKN?PRKCLRQ UGRF ?PPMUQ QFMUGLE RFC PCJ?RGMLQFGN ?LB RP?LQDCP MD CLCPEW MP LSRPGCLRQ @CRUCCL RFCK. 5FSQ, RFCQC BG?EP?KQ ?PC QMKCRGKCQ A?JJCB AMKN?PRKCLR KMBCJQ. 5M KMBCJ RFC AWAJGLE MD KGLCP?J LSRPGCLRQ, MPE?LGA ?LB GLMPE?LGA LSRPGCLRQ ?PC QS@BGTGBCB GLRM RFMQC RF?R ?PC @GM?T?GJ?@JC (PC?BW RM @C GLAMPNMP?RCB GLRM @GMJMEGA?J K?APMKMJCASJCQ) ?LB RFMQC RF?R ?PC LMR. 'MP CV?KNJC, GL ? RCPPCQRPG?J CAMQWQRCK LC?P ? BCNMQGR MD AM?J, A?P@ML UGJJ @C ?T?GJ?@JC RM RFC NJ?LRQ MD RFGQ CAMQWQRCK ?Q A?P@ML BGMVGBC E?Q GL ? QFMPR-RCPK NCPGMB, LMR DPMK RFC A?P@ML-PGAF AM?J GRQCJD. )MUCTCP, MTCP ? JMLECP NCPGMB, KGAPMMPE?LGQKQ A?N?@JC MD BGECQRGLE AM?J UGJJ GLAMPNMP?RC GRQ A?P@ML MP PCJC?QC GR ?Q L?RSP?J E?Q (KCRF?LC, $)4), AF?LEGLE RFGQ SL?T?GJ?@JC MPE?LGA QMSPAC GLRM ?L ?T?GJ?@JC MLC. 5FGQ AMLTCPQGML GQ EPC?RJW ?AACJCP?RCB @W RFC AMK@SQRGML MD DMQQGJ DSCJQ @W FSK?LQ, UFGAF PCJC?QCQ J?PEC ?KMSLRQ MD A?P@ML BGMVGBC GLRM RFC ?RKMQNFCPC. 5FGQ GQ RFMSEFR RM @C ? K?HMP D?ARMP GL RFC PGQC MD RFC ?RKMQNFCPGA A?P@ML BGMVGBC JCTCJQ GL RFC GLBSQRPG?J ?EC. 5FC A?P@ML BGMVGBC PCJC?QCB DPMK @SPLGLE DMQQGJ DSCJQ GQ NPMBSACB D?QRCP RF?L NFMRMQWLRFCRGA MPE?LGQKQ A?L SQC GR. 5FGQ NPMACQQ GQ GLRCLQGDGCB @W RFC PCBSARGML MD NFMRMQWLRFCRGA RPCCQ @CA?SQC MD UMPJBUGBC BCDMPCQR?RGML. .MQR QAGCLRGQRQ ?EPCC RF?R FGEF ?RKMQNFCPGA A?P@ML BGMVGBC GQ ? K?HMP A?SQC MD EJM@?J AJGK?RC AF?LEC. $MLACNRS?J KMBCJQ ?PC ?JQM SQCB RM QFMU RFC DJMU MD CLCPEW RFPMSEF N?PRGASJ?P CAMQWQRCKQ. Figure 46.8 GQ @?QCB ML )MU?PB 5. 0BSKsQ AJ?QQGA?J QRSBW MD RFC 4GJTCP 4NPGLEQ, 'JMPGB?, FMJGQRGA CAMQWQRCK
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1375
GL RFC KGB-RUCLRGCRF ACLRSPW. CAMQWQRCK AMKN?PRKCLRQ.
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5FGQ QRSBW QFMUQ RFC CLCPEW AMLRCLR ?LB RP?LQDCP @CRUCCL T?PGMSQ
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2. )MU?PB 5. 0BSK, t5PMNFGA 4RPSARSPC ?LB 1PMBSARGTGRW MD 4GJTCP 4NPGLEQ, 'JMPGB?,u E*636.0*(3 M656.9(7/: 27, LM. 1 (1957): 47p112.
1376
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sQ CAMQWQRCKQ. 5FCQC CAMQWQRCKQ ?PC MDRCL BCQAPG@CB @W EP?XGLE DMMB UC@Q. 1FMRM?SRMRPMNFQ F?PLCQQ RFC QMJ?P CLCPEW MD RFC QSL @W AMLTCPRGLE GR RM AFCKGA?J CLCPEW GL RFC DMPK MD "51 (?LB /"%1). 5FC CLCPEW QRMPCB GL "51 GQ SQCB RM QWLRFCQGXC AMKNJCV MPE?LGA KMJCASJCQ, QSAF ?Q EJSAMQC.
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1377
Chemoautotrophs ?PC NPGK?PGJW @?ARCPG? RF?R ?PC DMSLB GL P?PC CAMQWQRCKQ UFCPC QSLJGEFR GQ LMR ?T?GJ?@JC, QSAF ?Q GL RFMQC ?QQMAG?RCB UGRF B?PI A?TCQ MP FWBPMRFCPK?J TCLRQ ?R RFC @MRRMK MD RFC MAC?L (Figure 46.9). .?LW AFCKM?SRMRPMNFQ GL FWBPMRFCPK?J TCLRQ SQC FWBPMECL QSJDGBC ()24), UFGAF GQ PCJC?QCB DPMK RFC TCLRQ ?Q ? QMSPAC MD AFCKGA?J CLCPEW. 5FGQ ?JJMUQ AFCKM?SRMRPMNFQ RM QWLRFCQGXC AMKNJCV MPE?LGA KMJCASJCQ, QSAF ?Q EJSAMQC, DMP RFCGP MUL CLCPEW ?LB GL RSPL QSNNJGCQ CLCPEW RM RFC PCQR MD RFC CAMQWQRCK.
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1378
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(CLRPMNW) GL RFC QWQRCK. *L @GMJMEGA QWQRCKQ, RFGQ KC?LQ ? EPC?R BC?J MD CLCPEW GQ JMQR ?Q KCR?@MJGA FC?R UFCL RFC MPE?LGQKQ DPMK MLC RPMNFGA JCTCJ AMLQSKC RFC LCVR JCTCJ. *L RFC 4GJTCP 4NPGLEQ CAMQWQRCK CV?KNJC (Figure 46.8), UC QCC RF?R RFC NPGK?PW AMLQSKCPQ NPMBSACB 1103 IA?J/K2/WP DPMK RFC 7618 IA?J/K2/WP MD CLCPEW ?T?GJ?@JC RM RFCK DPMK RFC NPGK?PW NPMBSACPQ. 5FC KC?QSPCKCLR MD CLCPEW RP?LQDCP CDDGAGCLAW @CRUCCL RUM QSAACQQGTC RPMNFGA JCTCJQ GQ RCPKCB RFC trophic level transfer efficiency (TLTE) ?LB GQ BCDGLCB @W RFC DMPKSJ?:
TLTE =
production at present trophic level 100 production at previous trophic level
*L 4GJTCP 4NPGLEQ, RFC 5-5& @CRUCCL RFC DGPQR RUM RPMNFGA JCTCJQ U?Q ?NNPMVGK?RCJW 14.8 NCPACLR. 5FC JMU CDDGAGCLAW MD CLCPEW RP?LQDCP @CRUCCL RPMNFGA JCTCJQ GQ SQS?JJW RFC K?HMP D?ARMP RF?R JGKGRQ RFC JCLERF MD DMMB AF?GLQ M@QCPTCB GL ? DMMB UC@. 5FC D?AR GQ, ?DRCP DMSP RM QGV CLCPEW RP?LQDCPQ, RFCPC GQ LMR CLMSEF CLCPEW JCDR RM QSNNMPR ?LMRFCP RPMNFGA JCTCJ. *L RFC -?IC 0LR?PGM CV?KNJC QFMUL GL Figure 46.6, MLJW RFPCC CLCPEW RP?LQDCPQ MAASPPCB @CRUCCL RFC NPGK?PW NPMBSACP, (EPCCL ?JE?C), ?LB RFC ?NCV AMLQSKCP ($FGLMMI Q?JKML). &AMJMEGQRQ F?TC K?LW BGDDCPCLR KCRFMBQ MD KC?QSPGLE CLCPEW RP?LQDCPQ UGRFGL CAMQWQRCKQ. 4MKC RP?LQDCPQ ?PC C?QGCP MP KMPC BGDDGASJR RM KC?QSPC BCNCLBGLE ML RFC AMKNJCVGRW MD RFC CAMQWQRCK ?LB FMU KSAF ?AACQQ QAGCLRGQRQ F?TC RM M@QCPTC RFC CAMQWQRCK. *L MRFCP UMPBQ, QMKC CAMQWQRCKQ ?PC KMPC BGDDGASJR RM QRSBW RF?L MRFCPQ, ?LB QMKCRGKCQ RFC OS?LRGDGA?RGML MD CLCPEW RP?LQDCPQ F?Q RM @C CQRGK?RCB. "LMRFCP K?GL N?P?KCRCP RF?R GQ GKNMPR?LR GL AF?P?ARCPGXGLE CLCPEW DJMU UGRFGL ?L CAMQWQRCK GQ RFC LCR NPMBSARGML CDDGAGCLAW. Net production efficiency (NPE) ?JJMUQ CAMJMEGQRQ RM OS?LRGDW FMU CDDGAGCLRJW MPE?LGQKQ MD ? N?PRGASJ?P RPMNFGA JCTCJ GLAMPNMP?RC RFC CLCPEW RFCW PCACGTC GLRM @GMK?QQ; GR GQ A?JASJ?RCB SQGLE RFC DMJJMUGLE DMPKSJ?:
NPE =
net consumer productivity 100 assimilation
Net consumer productivity GQ RFC CLCPEW AMLRCLR ?T?GJ?@JC RM RFC MPE?LGQKQ MD RFC LCVR RPMNFGA JCTCJ. Assimilation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
(F;<C@E> :FJPJK<DJ EP !CFN: :FCF>@:8C +PI8D@;J

5FC QRPSARSPC MD CAMQWQRCKQ A?L @C TGQS?JGXCB UGRF CAMJMEGA?J NWP?KGBQ, UFGAF UCPC DGPQR BCQAPG@CB @W RFC NGMLCCPGLE QRSBGCQ MD $F?PJCQ &JRML GL RFC 1920Q. Ecological pyramids QFMU RFC PCJ?RGTC ?KMSLRQ MD T?PGMSQ N?P?KCRCPQ (QSAF ?Q LSK@CP MD MPE?LGQKQ, CLCPEW, ?LB @GMK?QQ) ?APMQQ RPMNFGA JCTCJQ. 1WP?KGBQ MD LSK@CPQ A?L @C CGRFCP SNPGEFR MP GLTCPRCB, BCNCLBGLE ML RFC CAMQWQRCK. "Q QFMUL GL Figure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1379
"LMRFCP U?W RM TGQS?JGXC CAMQWQRCK QRPSARSPC GQ UGRF NWP?KGBQ MD @GMK?QQ. 5FGQ NWP?KGB KC?QSPCQ RFC ?KMSLR MD CLCPEW AMLTCPRCB GLRM JGTGLE RGQQSC ?R RFC BGDDCPCLR RPMNFGA JCTCJQ. 6QGLE RFC 4GJTCP 4NPGLEQ CAMQWQRCK CV?KNJC, RFGQ B?R? CVFG@GRQ ?L SNPGEFR @GMK?QQ NWP?KGB (Figure 46.10), UFCPC?Q RFC NWP?KGB DPMK RFC &LEJGQF $F?LLCJ CV?KNJC GQ GLTCPRCB. 5FC NJ?LRQ (NPGK?PW NPMBSACPQ) MD RFC 4GJTCP 4NPGLEQ CAMQWQRCK K?IC SN ? J?PEC NCPACLR?EC MD RFC @GMK?QQ DMSLB RFCPC. )MUCTCP, RFC NFWRMNJ?LIRML GL RFC &LEJGQF $F?LLCJ CV?KNJC K?IC SN JCQQ @GMK?QQ RF?L RFC NPGK?PW AMLQSKCPQ, RFC XMMNJ?LIRML. "Q UGRF GLTCPRCB NWP?KGBQ MD LSK@CPQ, RFGQ GLTCPRCB NWP?KGB GQ LMR BSC RM ? J?AI MD NPMBSARGTGRW DPMK RFC NPGK?PW NPMBSACPQ, @SR PCQSJRQ DPMK RFC FGEF RSPLMTCP P?RC MD RFC NFWRMNJ?LIRML. 5FC NFWRMNJ?LIRML ?PC AMLQSKCB P?NGBJW @W RFC NPGK?PW AMLQSKCPQ, RFSQ, KGLGKGXGLE RFCGP @GMK?QQ ?R ?LW N?PRGASJ?P NMGLR GL RGKC. )MUCTCP, NFWRMNJ?LIRML PCNPMBSAC OSGAIJW, RFSQ RFCW ?PC ?@JC RM QSNNMPR RFC PCQR MD RFC CAMQWQRCK. 1WP?KGB CAMQWQRCK KMBCJGLE A?L ?JQM @C SQCB RM QFMU CLCPEW DJMU RFPMSEF RFC RPMNFGA JCTCJQ. /MRGAC RF?R RFCQC LSK@CPQ ?PC RFC Q?KC ?Q RFMQC SQCB GL RFC CLCPEW DJMU AMKN?PRKCLR BG?EP?K GL Figure 46.8. 1WP?KGBQ MD CLCPEW ?PC ?JU?WQ SNPGEFR, ?LB ?L CAMQWQRCK UGRFMSR QSDDGAGCLR NPGK?PW NPMBSARGTGRW A?LLMR @C QSNNMPRCB. "JJ RWNCQ MD CAMJMEGA?J NWP?KGBQ ?PC SQCDSJ DMP AF?P?ARCPGXGLE CAMQWQRCK QRPSARSPC. )MUCTCP, GL RFC QRSBW MD CLCPEW DJMU RFPMSEF RFC CAMQWQRCK, NWP?KGBQ MD CLCPEW ?PC RFC KMQR AMLQGQRCLR ?LB PCNPCQCLR?RGTC KMBCJQ MD CAMQWQRCK QRPSARSPC (Figure 46.10).
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0LC MD RFC KMQR GKNMPR?LR CLTGPMLKCLR?J AMLQCOSCLACQ MD CAMQWQRCK BWL?KGAQ GQ @GMK?ELGDGA?RGML. Biomagnification GQ RFC GLAPC?QGLE AMLACLRP?RGML MD NCPQGQRCLR, RMVGA QS@QR?LACQ GL MPE?LGQKQ ?R C?AF RPMNFGA JCTCJ, DPMK RFC NPGK?PW NPMBSACPQ RM RFC ?NCV AMLQSKCPQ. .?LW QS@QR?LACQ F?TC @CCL QFMUL RM @GM?AASKSJ?RC, GLAJSBGLE AJ?QQGA?J QRSBGCQ UGRF RFC NCQRGAGBC dGAFJMPMdGNFCLWJtPGAFJMPMCRF?LC (%%5), UFGAF U?Q NS@JGQFCB GL RFC 1960Q @CQRQCJJCP, 03,5; 7905., @W 3?AFCJ $?PQML. %%5 U?Q ? AMKKMLJW SQCB NCQRGAGBC @CDMPC GRQ B?LECPQ @CA?KC ILMUL. *L QMKC ?OS?RGA CAMQWQRCKQ, MPE?LGQKQ DPMK C?AF RPMNFGA JCTCJ AMLQSKCB K?LW MPE?LGQKQ MD RFC JMUCP JCTCJ, UFGAF A?SQCB %%5 RM GLAPC?QC GL @GPBQ (?NCV
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46.3 R B@F><F:?<D@:8C CP:C<J

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#(!04%2 46 X %#/3934%-3
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!@>LI< 46.12 /KIV 2.5 MBO@BKQ LC T>QBO LK %>OQE FP COBPE T>QBO, >KA IBPP QE>K 1 MBO@BKQ LC COBPE T>QBO FP B>PFIV >@@BPPF?IB QL IFSFKD QEFKDP.
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1382
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1385
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1386
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s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
&AMQWQRCKQ CVGQR ML J?LB, ?R QC?, GL RFC ?GP, ?LB SLBCPEPMSLB. %GDDCPCLR U?WQ MD KMBCJGLE CAMQWQRCKQ ?PC LCACQQ?PW RM SLBCPQR?LB FMU CLTGPMLKCLR?J BGQRSP@?LACQ UGJJ ?DDCAR CAMQWQRCK QRPSARSPC ?LB BWL?KGAQ. $MLACNRS?J KMBCJQ ?PC SQCDSJ RM QFMU RFC ECLCP?J PCJ?RGMLQFGNQ @CRUCCL MPE?LGQKQ ?LB RFC DJMU MD K?RCPG?JQ MP CLCPEW @CRUCCL RFCK. "L?JWRGA?J KMBCJQ ?PC SQCB RM BCQAPG@C JGLC?P DMMB AF?GLQ, ?LB QGKSJ?RGML KMBCJQ UMPI @CQR UGRF FMJGQRGA DMMB UC@Q.
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0PE?LGQKQ GL ?L CAMQWQRCK ?AOSGPC CLCPEW GL ? T?PGCRW MD U?WQ, UFGAF GQ RP?LQDCPPCB @CRUCCL RPMNFGA JCTCJQ ?Q RFC CLCPEW DJMUQ DPMK RFC @MRRMK RM RFC RMN MD RFC DMMB UC@, UGRF CLCPEW @CGLE JMQR ?R C?AF RP?LQDCP. 5FC CDDGAGCLAW MD RFCQC RP?LQDCPQ GQ GKNMPR?LR DMP SLBCPQR?LBGLE RFC BGDDCPCLR @CF?TGMPQ ?LB
1394
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1. Figure 46.8 8FW BM WMS RFGLI RFC T?JSC DMP EPMQQ NPMBSARGTGRW MD RFC NPGK?PW NPMBSACPQ GQ RFC Q?KC ?Q RFC T?JSC DMP RMR?J FC?R ?LB PCQNGP?RGML (20,810 IA?J/K2/WP) 2. Figure 46.10 1WP?KGBQ BCNGARGLE RFC LSK@CP MD MPE?LGQKQ MP @GMK?QQ K?W @C GLTCPRCB, SNPGEFR, MP CTCL BG?KMLB-QF?NCB. &LCPEW NWP?KGBQ, FMUCTCP, ?PC ?JU?WQ SNPGEFR. 8FW 3. Figure 46.17 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ ?@MSR RFC LGRPMECL AWAJC GQ D?JQC >. "KKMLGDGA?RGML AMLTCPRQ MPE?LGA LGRPMECLMSQ K?RRCP DPMK JGTGLE MPE?LGQKQ GLRM ?KKMLGSK (/)4+). ?. %CLGRPGDGA?RGML @W @?ARCPG? AMLTCPRQ LGRP?RCQ (/03y) RM LGRPMECL E?Q (/2). @. /GRPGDGA?RGML @W @?ARCPG? AMLTCPRQ LGRP?RCQ (/03y) RM LGRPGRCQ (/02y). A. /GRPMECL DGVGLE @?ARCPG? AMLTCPR LGRPMECL E?Q (/2) GLRM MPE?LGA AMKNMSLBQ.
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4. 5FC ?@GJGRW MD ?L CAMQWQRCK RM PCRSPL RM GRQ COSGJG@PGSK QR?RC ?DRCP ?L CLTGPMLKCLR?J BGQRSP@?LAC GQ A?JJCB >>>>>>>>. >. PCQGQR?LAC ?. PCQRMP?RGML @. PCDMPK?RGML A. PCQGJGCLAC 5. " PC-APC?RCB CAMQWQRCK GL ? J?@MP?RMPW CLTGPMLKCLR GQ ILMUL ?Q ? >>>>>>>>. >. KCQMAMQK ?. QGKSJ?RGML @. KGAPMAMQK A. PCNPMBSARGML 6. %CAMKNMQCPQ ?PC ?QQMAG?RCB UGRF UFGAF AJ?QQ MD DMMB UC@ >. EP?XGLE ?. BCRPGR?J @. GLTCPRCB A. ?OS?RGA 7. 5FC NPGK?PW NPMBSACPQ GL ?L MAC?L EP?XGLE DMMB UC@ ?PC SQS?JJW >>>>>>>>. >. NJ?LRQ ?. ?LGK?JQ @. DSLEG A. NFWRMNJ?LIRML 8. 8F?R RCPK BCQAPG@CQ RFC SQC MD K?RFCK?RGA?J COS?RGMLQ GL RFC KMBCJGLE MD JGLC?P ?QNCARQ MD CAMQWQRCKQ >. ?L?JWRGA?J KMBCJGLE ?. QGKSJ?RGML KMBCJGLE @. AMLACNRS?J KMBCJGLE A. GLBGTGBS?J-@?QCB KMBCJGLE 9. 5FC NMQGRGML MD ?L MPE?LGQK ?JMLE ? DMMB AF?GL GQ ILMUL ?Q GRQ >>>>>>>>. >. JMASQ ?. JMA?RGML @. RPMNFGA JCTCJ A. KGAPMAMQK 10. 5FC UCGEFR MD JGTGLE MPE?LGQKQ GL ?L CAMQWQRCK ?R ? N?PRGASJ?P NMGLR GL RGKC GQ A?JJCB: >. ?. @. A. CLCPEW NPMBSARGML CLRPMNW @GMK?QQ
11. 8FGAF RCPK BCQAPG@CQ RFC NPMACQQ UFCPC@W RMVGA QS@QR?LACQ GLAPC?QC ?JMLE RPMNFGA JCTCJQ MD ?L CAMQWQRCK >. @GMK?QQGDGA?RGML ?. @GMK?ELGDGA?RGML @. @GMCLRPMNW A. FCRCPMRPMNFW 12. 0PE?LGQKQ RF?R A?L K?IC RFCGP MUL DMMB SQGLE GLMPE?LGA KMJCASJCQ ?PC A?JJCB: >. ?SRMRPMNFQ ?. FCRCPMRPMNFQ @. NFMRM?SRMRPMNFQ A. AFCKM?SRMRPMNFQ 13. *L RFC &LEJGQF $F?LLCJ CAMQWQRCK, RFC LSK@CP MD NPGK?PW NPMBSACPQ GQ QK?JJCP RF?L RFC LSK@CP
#(!04%2 46 X %#/3934%-3
1395
MD NPGK?PW AMLQSKCPQ @CA?SQC>>>>>>>>. >. RFC ?NCV AMLQSKCPQ F?TC ? JMU RSPLMTCP P?RC ?. RFC NPGK?PW NPMBSACPQ F?TC ? JMU RSPLMTCP P?RC @. RFC NPGK?PW NPMBSACPQ F?TC ? FGEF RSPLMTCP P?RC A. RFC NPGK?PW AMLQSKCPQ F?TC ? FGEF RSPLMTCP P?RC 14. 8F?R J?U MD AFCKGQRPW BCRCPKGLCQ FMU KSAF CLCPEW A?L @C RP?LQDCPPCB UFCL GR GQ AMLTCPRCB DPMK MLC DMPK RM ?LMRFCP >. RFC DGPQR J?U MD RFCPKMBWL?KGAQ ?. RFC QCAMLB J?U MD RFCPKMBWL?KGAQ @. RFC AMLQCPT?RGML MD K?RRCP A. RFC AMLQCPT?RGML MD CLCPEW 15. 5FC KMTCKCLR MD KGLCP?J LSRPGCLRQ RFPMSEF MPE?LGQKQ ?LB RFCGP CLTGPMLKCLR GQ A?JJCB ? >>>>>>>> AWAJC. >. @GMJMEGA?J ?. @GM?AASKSJ?RGML @. @GMECMAFCKGA?J A. @GMAFCKGA?J 16. $?P@ML GQ NPCQCLR GL RFC ?RKMQNFCPC ?Q >>>>>>>>. >. A?P@ML BGMVGBC ?. A?P@ML?RC GML @. A?P@ML BSQR
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18. 5FC ?TCP?EC RGKC ? KMJCASJC QNCLBQ GL GRQ PCQCPTMGP GQ ILMUL ?Q >>>>>>>>. >. PCQGBCLAC RGKC ?. PCQRPGARGML RGKC @. PCQGJGCLAC RGKC A. QRMP?EC RGKC 19. 5FC NPMACQQ UFCPC@W MVWECL GQ BCNJCRCB @W RFC EPMURF MD KGAPMMPE?LGQKQ BSC RM CVACQQ LSRPGCLRQ GL ?OS?RGA QWQRCKQ GQ A?JJCB >>>>>>>>. >. ?. @. A. BC?B XMLGLE CSRPMNFGA?RGML PCRPMDGA?RGML BCNJCRGML
20. 5FC NPMACQQ UFCPC@W LGRPMECL GQ @PMSEFR GLRM MPE?LGA KMJCASJCQ GQ A?JJCB >>>>>>>>. >. LGRPGDGA?RGML ?. BCLGRPGDGA?RGML @. LGRPMECL DGV?RGML A. LGRPMECL AWAJGLE
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1398
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#W RFC CLB MD RFGQ QCARGML, WMS UGJJ @C ?@JC RM: z %CDGLC @GMBGTCPQGRW z %CQAPG@C @GMBGTCPQGRW ?Q RFC COSGJG@PGSK MD L?RSP?JJW DJSARS?RGLE P?RCQ MD CVRGLARGML ?LB QNCAG?RGML z *BCLRGDW FGQRMPGA?J A?SQCQ MD FGEF CVRGLARGML P?RCQ GL &?PRFsQ FGQRMPW 5P?BGRGML?JJW, CAMJMEGQRQ F?TC KC?QSPCB biodiversity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adaptive radiation. "L ?B?NRGTC P?BG?RGML GQ ? P?NGB (JCQQ RF?L RFPCC KGJJGML WC?PQ GL RFC A?QC MD RFC -?IC 7GARMPG? AGAFJGBQ) @P?LAFGLE RFPMSEF QNCAG?RGML MD ? NFWJMECLCRGA RPCC GLRM K?LW AJMQCJW PCJ?RCB QNCAGCQ; RWNGA?JJW, RFC QNCAGCQ tP?BG?RCu GLRM BGDDCPCLR F?@GR?RQ ?LB LGAFCQ. 5FC (?JaN?EMQ DGLAFCQ ?PC ?L CV?KNJC MD ? KMBCQR ?B?NRGTC P?BG?RGML UGRF 15 QNCAGCQ. 5FC AGAFJGBQ MD -?IC 7GARMPG? ?PC ?L CV?KNJC MD ? QNCAR?ASJ?P ?B?NRGTC P?BG?RGML RF?R GLAJSBCQ ?@MSR 500 QNCAGCQ. "R RFC RGKC @GMJMEGQRQ UCPC K?IGLE RFGQ BGQAMTCPW, QMKC QNCAGCQ @CE?L RM OSGAIJW BGQ?NNC?P. " ASJNPGR GL RFCQC BCAJGLCQ U?Q ? QNCAGCQ MD J?PEC DGQF RF?R U?Q GLRPMBSACB RM -?IC 7GARMPG? @W DGQFCPGCQ RM DCCB RFC NCMNJC JGTGLE ?PMSLB RFC J?IC. 5FC /GJC NCPAF U?Q GLRPMBSACB GL 1963, @SR J?W JMU SLRGJ RFC 1980Q UFCL GRQ NMNSJ?RGMLQ @CE?L RM QSPEC. 5FC /GJC NCPAF NMNSJ?RGML EPCU @W AMLQSKGLE AGAFJGBQ, BPGTGLE QNCAGCQ ?DRCP QNCAGCQ RM RFC NMGLR MD extinction (RFC BGQ?NNC?P?LAC MD ? QNCAGCQ). *L D?AR, RFCPC UCPC QCTCP?J D?ARMPQ RF?R NJ?WCB ? PMJC GL RFC CVRGLARGML MD NCPF?NQ 200 AGAFJGB QNCAGCQ GL -?IC 7GARMPG?: RFC /GJC NCPAF, BCAJGLGLE J?IC U?RCP OS?JGRW BSC RM ?EPGASJRSPC ?LB J?LB AJC?PGLE ML RFC QFMPCQ MD -?IC 7GARMPG?, ?LB GLAPC?QCB DGQFGLE NPCQQSPC. 4AGCLRGQRQ F?B LMR CTCL A?R?JMESCB ?JJ MD RFC QNCAGCQ NPCQCLRqQM K?LW UCPC JMQR RF?R UCPC LCTCP L?KCB. 5FC BGTCPQGRW GQ LMU ? QF?BMU MD UF?R GR MLAC U?Q. 5FC AGAFJGBQ MD -?IC 7GARMPG? ?PC ? RFSK@L?GJ QICRAF MD AMLRCKNMP?PW P?NGB QNCAGCQ JMQQ RF?R MAASPQ ?JJ MTCP &?PRF ?LB GQ A?SQCB @W FSK?L ?ARGTGRW. &VRGLARGML GQ ? L?RSP?J NPMACQQ MD K?APMCTMJSRGML RF?R MAASPQ ?R RFC P?RC MD ?@MSR MLC MSR MD 1 KGJJGML QNCAGCQ @CAMKGLE CVRGLAR NCP WC?P. 5FC DMQQGJ PCAMPB PCTC?JQ RF?R RFCPC F?TC @CCL DGTC NCPGMBQ MD K?QQ CVRGLARGML GL FGQRMPW UGRF KSAF FGEFCP P?RCQ MD QNCAGCQ JMQQ, ?LB RFC P?RC MD QNCAGCQ JMQQ RMB?W GQ AMKN?P?@JC RM RFMQC NCPGMBQ MD K?QQ CVRGLARGML. )MUCTCP, RFCPC GQ ? K?HMP BGDDCPCLAC @CRUCCL RFC NPCTGMSQ K?QQ CVRGLARGMLQ ?LB RFC ASPPCLR CVRGLARGML UC ?PC CVNCPGCLAGLE: FSK?L ?ARGTGRW. 4NCAGDGA?JJW, RFPCC FSK?L ?ARGTGRGCQ F?TC ? K?HMP GKN?AR: BCQRPSARGML MD F?@GR?R, GLRPMBSARGML MD CVMRGA QNCAGCQ, ?LB MTCP-F?PTCQRGLE. 1PCBGARGMLQ MD QNCAGCQ JMQQ UGRFGL RFC LCVR ACLRSPW, ? RGLW ?KMSLR MD RGKC ML ECMJMEGA?J RGKCQA?JCQ, P?LEC DPMK 10 NCPACLR RM 50 NCPACLR. &VRGLARGMLQ ML RFGQ QA?JC F?TC MLJW F?NNCLCB DGTC MRFCP RGKCQ GL RFC FGQRMPW MD RFC NJ?LCR, ?LB RFCW F?TC @CCL A?SQCB @W A?R?AJWQKGA CTCLRQ RF?R AF?LECB RFC AMSPQC MD RFC FGQRMPW MD JGDC GL C?AF GLQR?LAC. &?PRF GQ LMU GL MLC MD RFMQC RGKCQ.
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1400
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RFC QSL ?LB K?W ?AAMSLR DMP AJGK?RC AF?LECQ M@QCPTCB ?R RFC RGKC. 5FC FWNMRFCQGQ GQ QNCASJ?RGTC, @SR CVRP?RCPPCQRPG?J GLDJSCLACQ ML &?PRFsQ FGQRMPW ?PC ?L ?ARGTC JGLC MD PCQC?PAF. 3CAMTCPW MD @GMBGTCPQGRW ?DRCP RFC K?QQ CVRGLARGML RMMI DPMK 5 RM 20 KGJJGML WC?PQ, BCNCLBGLE ML RFC JMA?RGML. 5FC J?RC %CTMLG?L CVRGLARGML K?W F?TC MAASPPCB MTCP ? PCJ?RGTCJW JMLE NCPGMB MD RGKC. *R ?NNC?PQ RM F?TC ?DDCARCB K?PGLC QNCAGCQ ?LB LMR RFC NJ?LRQ MP ?LGK?JQ GLF?@GRGLE RCPPCQRPG?J F?@GR?RQ. 5FC A?SQCQ MD RFGQ CVRGLARGML ?PC NMMPJW SLBCPQRMMB. 5FC CLB-1CPKG?L CVRGLARGML U?Q RFC J?PECQR GL RFC FGQRMPW MD JGDC. *LBCCB, ?L ?PESKCLR AMSJB @C K?BC RF?R &?PRF LC?PJW @CA?KC BCTMGB MD JGDC BSPGLE RFGQ CVRGLARGML CTCLR. 5FC NJ?LCR JMMICB TCPW BGDDCPCLR @CDMPC ?LB ?DRCP RFGQ CTCLR. &QRGK?RCQ ?PC RF?R 96 NCPACLR MD ?JJ K?PGLC QNCAGCQ ?LB 70 NCPACLR MD ?JJ RCPPCQRPG?J QNCAGCQ UCPC JMQR. *R U?Q ?R RFGQ RGKC, DMP CV?KNJC, RF?R RFC RPGJM@GRCQ, ? EPMSN RF?R QSPTGTCB RFC 0PBMTGAG?Lp4GJSPG?L CVRGLARGML, @CA?KC CVRGLAR. 5FC A?SQCQ DMP RFGQ K?QQ CVRGLARGML ?PC LMR AJC?P, @SR RFC JC?BGLE QSQNCAR GQ CVRCLBCB ?LB UGBCQNPC?B TMJA?LGA ?ARGTGRW RF?R JCB RM ? PSL?U?W EJM@?J-U?PKGLE CTCLR. 5FC MAC?LQ @CA?KC J?PECJW ?LMVGA, QSDDMA?RGLE K?PGLC JGDC. 5CPPCQRPG?J RCRP?NMB BGTCPQGRW RMMI 30 KGJJGML WC?PQ RM PCAMTCP ?DRCP RFC CLB-1CPKG?L CVRGLARGML. 5FC 1CPKG?L CVRGLARGML BP?K?RGA?JJW ?JRCPCB &?PRFsQ @GMBGTCPQGRW K?ICSN ?LB RFC AMSPQC MD CTMJSRGML. 5FC A?SQCQ MD RFC 5PG?QQGAp+SP?QQGA CVRGLARGML CTCLR ?PC LMR AJC?P ?LB FWNMRFCQCQ MD AJGK?RC AF?LEC, ?QRCPMGB GKN?AR, ?LB TMJA?LGA CPSNRGMLQ F?TC @CCL ?PESCB. 5FC CVRGLARGML CTCLR MAASPPCB HSQR @CDMPC RFC @PC?ISN MD RFC QSNCPAMLRGLCLR 1?LE?C?, ?JRFMSEF PCACLR QAFMJ?PQFGN QSEECQRQ RF?R RFC CVRGLARGMLQ K?W F?TC MAASPPCB KMPC EP?BS?JJW RFPMSEFMSR RFC 5PG?QQGA. 5FC A?SQCQ MD RFC CLB-$PCR?ACMSQ CVRGLARGML CTCLR ?PC RFC MLCQ RF?R ?PC @CQR SLBCPQRMMB. *R U?Q BSPGLE RFGQ CVRGLARGML CTCLR ?@MSR 65 KGJJGML WC?PQ ?EM RF?R RFC BGLMQ?SPQ, RFC BMKGL?LR TCPRC@P?RC EPMSN DMP KGJJGMLQ MD WC?PQ, BGQ?NNC?PCB DPMK RFC NJ?LCR (UGRF RFC CVACNRGML MD ? RFCPMNMB AJ?BC RF?R E?TC PGQC RM @GPBQ). *LBCCB, CTCPW J?LB ?LGK?J RF?R UCGEFCB KMPC RFCL 25 IE @CA?KC CVRGLAR. 5FC A?SQC MD RFGQ CVRGLARGML GQ LMU SLBCPQRMMB RM @C RFC PCQSJR MD ? A?R?AJWQKGA GKN?AR MD ? J?PEC KCRCMPGRC, MP ?QRCPMGB, MDD RFC AM?QR MD UF?R GQ LMU RFC :SA?RaL 1CLGLQSJ?. 5FGQ FWNMRFCQGQ, NPMNMQCB DGPQR GL 1980, U?Q ? P?BGA?J CVNJ?L?RGML @?QCB ML ? QF?PN QNGIC GL RFC JCTCJQ MD GPGBGSK (UFGAF P?GLQ BMUL DPMK QN?AC GL KCRCMPQ ?R ? D?GPJW AMLQR?LR P?RC @SR GQ MRFCPUGQC ?@QCLR ML &?PRFsQ QSPD?AC) ?R RFC PMAI QRP?RSK RF?R K?PIQ RFC @MSLB?PW @CRUCCL RFC $PCR?ACMSQ ?LB 1?JCMECLC NCPGMBQ (Figure 47.6). 5FGQ @MSLB?PW K?PICB RFC BGQ?NNC?P?LAC MD RFC BGLMQ?SPQ GL DMQQGJQ ?Q UCJJ ?Q K?LW MRFCP R?V?. 5FC PCQC?PAFCPQ UFM BGQAMTCPCB RFC GPGBGSK QNGIC GLRCPNPCRCB GR ?Q ? P?NGB GLDJSV MD GPGBGSK DPMK QN?AC RM RFC ?RKMQNFCPC (GL RFC DMPK MD ? J?PEC ?QRCPMGB) P?RFCP RF?L ? QJMUGLE GL RFC BCNMQGRGML MD QCBGKCLRQ BSPGLE RF?R NCPGMB. *R U?Q ? P?BGA?J CVNJ?L?RGML, @SR RFC PCNMPR MD ?L ?NNPMNPG?RCJW ?ECB ?LB QGXCB GKN?AR AP?RCP GL 1991 K?BC RFC FWNMRFCQGQ KMPC @CJGCT?@JC. /MU ?L ?@SLB?LAC MD ECMJMEGA?J CTGBCLAC QSNNMPRQ RFC RFCMPW. 3CAMTCPW RGKCQ DMP @GMBGTCPQGRW ?DRCP RFC CLB-$PCR?ACMSQ CVRGLARGML ?PC QFMPRCP, GL ECMJMEGA?J RGKC, RF?L DMP RFC CLB-1CPKG?L CVRGLARGML, ML RFC MPBCP MD 10 KGJJGML WC?PQ.
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3@FBKQFPQP JB>PROBA QEB OBI>QFSB >?RKA>K@B LC CBOK PMLOBP >?LSB >KA ?BILT QEB +t0D ?LRKA>OV FK QEFP OL@H P>JMIB. 7EF@E LC QEB CLIILTFKD PQ>QBJBKQP JLPQ IFHBIV OBMOBPBKQP QEBFO CFKAFKDP >. !K >?RKA>K@B LC CBOK PMLOBP COLJ PBSBO>I PMB@FBP T>P CLRKA ?BILT QEB +t0D ?LRKA>OV, ?RQ KLKB T>P CLRKA >?LSB. ?. !K >?RKA>K@B LC CBOK PMLOBP COLJ PBSBO>I PMB@FBP T>P CLRKA >?LSB QEB +t0D ?LRKA>OV, ?RQ KLKB T>P CLRKA ?BILT. @. !K >?RKA>K@B LC CBOK PMLOBP T>P CLRKA ?LQE >?LSB >KA ?BILT QEB +t0D ?LRKA>OV, ?RQ LKIV LKB PMB@FBP T>P CLRKA ?BILT QEB ?LRKA>OV, >KA J>KV PMB@FBP TBOB CLRKA >?LSB QEB ?LRKA>OV. A. ->KV PMB@FBP LC CBOK PMLOBP TBOB CLRKA ?LQE >?LSB >KA ?BILT QEB ?LRKA>OV, ?RQ QEB QLQ>I KRJ?BO LC PMLOBP T>P DOB>QBO ?BILT QEB ?LRKA>OV.
&VNJMPC RFGQ interactive website (http://openstaxcollege.org/l/extinctions) ?@MSR K?QQ CVRGLARGMLQ. ".+ P1+/894)+3+ E=9/3)9/43 5FC 1JCGQRMACLC &VRGLARGML GQ MLC MD RFC JCQQCP CVRGLARGMLQ, ?LB ? PCACLR MLC. *R GQ UCJJ ILMUL RF?R RFC /MPRF "KCPGA?L, ?LB RM QMKC BCEPCC &SP?QG?L, megafauna, MP J?PEC ?LGK?JQ, BGQ?NNC?PCB RMU?PB RFC CLB MD RFC J?QR EJ?AG?RGML NCPGMB. 5FC CVRGLARGML ?NNC?PQ RM F?TC F?NNCLCB GL ? PCJ?RGTCJW PCQRPGARCB RGKC NCPGMB MD 10,000p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"KCPGA?, RFC CVRGLARGMLQ MD ?JKMQR ?JJ MD RFC J?PEC K?KK?JQ MAASPPCB 10,000p12,000 WC?PQ ?EM. "JJ RF?R ?PC JCDR ?PC RFC QK?JJCP K?KK?JQ QSAF ?Q @C?PQ, CJI, KMMQC, ?LB AMSE?PQ. 'GL?JJW, ML K?LW PCKMRC MAC?LGA GQJ?LBQ, RFC CVRGLARGMLQ MD K?LW QNCAGCQ MAASPPCB AMGLAGBCLR UGRF FSK?L ?PPGT?JQ. /MR ?JJ MD RFC GQJ?LBQ F?B J?PEC ?LGK?JQ, @SR UFCL RFCPC UCPC J?PEC ?LGK?JQ, RFCW UCPC JMQR. .?B?E?QA?P U?Q AMJMLGXCB ?@MSR 2,000 WC?PQ ?EM ?LB RFC J?PEC K?KK?JQ RF?R JGTCB RFCPC @CA?KC CVRGLAR. &SP?QG? ?LB "DPGA? BM LMR QFMU RFGQ N?RRCPL, @SR RFCW ?JQM BGB LMR CVNCPGCLAC ? PCACLR ?PPGT?J MD FSK?LQ. )SK?LQ ?PPGTCB GL &SP?QG? FSLBPCBQ MD RFMSQ?LBQ MD WC?PQ ?EM ?DRCP RFC MPGEGL MD RFC QNCAGCQ GL "DPGA?. 5FGQ RMNGA PCK?GLQ ?L ?PC? MD ?ARGTC PCQC?PAF ?LB FWNMRFCQGXGLE. *R QCCKQ AJC?P RF?R CTCL GD AJGK?RC NJ?WCB ? PMJC, GL KMQR A?QCQ FSK?L FSLRGLE NPCAGNGR?RCB RFC CVRGLARGMLQ. P7+8+39-"/2+ E=9/3)9/438 5FC QGVRF, MP )MJMACLC, K?QQ CVRGLARGML ?NNC?PQ RM F?TC @CESL C?PJGCP RF?L NPCTGMSQJW @CJGCTCB ?LB F?Q KMQRJW RM BM UGRF RFC ?ARGTGRGCQ MD H646 :(70,5:. 4GLAC RFC @CEGLLGLE MD RFC )MJMACLC NCPGMB, RFCPC ?PC LSKCPMSQ PCACLR CVRGLARGMLQ MD GLBGTGBS?J QNCAGCQ RF?R ?PC PCAMPBCB GL FSK?L UPGRGLEQ. .MQR MD RFCQC ?PC AMGLAGBCLR UGRF RFC CVN?LQGML MD RFC &SPMNC?L AMJMLGCQ QGLAC RFC 1500Q. 0LC MD RFC C?PJGCP ?LB NMNSJ?PJW ILMUL CV?KNJCQ GQ RFC BMBM @GPB. 5FC BMBM @GPB JGTCB GL RFC DMPCQRQ MD .?SPGRGSQ, ?L GQJ?LB GL RFC *LBG?L 0AC?L. 5FC BMBM @GPB @CA?KC CVRGLAR ?PMSLB 1662. *R U?Q FSLRCB DMP GRQ KC?R @W Q?GJMPQ ?LB U?Q C?QW NPCW @CA?SQC RFC BMBM, UFGAF BGB LMR CTMJTC UGRF FSK?LQ, UMSJB ?NNPM?AF NCMNJC UGRFMSR DC?P. *LRPMBSACB NGEQ, P?RQ, ?LB BMEQ @PMSEFR RM RFC GQJ?LB @W &SPMNC?L QFGNQ ?JQM IGJJCB BMBM WMSLE ?LB CEEQ. 4RCJJCP'Q QC? AMU @CA?KC CVRGLAR GL 1768; GR U?Q PCJ?RCB RM RFC K?L?RCC ?LB NPM@?@JW MLAC JGTCB ?JMLE RFC LMPRFUCQR AM?QR MD /MPRF "KCPGA?. 4RCJJCP'Q QC? AMU U?Q DGPQR BGQAMTCPCB @W &SPMNC?LQ GL 1741 ?LB U?Q FSLRCB DMP KC?R ?LB MGJ. 5FC J?QR QC? AMU U?Q IGJJCB GL 1768. 5F?R ?KMSLRQ RM 27 WC?PQ @CRUCCL RFC QC? AMUsQ DGPQR AMLR?AR UGRF &SPMNC?LQ ?LB CVRGLARGML MD RFC QNCAGCQ. *L 1914, RFC J?QR JGTGLE N?QQCLECP NGECML BGCB GL ? XMM GL $GLAGLL?RG, 0FGM. 5FGQ QNCAGCQ F?B MLAC B?PICLCB RFC QIGCQ MD /MPRF "KCPGA? BSPGLE GRQ KGEP?RGMLQ, @SR GR U?Q FSLRCB ?LB QSDDCPCB DPMK F?@GR?R JMQQ RFPMSEF RFC AJC?PGLE MD DMPCQRQ DMP D?PKJ?LB. *L 1918, RFC J?QR JGTGLE $?PMJGL? N?P?ICCR BGCB GL A?NRGTGRW. 5FGQ QNCAGCQ U?Q MLAC AMKKML GL RFC C?QRCPL 6LGRCB 4R?RCQ, @SR GR QSDDCPCB DPMK F?@GR?R JMQQ. 5FC QNCAGCQ U?Q ?JQM FSLRCB @CA?SQC GR ?RC MPAF?PB DPSGR UFCL GRQ L?RGTC DMMBQ UCPC BCQRPMWCB RM K?IC U?W DMP D?PKJ?LB. 5FC +?N?LCQC QC? JGML, UFGAF GLF?@GRCB ? @PM?B ?PC? ?PMSLB +?N?L ?LB RFC AM?QR MD ,MPC?, @CA?KC CVRGLAR GL RFC 1950Q BSC RM DGQFCPKCL. 5FC $?PG@@C?L KMLI QC?J U?Q BGQRPG@SRCB RFPMSEFMSR RFC $?PG@@C?L 4C? @SR U?Q BPGTCL RM CVRGLARGML TG? FSLRGLE @W 1952. 5FCQC ?PC MLJW ? DCU MD RFC PCAMPBCB CVRGLARGMLQ GL RFC N?QR 500 WC?PQ. 5FC *LRCPL?RGML?J 6LGML DMP $MLQCPT?RGML MD /?RSPC (*6$/) ICCNQ ? JGQR MD CVRGLAR ?LB CLB?LECPCB QNCAGCQ A?JJCB RFC 3CB -GQR. 5FC JGQR GQ LMR AMKNJCRC, @SR GR BCQAPG@CQ 380 CVRGLAR QNCAGCQ MD TCPRC@P?RCQ ?DRCP 1500 "%, 86 MD UFGAF UCPC BPGTCL CVRGLAR @W MTCPFSLRGLE MP MTCPDGQFGLE. E89/2'9+8 4, P7+8+39-"/2+ E=9/3)9/43 '9+8 &QRGK?RCQ MD extinction rates ?PC F?KNCPCB @W RFC D?AR RF?R KMQR CVRGLARGMLQ ?PC NPM@?@JW F?NNCLGLE UGRFMSR M@QCPT?RGML. 5FC CVRGLARGML MD ? @GPB MP K?KK?J GQ JGICJW RM @C LMRGACB @W FSK?LQ, CQNCAG?JJW GD GR F?Q @CCL FSLRCB MP SQCB GL QMKC MRFCP U?W. #SR RFCPC ?PC K?LW MPE?LGQKQ RF?R ?PC MD JCQQ GLRCPCQR RM FSK?LQ (LMR LCACQQ?PGJW MD JCQQ T?JSC) ?LB K?LW RF?R ?PC SLBCQAPG@CB. 5FC @?AIEPMSLB CVRGLARGML P?RC GQ CQRGK?RCB RM @C ?@MSR MLC NCP KGJJGML QNCAGCQ NCP WC?P (&/.4:). 'MP CV?KNJC, ?QQSKGLE RFCPC ?PC ?@MSR RCL KGJJGML QNCAGCQ GL CVGQRCLAC, RFC CVNCAR?RGML GQ RF?R RCL QNCAGCQ UMSJB @CAMKC CVRGLAR C?AF WC?P (C?AF WC?P PCNPCQCLRQ RCL KGJJGML QNCAGCQ NCP WC?P). 0LC AMLRCKNMP?PW CVRGLARGML P?RC CQRGK?RC SQCQ RFC CVRGLARGMLQ GL RFC UPGRRCL PCAMPB QGLAC RFC WC?P 1500. 'MP @GPBQ ?JMLC RFGQ KCRFMB WGCJBQ ?L CQRGK?RC MD 26 &/.4:. )MUCTCP, RFGQ T?JSC K?W @C SLBCPCQRGK?RCB DMP RFPCC PC?QMLQ. 'GPQR, K?LW QNCAGCQ UMSJB LMR F?TC @CCL BCQAPG@CB SLRGJ KSAF J?RCP GL RFC RGKC NCPGMB, QM RFCGP JMQQ UMSJB F?TC EMLC SLLMRGACB. 4CAMLB, RFC LSK@CP MD PCACLRJW CVRGLAR QNCAGCQ GQ GLAPC?QGLE @CA?SQC CVRGLAR QNCAGCQ LMU ?PC @CGLE BCQAPG@CB DPMK QICJCR?J PCK?GLQ. "LB RFGPB, QMKC QNCAGCQ ?PC NPM@?@JW ?JPC?BW CVRGLAR CTCL RFMSEF AMLQCPT?RGMLGQRQ ?PC PCJSAR?LR RM L?KC RFCK ?Q QSAF. 5?IGLE RFCQC D?ARMPQ GLRM ?AAMSLR P?GQCQ RFC CQRGK?RCB CVRGLARGML P?RC AJMQCP RM 100 &/.4:. 5FC NPCBGARCB P?RC @W RFC CLB MD RFC ACLRSPW GQ 1500 &/.4:. " QCAMLB ?NNPM?AF RM CQRGK?RGLE NPCQCLR-RGKC CVRGLARGML P?RCQ GQ RM AMPPCJ?RC QNCAGCQ JMQQ UGRF F?@GR?R JMQQ @W KC?QSPGLE DMPCQR-?PC? JMQQ ?LB SLBCPQR?LBGLE QNCAGCQ-?PC? PCJ?RGMLQFGNQ. 5FC species-area relationship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UMPI F?Q ?JQM A?JJCB GLRM OSCQRGML RFC ?NNJGA?@GJGRW MD RFC QNCAGCQ-?PC? PCJ?RGMLQFGN UFCL CQRGK?RGLE RFC JMQQ MD QNCAGCQ. 5FGQ UMPI ?PESCQ RF?R RFC QNCAGCQ-?PC? PCJ?RGMLQFGN JC?BQ RM ?L MTCPCQRGK?RC MD CVRGLARGML P?RCQ. " @CRRCP PCJ?RGMLQFGN RM SQC K?W @C RFC CLBCKGAQ-?PC? PCJ?RGMLQFGN. 6QGLE RFGQ KCRFMB UMSJB @PGLE CQRGK?RCQ BMUL RM ?PMSLB 500 &/.4: GL RFC AMKGLE ACLRSPW. /MRC RF?R RFGQ T?JSC GQ QRGJJ 500 RGKCQ RFC @?AIEPMSLB P?RC.
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1408
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D?KGLC UFCL RFC QGLEJC T?PGCRW EPMUL GL *PCJ?LB @CA?KC QSQACNRG@JC RM ? NMR?RM @JGEFR, UGNGLE MSR RFC APMN. 5FC JMQQ MD RFC APMN JCB RM D?KGLC, BC?RF, ?LB K?QQ CKGEP?RGML. 3CQGQR?LAC RM BGQC?QC GQ ? AFGCD @CLCDGR RM K?GLR?GLGLE APMN @GMBGTCPQGRW, ?LB J?AI MD BGTCPQGRW GL AMLRCKNMP?PW APMN QNCAGCQ A?PPGCQ QGKGJ?P PGQIQ. 4CCB AMKN?LGCQ, UFGAF ?PC RFC QMSPAC MD KMQR APMN T?PGCRGCQ GL BCTCJMNCB AMSLRPGCQ, KSQR AMLRGLS?JJW @PCCB LCU T?PGCRGCQ RM ICCN SN UGRF CTMJTGLE NCQR MPE?LGQKQ. 5FCQC Q?KC QCCB AMKN?LGCQ, FMUCTCP, F?TC N?PRGAGN?RCB GL RFC BCAJGLC MD RFC LSK@CP MD T?PGCRGCQ ?T?GJ?@JC ?Q RFCW DMASQ ML QCJJGLE DCUCP T?PGCRGCQ GL KMPC ?PC?Q MD RFC UMPJB. 5FC ?@GJGRW RM APC?RC LCU APMN T?PGCRGCQ PCJGCQ ML RFC BGTCPQGRW MD T?PGCRGCQ ?T?GJ?@JC ?LB RFC ?AACQQG@GJGRW MD UGJB DMPKQ PCJ?RCB RM RFC APMN NJ?LR. 5FCQC UGJB DMPKQ ?PC MDRCL RFC QMSPAC MD LCU ECLC T?PG?LRQ RF?R A?L @C @PCB UGRF CVGQRGLE T?PGCRGCQ RM APC?RC T?PGCRGCQ UGRF LCU ?RRPG@SRCQ. -MQQ MD UGJB QNCAGCQ PCJ?RCB RM ? APMN UGJJ KC?L RFC JMQQ MD NMRCLRG?J GL APMN GKNPMTCKCLR. .?GLR?GLGLE RFC ECLCRGA BGTCPQGRW MD UGJB QNCAGCQ PCJ?RCB RM BMKCQRGA?RCB QNCAGCQ CLQSPCQ MSP AMLRGLSCB DMMB QSNNJW. 4GLAC RFC 1920Q, EMTCPLKCLR ?EPGASJRSPC BCN?PRKCLRQ F?TC K?GLR?GLCB QCCB @?LIQ MD APMN T?PGCRGCQ ?Q ? U?W RM K?GLR?GL APMN BGTCPQGRW. 5FGQ QWQRCK F?Q DJ?UQ @CA?SQC, MTCP RGKC, QCCB @?LIQ ?PC JMQR RFPMSEF ?AAGBCLRQ, ?LB RFCPC GQ LM U?W RM PCNJ?AC RFCK. *L 2008, RFC 4T?J@?PB (JM@?J 4CCB 7?SJR (Figure 47.9) @CE?L QRMPGLE QCCBQ DPMK ?PMSLB RFC UMPJB ?Q ? @?AISN QWQRCK RM RFC PCEGML?J QCCB @?LIQ. *D ? PCEGML?J QCCB @?LI QRMPCQ T?PGCRGCQ GL 4T?J@?PB, JMQQCQ A?L @C PCNJ?ACB DPMK 4T?J@?PB. 5FC QCCB T?SJR GQ JMA?RCB BCCN GLRM RFC PMAI MD ?L ?PARGA GQJ?LB. $MLBGRGMLQ UGRFGL RFC T?SJR ?PC K?GLR?GLCB ?R GBC?J RCKNCP?RSPC ?LB FSKGBGRW DMP QCCB QSPTGT?J, @SR RFC BCCN SLBCPEPMSLB JMA?RGML MD RFC T?SJR GL RFC ?PARGA KC?LQ RF?R D?GJSPC MD RFC T?SJRsQ QWQRCKQ UGJJ LMR AMKNPMKGQC RFC AJGK?RGA AMLBGRGMLQ GLQGBC RFC T?SJR.
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1410
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@CC QNCAGCQ ?LB T?PGMSQ GLQCARQ ?LB @GPBQ. -MQQ MD RFCQC QNCAGCQ UMSJB K?IC EPMUGLE APMNQ PCOSGPGLE NMJJGL?RGML GKNMQQG@JC, GLAPC?QGLE BCNCLBCLAC ML MRFCP APMNQ. 'GL?JJW, FSK?LQ AMKNCRC DMP RFCGP DMMB UGRF APMN NCQRQ, KMQR MD UFGAF ?PC GLQCARQ. 1CQRGAGBCQ AMLRPMJ RFCQC AMKNCRGRMPQ; FMUCTCP, NCQRGAGBCQ ?PC AMQRJW ?LB JMQC RFCGP CDDCARGTCLCQQ MTCP RGKC ?Q NCQR NMNSJ?RGMLQ ?B?NR. 5FCW ?JQM JC?B RM AMJJ?RCP?J B?K?EC @W IGJJGLE LML-NCQR QNCAGCQ ?LB PGQIGLE RFC FC?JRF MD AMLQSKCPQ ?LB ?EPGASJRSP?J UMPICPQ. &AMJMEGQRQ @CJGCTC RF?R RFC @SJI MD RFC UMPI GL PCKMTGLE NCQRQ GQ ?ARS?JJW BMLC @W NPCB?RMPQ ?LB N?P?QGRCQ MD RFMQC NCQRQ, @SR RFC GKN?AR F?Q LMR @CCL UCJJ QRSBGCB. " PCTGCU DMSLB RF?R GL 74 NCPACLR MD QRSBGCQ RF?R JMMICB DMP ?L CDDCAR MD J?LBQA?NC AMKNJCVGRW ML L?RSP?J CLCKGCQ MD NCQRQ, RFC EPC?RCP RFC AMKNJCVGRW, RFC EPC?RCP RFC CDDCAR MD NCQR-QSNNPCQQGLE MPE?LGQKQ. "L CVNCPGKCLR?J QRSBW DMSLB RF?R GLRPMBSAGLE KSJRGNJC CLCKGCQ MD NC? ?NFGBQ (?L GKNMPR?LR ?JD?JD? NCQR) GLAPC?QCB RFC WGCJB MD ?JD?JD? QGELGDGA?LRJW. 5FGQ QRSBW QFMUQ RFC GKNMPR?LAC MD J?LBQA?NC BGTCPQGRW TG? RFC OSCQRGML MD UFCRFCP ? BGTCPQGRW MD NCQRQ GQ KMPC CDDCARGTC ?R AMLRPMJ RF?L MLC QGLEJC NCQR; RFC PCQSJRQ QFMUCB RFGQ RM @C RFC A?QC. -MQQ MD BGTCPQGRW GL NCQR CLCKGCQ UGJJ GLCTGR?@JW K?IC GR KMPC BGDDGASJR ?LB AMQRJW RM EPMU DMMB.
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1411
5FC AMPC RFPC?R RM @GMBGTCPQGRW ML RFC NJ?LCR, ?LB RFCPCDMPC ? RFPC?R RM FSK?L UCJD?PC, GQ RFC AMK@GL?RGML MD FSK?L NMNSJ?RGML EPMURF ?LB PCQMSPAC CVNJMGR?RGML. 5FC FSK?L NMNSJ?RGML PCOSGPCQ PCQMSPACQ RM QSPTGTC ?LB EPMU, ?LB RFMQC PCQMSPACQ ?PC @CGLE PCKMTCB SLQSQR?GL?@JW DPMK RFC CLTGPMLKCLR. 5FC RFPCC EPC?RCQR NPMVGK?RC RFPC?RQ RM @GMBGTCPQGRW ?PC F?@GR?R JMQQ, MTCPF?PTCQRGLE, ?LB GLRPMBSARGML MD CVMRGA QNCAGCQ. 5FC DGPQR RUM MD RFCQC ?PC ? BGPCAR PCQSJR MD FSK?L NMNSJ?RGML EPMURF ?LB PCQMSPAC SQC. 5FC RFGPB PCQSJRQ DPMK GLAPC?QCB KM@GJGRW ?LB RP?BC. " DMSPRF K?HMP A?SQC MD CVRGLARGML, ?LRFPMNMECLGA AJGK?RC AF?LEC, F?Q LMR WCR F?B ? J?PEC GKN?AR, @SR GR GQ NPCBGARCB RM @CAMKC QGELGDGA?LR BSPGLE RFGQ ACLRSPW. (JM@?J AJGK?RC AF?LEC GQ ?JQM ? AMLQCOSCLAC MD FSK?L NMNSJ?RGML LCCBQ DMP CLCPEW ?LB RFC SQC MD DMQQGJ DSCJQ RM KCCR RFMQC LCCBQ (Figure 47.10). &LTGPMLKCLR?J GQQSCQ, QSAF ?Q RMVGA NMJJSRGML, F?TC QNCAGDGA R?PECRCB CDDCARQ ML QNCAGCQ, @SR RFCW ?PC LMR ECLCP?JJW QCCL ?Q RFPC?RQ ?R RFC K?ELGRSBC MD RFC MRFCPQ.
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)SK?LQ PCJW ML RCAFLMJMEW RM KMBGDW RFCGP CLTGPMLKCLR ?LB PCNJ?AC ACPR?GL DSLARGMLQ RF?R UCPC MLAC NCPDMPKCB @W RFC L?RSP?J CAMQWQRCK. 0RFCP QNCAGCQ A?LLMR BM RFGQ. &JGKGL?RGML MD RFCGP CAMQWQRCKqUFCRFCP GR GQ ? DMPCQR, ? BCQCPR, ? EP?QQJ?LB, ? DPCQFU?RCP CQRS?PGLC, MP ? K?PGLC CLTGPMLKCLRqUGJJ IGJJ RFC GLBGTGBS?JQ GL RFC QNCAGCQ. 3CKMTC RFC CLRGPC F?@GR?R UGRFGL RFC P?LEC MD ? QNCAGCQ ?LB, SLJCQQ RFCW ?PC MLC MD RFC DCU QNCAGCQ RF?R BM UCJJ GL FSK?L-@SGJR CLTGPMLKCLRQ, RFC QNCAGCQ UGJJ @CAMKC CVRGLAR. )SK?L BCQRPSARGML MD F?@GR?RQ ?AACJCP?RCB GL RFC J?RRCP F?JD MD RFC RUCLRGCRF ACLRSPW. $MLQGBCP RFC CVACNRGML?J @GMBGTCPQGRW MD 4SK?RP?: GR GQ FMKC RM MLC QNCAGCQ MD MP?LESR?L, ? QNCAGCQ MD APGRGA?JJW CLB?LECPCB CJCNF?LR, ?LB RFC 4SK?RP?L RGECP, @SR F?JD MD 4SK?RP?sQ DMPCQR GQ LMU EMLC. 5FC LCGEF@MPGLE GQJ?LB MD #MPLCM, FMKC RM RFC MRFCP QNCAGCQ MD MP?LESR?L, F?Q JMQR ? QGKGJ?P ?PC? MD DMPCQR. 'MPCQR JMQQ AMLRGLSCQ GL NPMRCARCB ?PC?Q MD #MPLCM. 5FC MP?LESR?L GL #MPLCM GQ JGQRCB ?Q CLB?LECPCB @W RFC *LRCPL?RGML?J 6LGML DMP $MLQCPT?RGML MD /?RSPC (*6$/), @SR GR GQ QGKNJW RFC KMQR TGQG@JC MD RFMSQ?LBQ MD QNCAGCQ RF?R UGJJ LMR QSPTGTC RFC BGQ?NNC?P?LAC MD RFC DMPCQRQ MD #MPLCM. 5FC DMPCQRQ ?PC PCKMTCB DMP RGK@CP ?LB RM NJ?LR N?JK MGJ NJ?LR?RGMLQ (Figure 47.11). 1?JK MGJ GQ SQCB GL K?LW NPMBSARQ GLAJSBGLE DMMB NPMBSARQ, AMQKCRGAQ, ?LB @GMBGCQCJ GL &SPMNC. " DGTC-WC?P CQRGK?RC MD EJM@?J DMPCQR AMTCP JMQQ DMP RFC WC?PQ 2000p2005 U?Q 3.1 NCPACLR. *L RFC FSKGB RPMNGAQ UFCPC DMPCQR JMQQ GQ NPGK?PGJW DPMK RGK@CP CVRP?ARGML, 272,000 IK2 U?Q JMQR MSR MD ? EJM@?J RMR?J MD 11,564,000 IK2 (MP 2.4 NCPACLR). *L RFC RPMNGAQ, RFCQC JMQQCQ ACPR?GLJW ?JQM PCNPCQCLR RFC CVRGLARGML MD QNCAGCQ @CA?SQC MD FGEF JCTCJQ MD CLBCKGQK.
1412
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1413
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-LPQ @LKPRJBOP AL KLQ FJ>DFKB QE>Q QEB ELJB FJMOLSBJBKQ MOLAR@QP QEBV ?RV JFDEQ ?B @LKQOF?RQFKD QL E>?FQ>Q ILPP >KA PMB@FBP BUQFK@QFLKP. 9BQ QEB J>OHBQ CLO FIIBD>IIV E>OSBPQBA QOLMF@>I QFJ?BO FP ERDB, >KA QEB TLLA MOLAR@QP LCQBK CFKA QEBJPBISBP FK ?RFIAFKD PRMMIV PQLOBP FK QEB 5KFQBA 3Q>QBP. /KB BPQFJ>QB FP QE>Q 10 MBO@BKQ LC QEB FJMLOQBA QFJ?BO PQOB>J FK QEB 5KFQBA 3Q>QBP, TEF@E FP QEB TLOIAw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w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sQ RCPPGRMPG?J U?RCPQ. $MKKML PCQMSPACQ ?PC QS@HCAR RM ?L CAMLMKGA NPCQQSPC ILMUL ?Q RFC tragedy of the commons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qUF?JCQ ?PC ?L CV?KNJCqCAMLMKGA DMPACQ UGJJ ?JU?WQ BPGTC RMU?PB DGQFGLE RFC NMNSJ?RGML RM CVRGLARGML.
1414
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&VNJMPC ? 6.4. 'GQF & 8GJBJGDC 4CPTGAC interactive map (http://openstaxcollege.org/l/habitat_map) MD APGRGA?J F?@GR?R DMP CLB?LECPCB ?LB RFPC?RCLCB QNCAGCQ GL RFC 6LGRCB 4R?RCQ. 5M @CEGL, QCJCAR t7GQGR RFC MLJGLC K?NNCP.u 'MP RFC KMQR N?PR, DGQFCPW CVRGLARGML GQ LMR COSGT?JCLR RM @GMJMEGA?J CVRGLARGMLqRFC J?QR DGQF MD ? QNCAGCQ GQ P?PCJW DGQFCB MSR MD RFC MAC?L. "R RFC Q?KC RGKC, DGQFCPW CVRGLARGML GQ QRGJJ F?PKDSJ RM DGQF QNCAGCQ ?LB RFCGP CAMQWQRCKQ. 5FCPC ?PC QMKC GLQR?LACQ GL UFGAF RPSC CVRGLARGML GQ ? NMQQG@GJGRW. 8F?JCQ F?TC QJMU-EPMUGLE NMNSJ?RGMLQ ?LB ?PC ?R PGQI MD AMKNJCRC CVRGLARGML RFPMSEF FSLRGLE. 5FCPC ?PC QMKC QNCAGCQ MD QF?PIQ UGRF PCQRPGARCB BGQRPG@SRGMLQ RF?R ?PC ?R PGQI MD CVRGLARGML. 5FC EPMSNCPQ ?PC ?LMRFCP NMNSJ?RGML MD ECLCP?JJW QJMU-EPMUGLE DGQFCQ RF?R, GL RFC $?PG@@C?L, GLAJSBCQ ? LSK@CP MD QNCAGCQ RF?R ?PC ?R PGQI MD CVRGLARGML DPMK MTCPDGQFGLE. $MP?J PCCDQ ?PC CVRPCKCJW BGTCPQC K?PGLC CAMQWQRCKQ RF?R D?AC NCPGJ DPMK QCTCP?J NPMACQQCQ. 3CCDQ ?PC FMKC RM 1/3 MD RFC UMPJBsQ K?PGLC DGQF QNCAGCQq?@MSR 4,000 QNCAGCQq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
7GCU ? brief video (http://openstaxcollege.org/l/ocean_matters) BGQASQQGLE RFC PMJC MD K?PGLC CAMQWQRCKQ GL QSNNMPRGLE FSK?L UCJD?PC ?LB RFC BCAJGLC MD MAC?L CAMQWQRCKQ. Bush meat GQ RFC ECLCPGA RCPK SQCB DMP UGJB ?LGK?JQ IGJJCB DMP DMMB. )SLRGLE GQ NP?ARGACB RFPMSEFMSR RFC UMPJB, @SR FSLRGLE NP?ARGACQ, N?PRGASJ?PJW GL COS?RMPG?J "DPGA? ?LB N?PRQ MD "QG?, ?PC @CJGCTCB RM RFPC?RCL QCTCP?J QNCAGCQ UGRF CVRGLARGML. 5P?BGRGML?JJW, @SQF KC?R GL "DPGA? U?Q FSLRCB RM DCCB D?KGJGCQ BGPCARJW; FMUCTCP, PCACLR AMKKCPAG?JGX?RGML MD RFC NP?ARGAC LMU F?Q @SQF KC?R ?T?GJ?@JC GL EPMACPW QRMPCQ, UFGAF F?Q GLAPC?QCB F?PTCQR P?RCQ RM RFC JCTCJ MD SLQSQR?GL?@GJGRW. "BBGRGML?JJW, FSK?L NMNSJ?RGML EPMURF F?Q GLAPC?QCB RFC LCCB DMP NPMRCGL DMMBQ RF?R ?PC LMR @CGLE KCR DPMK ?EPGASJRSPC. 4NCAGCQ RFPC?RCLCB @W RFC @SQF KC?R RP?BC ?PC KMQRJW K?KK?JQ GLAJSBGLE K?LW NPGK?RCQ JGTGLE GL RFC $MLEM @?QGL.
OFK@: .G<:@<J
Exotic species ?PC QNCAGCQ RF?R F?TC @CCL GLRCLRGML?JJW MP SLGLRCLRGML?JJW GLRPMBSACB @W FSK?LQ GLRM ?L CAMQWQRCK GL UFGAF RFCW BGB LMR CTMJTC. 4SAF GLRPMBSARGMLQ JGICJW MAASP DPCOSCLRJW ?Q L?RSP?J NFCLMKCL?. 'MP CV?KNJC, ,SBXS (P<,9(90( 36)(;(), UFGAF GQ L?RGTC RM +?N?L, U?Q GLRPMBSACB GL RFC 6LGRCB 4R?RCQ GL 1876. *R U?Q J?RCP NJ?LRCB DMP QMGJ AMLQCPT?RGML. 1PM@JCK?RGA?JJW, GR EPMUQ RMM UCJJ GL RFC QMSRFC?QRCPL 6LGRCB 4R?RCQqSN RM ? DMMR ? B?W. *R GQ LMU ? NCQR QNCAGCQ ?LB AMTCPQ MTCP 7 KGJJGML ?APCQ GL RFC QMSRFC?QRCPL 6LGRCB 4R?RCQ. *D ?L GLRPMBSACB QNCAGCQ GQ ?@JC RM QSPTGTC GL GRQ LCU F?@GR?R, RF?R GLRPMBSARGML GQ LMU PCDJCARCB GL RFC M@QCPTCB P?LEC MD RFC QNCAGCQ. )SK?L RP?LQNMPR?RGML MD NCMNJC ?LB EMMBQ, GLAJSBGLE RFC GLRCLRGML?J RP?LQNMPR MD MPE?LGQKQ DMP RP?BC, F?Q BP?K?RGA?JJW GLAPC?QCB RFC GLRPMBSARGML MD QNCAGCQ GLRM LCU CAMQWQRCKQ, QMKCRGKCQ ?R BGQR?LACQ RF?R ?PC UCJJ @CWMLB RFC A?N?AGRW MD RFC QNCAGCQ RM CTCP RP?TCJ GRQCJD ?LB MSRQGBC RFC P?LEC MD RFC QNCAGCQs L?RSP?J NPCB?RMPQ.
#(!04%2 47 X #/.3%26!4)/. ")/,/'9 !.$ ")/$)6%23)49
1415
.MQR CVMRGA QNCAGCQ GLRPMBSARGMLQ NPM@?@JW D?GJ @CA?SQC MD RFC JMU LSK@CP MD GLBGTGBS?JQ GLRPMBSACB MP NMMP ?B?NR?RGML RM RFC CAMQWQRCK RFCW CLRCP. 4MKC QNCAGCQ, FMUCTCP, NMQQCQQ NPC?B?NR?RGMLQ RF?R A?L K?IC RFCK CQNCAG?JJW QSAACQQDSJ GL ? LCU CAMQWQRCK. 5FCQC CVMRGA QNCAGCQ MDRCL SLBCPEM BP?K?RGA NMNSJ?RGML GLAPC?QCQ GL RFCGP LCU F?@GR?R ?LB PCQCR RFC CAMJMEGA?J AMLBGRGMLQ GL RFC LCU CLTGPMLKCLR, RFPC?RCLGLE RFC QNCAGCQ RF?R CVGQR RFCPC. 'MP RFGQ PC?QML, CVMRGA QNCAGCQ ?PC ?JQM A?JJCB GLT?QGTC QNCAGCQ. &VMRGA QNCAGCQ A?L RFPC?RCL MRFCP QNCAGCQ RFPMSEF AMKNCRGRGML DMP PCQMSPACQ, NPCB?RGML, MP BGQC?QC.
&VNJMPC ?L interactive global database (http://openstaxcollege.org/l/exotic_invasive) MD CVMRGA MP GLT?QGTC QNCAGCQ. -?ICQ ?LB GQJ?LBQ ?PC N?PRGASJ?PJW TSJLCP?@JC RM CVRGLARGML RFPC?RQ DPMK GLRPMBSACB QNCAGCQ. *L -?IC 7GARMPG?, ?Q KCLRGMLCB C?PJGCP, RFC GLRCLRGML?J GLRPMBSARGML MD RFC /GJC NCPAF U?Q J?PECJW PCQNMLQG@JC DMP RFC CVRGLARGML MD ?@MSR 200 QNCAGCQ MD AGAFJGBQ. 5FC ?AAGBCLR?J GLRPMBSARGML MD RFC @PMUL RPCC QL?IC TG? ?GPAP?DR (Figure 47.12) DPMK RFC 4MJMKML *QJ?LBQ RM (S?K GL 1950 F?Q JCB RM RFC CVRGLARGML MD RFPCC QNCAGCQ MD @GPBQ ?LB RFPCC RM DGTC QNCAGCQ MD PCNRGJCQ CLBCKGA RM RFC GQJ?LB. 4CTCP?J MRFCP QNCAGCQ ?PC QRGJJ RFPC?RCLCB. 5FC @PMUL RPCC QL?IC GQ ?BCNR ?R CVNJMGRGLE FSK?L RP?LQNMPR?RGML ?Q ? KC?LQ RM KGEP?RC; MLC U?Q CTCL DMSLB ML ?L ?GPAP?DR ?PPGTGLE GL $MPNSQ $FPGQRG, 5CV?Q. $MLQR?LR TGEGJ?LAC ML RFC N?PR MD ?GPNMPR, KGJGR?PW, ?LB AMKKCPAG?J ?GPAP?DR NCPQMLLCJ GQ PCOSGPCB RM NPCTCLR RFC QL?IC DPMK KMTGLE DPMK (S?K RM MRFCP GQJ?LBQ GL RFC 1?AGDGA, CQNCAG?JJW )?U?GG. *QJ?LBQ BM LMR K?IC SN ? J?PEC ?PC? MD J?LB ML RFC EJM@C, @SR RFCW BM AMLR?GL ? BGQNPMNMPRGML?RC LSK@CP MD CLBCKGA QNCAGCQ @CA?SQC MD RFCGP GQMJ?RGML DPMK K?GLJ?LB ?LACQRMPQ.
!@>LI< 47.12 4EB ?OLTK QOBB PK>HB, B60.( 099,.<3(90:, FP >K BULQF@ PMB@FBP QE>Q E>P @>RPBA KRJBOLRP BUQFK@QFLKP LK QEB FPI>KA LC 'R>J PFK@B FQP >@@FABKQ>I FKQOLAR@QFLK FK 1950. (@OBAFQ: .03)
*R LMU ?NNC?PQ RF?R RFC EJM@?J BCAJGLC GL ?KNFG@G?L QNCAGCQ PCAMELGXCB GL RFC 1990Q GQ, GL QMKC N?PR, A?SQCB @W RFC DSLESQ B(;9(*/6*/@;90<4 +,5+96)(;0+0:, UFGAF A?SQCQ RFC BGQC?QC chytridiomycosis (Figure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
1416
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1417
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1418
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1419
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1420
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AMKKGRRCB AMSLRPGCQ RM PCBSAGLE EPCCLFMSQC E?Q CKGQQGMLQ @W 2012, U?Q P?RGDGCB @W QMKC AMSLRPGCQ, @SR QNSPLCB @W MRFCPQ. 5UM GKNMPR?LR AMSLRPGCQ GL RCPKQ MD RFCGP NMRCLRG?J GKN?AR RF?R BGB LMR P?RGDW RFC ,WMRM 1PMRMAMJ UCPC RFC 6LGRCB 4R?RCQ ?LB $FGL?. 5FC 6LGRCB 4R?RCQ PCHCARCB GR ?Q ? PCQSJR MD ? NMUCPDSJ DMQQGJ DSCJ GLBSQRPW ?LB $FGL? @CA?SQC MD ? AMLACPL GR UMSJB QRGDJC RFC L?RGMLs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&QR?@JGQFKCLR MD UGJBJGDC ?LB CAMQWQRCK NPCQCPTCQ GQ MLC MD RFC ICW RMMJQ GL AMLQCPT?RGML CDDMPRQ. " NPCQCPTC GQ ?L ?PC? MD J?LB QCR ?QGBC UGRF T?PWGLE BCEPCCQ MD NPMRCARGML DMP RFC MPE?LGQKQ RF?R CVGQR UGRFGL RFC @MSLB?PGCQ MD RFC NPCQCPTC. 1PCQCPTCQ A?L @C CDDCARGTC GL RFC QFMPR RCPK DMP NPMRCARGLE @MRF QNCAGCQ ?LB CAMQWQRCKQ, @SR RFCW D?AC AF?JJCLECQ RF?R QAGCLRGQRQ ?PC QRGJJ CVNJMPGLE RM QRPCLERFCL RFCGP TG?@GJGRW ?Q JMLE-RCPK QMJSRGMLQ. H4< M:). A7+' 94 P7+8+7;+? %SC RM RFC U?W NPMRCARCB J?LBQ ?PC ?JJMA?RCB (RFCW RCLB RM AMLR?GL JCQQ CAMLMKGA?JJW T?JS?@JC PCQMSPACQ P?RFCP RF?L @CGLE QCR ?QGBC QNCAGDGA?JJW DMP RFC QNCAGCQ MP CAMQWQRCKQ ?R PGQI) ?LB RFC U?W @GMBGTCPQGRW GQ BGQRPG@SRCB, BCRCPKGLGLE ? R?PECR NCPACLR?EC MD J?LB MP K?PGLC F?@GR?R RF?R QFMSJB @C NPMRCARCB RM K?GLR?GL @GMBGTCPQGRW JCTCJQ GQ AF?JJCLEGLE. 5FC *6$/ 8MPJB 1?PIQ $MLEPCQQ CQRGK?RCB RF?R 11.5 NCPACLR MD &?PRFs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tGQJ?LBQu MD F?@GR?R UGRFGL t?L MAC?Lu MD LML-F?@GR?R. 'MP ? QNCAGCQ RM NCPQGQR GL ? NPCQCPTC, RFC NPCQCPTC KSQR @C J?PEC CLMSEF. 5FC APGRGA?J QGXC BCNCLBQ, GL N?PR, ML RFC FMKC P?LEC RF?R GQ AF?P?ARCPGQRGA MD RFC QNCAGCQ. " NPCQCPTC DMP UMJTCQ, UFGAF P?LEC FSLBPCBQ MD IGJMKCRCPQ, KSQR @C KSAF J?PECP RF?L ? NPCQCPTC DMP @SRRCPDJGCQ, UFGAF KGEFR P?LEC UGRFGL RCL IGJMKCRCPQ BSPGLE GRQ JGDCRGKC. #SR J?PECP NPCQCPTCQ F?TC KMPC AMPC ?PC? MD MNRGK?J F?@GR?R DMP GLBGTGBS?J QNCAGCQ, RFCW F?TC KMPC LGAFCQ RM QSNNMPR KMPC QNCAGCQ, ?LB RFCW ?RRP?AR KMPC QNCAGCQ @CA?SQC RFCW A?L @C DMSLB ?LB PC?AFCB KMPC C?QGJW. 1PCQCPTCQ NCPDMPK @CRRCP UFCL RFCPC ?PC @SDDCP XMLCQ ?PMSLB RFCK MD QS@MNRGK?J F?@GR?R. 5FC @SDDCP ?JJMUQ MPE?LGQKQ RM CVGR RFC @MSLB?PGCQ MD RFC NPCQCPTC UGRFMSR GKKCBG?RC LCE?RGTC AMLQCOSCLACQ DPMK NPCB?RGML MP J?AI MD PCQMSPACQ. 0LC J?PEC NPCQCPTC GQ @CRRCP RF?L RFC Q?KC ?PC? MD QCTCP?J QK?JJCP
5. 3M@CPR ). .?A"PRFSP ?LB &BU?PB 0. 8GJQML, &. 0., !/, !/,69@ 6- I:3(5+ B06.,6.9(7/@ (1PGLACRML, /.+.: 1PGLACRML 6LGTCPQGRW 1PCQQ, 1967).
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NPCQCPTCQ @CA?SQC RFCPC GQ KMPC AMPC F?@GR?R SL?DDCARCB @W CBECQ. 'MP RFGQ Q?KC PC?QML, NPCQCPTCQ GL RFC QF?NC MD ? QOS?PC MP AGPAJC UGJJ @C @CRRCP RF?L ? NPCQCPTC UGRF K?LW RFGL t?PKQ.u *D NPCQCPTCQ KSQR @C QK?JJCP, RFCL NPMTGBGLE UGJBJGDC AMPPGBMPQ @CRUCCL RFCK QM RF?R GLBGTGBS?JQ ?LB RFCGP ECLCQ A?L KMTC @CRUCCL RFC NPCQCPTCQ, DMP CV?KNJC ?JMLE PGTCPQ ?LB QRPC?KQ, UGJJ K?IC RFC QK?JJCP NPCQCPTCQ @CF?TC KMPC JGIC ? J?PEC MLC. "JJ MD RFCQC D?ARMPQ ?PC R?ICL GLRM AMLQGBCP?RGML UFCL NJ?LLGLE RFC L?RSPC MD ? NPCQCPTC @CDMPC RFC J?LB GQ QCR ?QGBC. *L ?BBGRGML RM RFC NFWQGA?J, @GMJMEGA?J, ?LB CAMJMEGA?J QNCAGDGA?RGMLQ MD ? NPCQCPTC, RFCPC ?PC ? T?PGCRW MD NMJGAW, JCEGQJ?RGTC, ?LB CLDMPACKCLR QNCAGDGA?RGMLQ PCJ?RCB RM SQCQ MD RFC NPCQCPTC DMP DSLARGMLQ MRFCP RF?L NPMRCARGML MD QNCAGCQ. 5FCQC A?L GLAJSBC ?LWRFGLE DPMK RGK@CP CVRP?ARGML, KGLCP?J CVRP?ARGML, PCESJ?RCB FSLRGLE, FSK?L F?@GR?RGML, ?LB LMLBCQRPSARGTC FSK?L PCAPC?RGML. .?LW MD RFCQC NMJGAW BCAGQGMLQ ?PC K?BC @?QCB ML NMJGRGA?J NPCQQSPCQ P?RFCP RF?L AMLQCPT?RGML AMLQGBCP?RGMLQ. *L QMKC A?QCQ, UGJBJGDC NPMRCARGML NMJGAGCQ F?TC @CCL QM QRPGAR RF?R QS@QGQRCLAC-JGTGLE GLBGECLMSQ NMNSJ?RGMLQ F?TC @CCL DMPACB DPMK ?LACQRP?J J?LBQ RF?R DCJJ UGRFGL ? NPCQCPTC. *L MRFCP A?QCQ, CTCL GD ? NPCQCPTC GQ BCQGELCB RM NPMRCAR UGJBJGDC, GD RFC NPMRCARGMLQ ?PC LMR MP A?LLMR @C CLDMPACB, RFC NPCQCPTC QR?RSQ UGJJ F?TC JGRRJC KC?LGLE GL RFC D?AC MD GJJCE?J NM?AFGLE ?LB RGK@CP CVRP?ARGML. 5FGQ GQ ? UGBCQNPC?B NPM@JCK UGRF NPCQCPTCQ GL ?PC?Q MD RFC RPMNGAQ. L/2/9'9/438 43 P7+8+7;+8 4MKC MD RFC JGKGR?RGMLQ ML NPCQCPTCQ ?Q AMLQCPT?RGML RMMJQ ?PC CTGBCLR DPMK RFC BGQASQQGML MD NPCQCPTC BCQGEL. 1MJGRGA?J ?LB CAMLMKGA NPCQQSPCQ RWNGA?JJW K?IC NPCQCPTCQ QK?JJCP, LCTCP J?PECP, QM QCRRGLE ?QGBC ?PC?Q RF?R ?PC J?PEC CLMSEF GQ BGDDGASJR. *D RFC ?PC? QCR ?QGBC GQ QSDDGAGCLRJW J?PEC, RFCPC K?W LMR @C QSDDGAGCLR ?PC? RM APC?RC ? @SDDCP ?PMSLB RFC NPCQCPTC. *L RFGQ A?QC, ?L ?PC? ML RFC MSRCP CBECQ MD RFC NPCQCPTC GLCTGR?@JW @CAMKCQ ? PGQIGCP QS@MNRGK?J F?@GR?R DMP RFC QNCAGCQ GL RFC NPCQCPTC. &LDMPACKCLR MD NPMRCARGMLQ GQ ?JQM ? QGELGDGA?LR GQQSC GL AMSLRPGCQ UGRFMSR RFC PCQMSPACQ MP NMJGRGA?J UGJJ RM NPCTCLR NM?AFGLE ?LB GJJCE?J PCQMSPAC CVRP?ARGML. $JGK?RC AF?LEC UGJJ APC?RC GLCTGR?@JC NPM@JCKQ UGRF RFC JMA?RGML MD NPCQCPTCQ. 5FC QNCAGCQ UGRFGL RFCK UGJJ KGEP?RC RM FGEFCP J?RGRSBCQ ?Q RFC F?@GR?R MD RFC NPCQCPTC @CAMKCQ JCQQ D?TMP?@JC. 4AGCLRGQRQ ?PC NJ?LLGLE DMP RFC CDDCARQ MD EJM@?J U?PKGLE ML DSRSPC NPCQCPTCQ ?LB QRPGTGLE RM NPCBGAR RFC LCCB DMP LCU NPCQCPTCQ RM ?AAMKKMB?RC ?LRGAGN?RCB AF?LECQ RM F?@GR?RQ; FMUCTCP, RFC CLB CDDCARGTCLCQQ GQ RCLSMSQ QGLAC RFCQC CDDMPRQ ?PC NPCBGARGML @?QCB. 'GL?JJW, ?L ?PESKCLR A?L @C K?BC RF?R AMLQCPT?RGML NPCQCPTCQ PCGLDMPAC RFC ASJRSP?J NCPACNRGML RF?R FSK?LQ ?PC QCN?P?RC DPMK L?RSPC, A?L CVGQR MSRQGBC MD GR, ?LB A?L MLJW MNCP?RC GL U?WQ RF?R BM B?K?EC RM @GMBGTCPQGRW. $PC?RGLE NPCQCPTCQ PCBSACQ RFC NPCQQSPC ML FSK?L ?ARGTGRGCQ MSRQGBC RFC NPCQCPTCQ RM @C QSQR?GL?@JC ?LB LML-B?K?EGLE RM @GMBGTCPQGRW. 6JRGK?RCJW, RFC NMJGRGA?J, CAMLMKGA, ?LB FSK?L BCKMEP?NFGA NPCQQSPCQ UGJJ BCEP?BC ?LB PCBSAC RFC QGXC MD AMLQCPT?RGML NPCQCPTCQ GD RFC ?ARGTGRGCQ MSRQGBC RFCK ?PC LMR ?JRCPCB RM @C JCQQ B?K?EGLE RM @GMBGTCPQGRW.
"L interactive global data system (http://openstaxcollege.org/l/protected_areas) MD NPMRCARCB ?PC?Q A?L @C DMSLB ?R UC@QGRC. 3CTGCU B?R? ?@MSR GLBGTGBS?J NPMRCARCB ?PC?Q @W JMA?RGML MP QRSBW QR?RGQRGAQ ML NPMRCARCB ?PC?Q @W AMSLRPW MP PCEGML. H'(/9'9 +8947'9/43 )?@GR?R PCQRMP?RGML FMJBQ AMLQGBCP?@JC NPMKGQC ?Q ? KCAF?LGQK DMP PCQRMPGLE ?LB K?GLR?GLGLE @GMBGTCPQGRW. 0D AMSPQC MLAC ? QNCAGCQ F?Q @CAMKC CVRGLAR, GRQ PCQRMP?RGML GQ GKNMQQG@JC. )MUCTCP, PCQRMP?RGML A?L GKNPMTC RFC @GMBGTCPQGRW MD BCEP?BCB CAMQWQRCKQ. 3CGLRPMBSAGLE UMJTCQ, ? RMN NPCB?RMP, RM :CJJMUQRMLC /?RGML?J 1?PI GL 1995 JCB RM BP?K?RGA AF?LECQ GL RFC CAMQWQRCK RF?R GLAPC?QCB @GMBGTCPQGRW. 5FC UMJTCQ (Figure 47.17) DSLARGML RM QSNNPCQQ CJI ?LB AMWMRC NMNSJ?RGMLQ ?LB NPMTGBC KMPC ?@SLB?LR PCQMSPACQ RM RFC ESGJB MD A?PPGML C?RCPQ. 3CBSAGLE CJI NMNSJ?RGMLQ F?Q ?JJMUCB PCTCECR?RGML MD PGN?PG?L ?PC?Q, UFGAF F?Q GLAPC?QCB RFC BGTCPQGRW MD QNCAGCQ GL RF?R F?@GR?R. %CAPC?QGLE RFC AMWMRC NMNSJ?RGML F?Q GLAPC?QCB RFC NMNSJ?RGMLQ MD QNCAGCQ RF?R UCPC NPCTGMSQJW QSNNPCQQCB @W RFGQ NPCB?RMP. 5FC LSK@CP MD QNCAGCQ MD A?PPGML C?RCPQ F?Q GLAPC?QCB @CA?SQC MD RFC NPCB?RMPW ?ARGTGRGCQ MD RFC UMJTCQ. *L RFGQ F?@GR?R, RFC UMJD GQ ? ICWQRMLC QNCAGCQ, KC?LGLE ? QNCAGCQ RF?R GQ GLQRPSKCLR?J GL K?GLR?GLGLE BGTCPQGRW GL ?L CAMQWQRCK. 3CKMTGLE ? ICWQRMLC QNCAGCQ DPMK ?L CAMJMEGA?J AMKKSLGRW
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K?W A?SQC ? AMJJ?NQC GL BGTCPQGRW. 5FC PCQSJRQ DPMK RFC :CJJMUQRMLC CVNCPGKCLR QSEECQR RF?R PCQRMPGLE ? ICWQRMLC QNCAGCQ A?L F?TC RFC CDDCAR MD PCQRMPGLE @GMBGTCPQGRW GL RFC AMKKSLGRW. &AMJMEGQRQ F?TC ?PESCB DMP RFC GBCLRGDGA?RGML MD ICWQRMLC QNCAGCQ UFCPC NMQQG@JC ?LB DMP DMASQGLE NPMRCARGML CDDMPRQ ML RFMQC QNCAGCQ; JGICUGQC, GR ?JQM K?ICQ QCLQC RM ?RRCKNR RM PCRSPL RFCK RM RFCGP CAMQWQRCK GD RFCW F?TC @CCL PCKMTCB.
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sQ ?RRGRSBCQ ?LB ?ARGMLQ PCE?PBGLE AMLQCPT?RGML; ?R NPCQCLR, RFC PCQSJRQ RCLB RM @C KGVCB.
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& 4 / -(.
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p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
C#A+/ - .0((A-4
47.1 /?< B@F;@M<IJ@KP CI@J@J
#GMBGTCPQGRW CVGQRQ ?R KSJRGNJC JCTCJQ MD MPE?LGX?RGML ?LB GQ KC?QSPCB GL BGDDCPCLR U?WQ BCNCLBGLE ML RFC EM?JQ MD RFMQC R?IGLE RFC KC?QSPCKCLRQ. 5FCQC KC?QSPCKCLRQ GLAJSBC LSK@CPQ MD QNCAGCQ, ECLCRGA BGTCPQGRW, AFCKGA?J BGTCPQGRW, ?LB CAMQWQRCK BGTCPQGRW. 5FC LSK@CP MD BCQAPG@CB QNCAGCQ GQ CQRGK?RCB RM @C 1.5 KGJJGML UGRF ?@MSR 17,000 LCU QNCAGCQ @CGLE BCQAPG@CB C?AF WC?P. &QRGK?RCQ DMP RFC RMR?J LSK@CP
1424
#(!04%2 47 X #/.3%26!4)/. ")/,/'9 !.$ ")/$)6%23)49
MD QNCAGCQ ML &?PRF T?PW @SR ?PC ML RFC MPBCP MD 10 KGJJGML. #GMBGTCPQGRW GQ LCE?RGTCJW AMPPCJ?RCB UGRF J?RGRSBC DMP KMQR R?V?, KC?LGLE RF?R @GMBGTCPQGRW GQ FGEFCP GL RFC RPMNGAQ. 5FC KCAF?LGQK DMP RFGQ N?RRCPL GQ LMR ILMUL UGRF ACPR?GLRW, @SR QCTCP?J NJ?SQG@JC FWNMRFCQCQ F?TC @CCL ?BT?LACB. 'GTC K?QQ CVRGLARGMLQ UGRF JMQQCQ MD KMPC RF?L 50 NCPACLR MD CVR?LR QNCAGCQ ?PC M@QCPT?@JC GL RFC DMQQGJ PCAMPB. #GMBGTCPQGRW PCAMTCPW RGKCQ ?DRCP K?QQ CVRGLARGMLQ T?PW, @SR F?TC @CCL SN RM 30 KGJJGML WC?PQ. 3CACLR CVRGLARGMLQ ?PC PCAMPBCB GL UPGRRCL FGQRMPW ?LB ?PC RFC @?QGQ DMP MLC KCRFMB MD CQRGK?RGLE AMLRCKNMP?PW CVRGLARGML P?RCQ. 5FC MRFCP KCRFMB SQCQ KC?QSPCQ MD F?@GR?R JMQQ ?LB QNCAGCQ-?PC? PCJ?RGMLQFGNQ. &QRGK?RCQ MD AMLRCKNMP?PW CVRGLARGML P?RCQ T?PW, @SR QMKC P?RCQ ?PC ?Q FGEF ?Q 500 RGKCQ RFC @?AIEPMSLB P?RC, ?Q BCRCPKGLCB DPMK RFC DMQQGJ PCAMPB, ?LB ?PC NPCBGARCB RM PGQC.
47.2 /?< $DGFIK8E:< F= B@F;@M<IJ@KP KF #LD8E '@=<

)SK?LQ SQC K?LW AMKNMSLBQ RF?R UCPC DGPQR BGQAMTCPCB MP BCPGTCB DPMK JGTGLE MPE?LGQKQ ?Q KCBGAGLCQ: QCAMLB?PW NJ?LR AMKNMSLBQ, ?LGK?J RMVGLQ, ?LB ?LRG@GMRGAQ NPMBSACB @W @?ARCPG? ?LB DSLEG. .MPC KCBGAGLCQ ?PC CVNCARCB RM @C BGQAMTCPCB GL L?RSPC. -MQQ MD @GMBGTCPQGRW UGJJ GKN?AR RFC LSK@CP MD NF?PK?ACSRGA?JQ ?T?GJ?@JC RM FSK?LQ. $PMN BGTCPQGRW GQ ? PCOSGPCKCLR DMP DMMB QCASPGRW, ?LB GR GQ @CGLE JMQR. 5FC JMQQ MD UGJB PCJ?RGTCQ RM APMNQ ?JQM RFPC?RCLQ @PCCBCPQs ?@GJGRGCQ RM APC?RC LCU T?PGCRGCQ. &AMQWQRCKQ NPMTGBC CAMQWQRCK QCPTGACQ RF?R QSNNMPR FSK?L ?EPGASJRSPC: NMJJGL?RGML, LSRPGCLR AWAJGLE, NCQR AMLRPMJ, ?LB QMGJ BCTCJMNKCLR ?LB K?GLRCL?LAC. -MQQ MD @GMBGTCPQGRW RFPC?RCLQ RFCQC CAMQWQRCK QCPTGACQ ?LB PGQIQ K?IGLE DMMB NPMBSARGML KMPC CVNCLQGTC MP GKNMQQG@JC. 8GJB DMMB QMSPACQ ?PC K?GLJW ?OS?RGA, @SR DCU ?PC @CGLE K?L?ECB DMP QSQR?GL?@GJGRW. 'GQFCPGCQs ?@GJGRW RM NPMTGBC NPMRCGL RM FSK?L NMNSJ?RGMLQ GQ RFPC?RCLCB UFCL CVRGLARGML MAASPQ. #GMBGTCPQGRW K?W NPMTGBC GKNMPR?LR NQWAFMJMEGA?J @CLCDGRQ RM FSK?LQ. "BBGRGML?JJW, RFCPC ?PC KMP?J ?PESKCLRQ DMP RFC K?GLRCL?LAC MD @GMBGTCPQGRW.
47.3 /?I<8KJ KF B@F;@M<IJ@KP

5FC AMPC RFPC?RQ RM @GMBGTCPQGRW ?PC FSK?L NMNSJ?RGML EPMURF ?LB SLQSQR?GL?@JC PCQMSPAC SQC. 5M B?RC, RFC KMQR QGELGDGA?LR A?SQCQ MD CVRGLARGMLQ ?PC F?@GR?R JMQQ, GLRPMBSARGML MD CVMRGA QNCAGCQ, ?LB MTCPF?PTCQRGLE. $JGK?RC AF?LEC GQ NPCBGARCB RM @C ? QGELGDGA?LR A?SQC MD CVRGLARGMLQ GL RFC AMKGLE ACLRSPW. )?@GR?R JMQQ MAASPQ RFPMSEF BCDMPCQR?RGML, B?KKGLE MD PGTCPQ, ?LB MRFCP ?ARGTGRGCQ. 0TCPF?PTCQRGLE GQ ? RFPC?R N?PRGASJ?PJW RM ?OS?RGA QNCAGCQ, UFGJC RFC R?IGLE MD @SQF KC?R GL RFC FSKGB RPMNGAQ RFPC?RCLQ K?LW QNCAGCQ GL "QG?, "DPGA?, ?LB RFC "KCPGA?Q. &VMRGA QNCAGCQ F?TC @CCL RFC A?SQC MD ? LSK@CP MD CVRGLARGMLQ ?LB ?PC CQNCAG?JJW B?K?EGLE RM GQJ?LBQ ?LB J?ICQ. &VMRGA QNCAGCQs GLRPMBSARGMLQ ?PC GLAPC?QGLE @CA?SQC MD RFC GLAPC?QCB KM@GJGRW MD FSK?L NMNSJ?RGMLQ ?LB EPMUGLE EJM@?J RP?BC ?LB RP?LQNMPR?RGML. $JGK?RC AF?LEC GQ DMPAGLE P?LEC AF?LECQ RF?R K?W JC?B RM CVRGLARGML. *R GQ ?JQM ?DDCARGLE ?B?NR?RGMLQ RM RFC RGKGLE MD PCQMSPAC ?T?GJ?@GJGRW RF?R LCE?RGTCJW ?DDCARQ QNCAGCQ GL QC?QML?J CLTGPMLKCLRQ. 5FC GKN?ARQ MD AJGK?RC AF?LEC ?PC EPC?RCQR GL RFC ?PARGA. (JM@?J U?PKGLE UGJJ ?JQM P?GQC QC? JCTCJQ, CJGKGL?RGLE QMKC GQJ?LBQ ?LB PCBSAGLE RFC ?PC? MD ?JJ MRFCPQ.
47.4 +I<J<IM@E> B@F;@M<IJ@KP

/CU RCAFLMJMEGA?J KCRFMBQ QSAF ?Q %/" @?PAMBGLE ?LB GLDMPK?RGML NPMACQQGLE ?LB ?AACQQG@GJGRW ?PC D?AGJGR?RGLE RFC A?R?JMEGLE MD RFC NJ?LCRsQ @GMBGTCPQGRW. 5FCPC GQ ?JQM ? JCEGQJ?RGTC DP?KCUMPI DMP @GMBGTCPQGRW NPMRCARGML. *LRCPL?RGML?J RPC?RGCQ QSAF ?Q $*5&4 PCESJ?RC RFC RP?LQNMPR?RGML MD CLB?LECPCB QNCAGCQ ?APMQQ GLRCPL?RGML?J @MPBCPQ. -CEGQJ?RGML UGRFGL GLBGTGBS?J AMSLRPGCQ NPMRCARGLE QNCAGCQ ?LB ?EPCCKCLRQ ML EJM@?J U?PKGLE F?TC F?B JGKGRCB QSAACQQ; RFCPC GQ ?R NPCQCLR LM GLRCPL?RGML?J ?EPCCKCLR ML R?PECRQ DMP EPCCLFMSQC E?Q CKGQQGMLQ. *L RFC 6LGRCB 4R?RCQ, RFC &LB?LECPCB 4NCAGCQ "AR NPMRCARQ JGQRCB QNCAGCQ @SR GQ F?KNCPCB @W NPMACBSP?J BGDDGASJRGCQ ?LB ? DMASQ ML GLBGTGBS?J QNCAGCQ. 5FC .GEP?RMPW #GPB "AR GQ ?L ?EPCCKCLR @CRUCCL $?L?B? ?LB RFC 6LGRCB 4R?RCQ RM NPMRCAR KGEP?RMPW @GPBQ. 5FC LML-NPMDGR QCARMP GQ ?JQM TCPW ?ARGTC GL AMLQCPT?RGML CDDMPRQ GL ? T?PGCRW MD U?WQ. $MLQCPT?RGML NPCQCPTCQ ?PC ? K?HMP RMMJ GL @GMBGTCPQGRW NPMRCARGML. 1PCQCLRJW, 11NCPACLR MD &?PRFsQ J?LB QSPD?AC GQ NPMRCARCB GL QMKC U?W. 5FC QAGCLAC MD GQJ?LB @GMECMEP?NFW F?Q GLDMPKCB RFC MNRGK?J BCQGEL MD NPCQCPTCQ; FMUCTCP, NPCQCPTCQ F?TC JGKGR?RGMLQ GKNMQCB @W NMJGRGA?J ?LB CAMLMKGA DMPACQ. *L ?BBGRGML, AJGK?RC AF?LEC UGJJ JGKGR RFC CDDCARGTCLCQQ MD NPCQCPTCQ GL RFC DSRSPC. " BMULQGBC MD NPCQCPTCQ GQ RF?R RFCW K?W JCQQCL RFC NPCQQSPC ML FSK?L QMAGCRGCQ RM DSLARGML KMPC QSQR?GL?@JW MSRQGBC RFC NPCQCPTCQ. )?@GR?R PCQRMP?RGML F?Q RFC NMRCLRG?J RM PCQRMPC CAMQWQRCKQ RM NPCTGMSQ @GMBGTCPQGRW JCTCJQ @CDMPC QNCAGCQ @CAMKC CVRGLAR. &V?KNJCQ MD PCQRMP?RGML GLAJSBC PCGLRPMBSARGML MD ICWQRMLC QNCAGCQ ?LB PCKMT?J MD B?KQ ML PGTCPQ. ;MMQ F?TC ?RRCKNRCB RM R?IC ? KMPC ?ARGTC PMJC GL AMLQCPT?RGML ?LB A?L F?TC ? JGKGRCB PMJC GL A?NRGTC @PCCBGLE NPMEP?KQ. ;MMQ ?JQM K?W F?TC ? SQCDSJ PMJC GL CBSA?RGML.
#(!04%2 47 X #/.3%26!4)/. ")/,/'9 !.$ ")/$)6%23)49
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A-/ C*)) C/$*) ,0 ./$*).

1. Figure 47.6 4AGCLRGQRQ KC?QSPCB RFC PCJ?RGTC ?@SLB?LAC MD DCPL QNMPCQ ?@MTC ?LB @CJMU RFC ,1E @MSLB?PW GL RFGQ PMAI Q?KNJC. 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ KMQR JGICJW PCNPCQCLRQ RFCGP DGLBGLEQ >. "L ?@SLB?LAC MD DCPL QNMPCQ DPMK QCTCP?J QNCAGCQ U?Q DMSLB @CJMU RFC ,1E @MSLB?PW, @SR LMLC U?Q DMSLB ?@MTC. ?. "L ?@SLB?LAC MD DCPL QNMPCQ DPMK QCTCP?J QNCAGCQ U?Q DMSLB ?@MTC RFC ,1E @MSLB?PW, @SR LMLC U?Q DMSLB @CJMU. @. "L ?@SLB?LAC MD DCPL QNMPCQ U?Q DMSLB @MRF ?@MTC ?LB @CJMU RFC ,-1E @MSLB?PW, @SR MLJW MLC QNCAGCQ U?Q DMSLB @CJMU RFC @MSLB?PW , ?LB K?LW QNCAGCQ UCPC DMSLB ?@MTC RFC @MSLB?PW. A. .?LW QNCAGCQ MD DCPL QNMPCQ UCPC DMSLB @MRF ?@MTC ?LB @CJMU RFC @MSLB?PW, @SR RFC RMR?J LSK@CP MD QNMPCQ U?Q EPC?RCP @CJMU RFC @MSLB?PW. 2. Figure 47.9 5FC 4T?J@?PB (JM@?J 4CCB 7?SJR GQ JMA?RCB ML 4NGRQ@CPECL GQJ?LB GL /MPU?W, UFGAF F?Q ?L ?PARGA AJGK?RC. 8FW KGEFR ?L ?PARGA AJGK?RC @C EMMB DMP QCCB QRMP?EC 3. $MLTCPRGLE ? NP?GPGC RM ? D?PK DGCJB GQ ?L CV?KNJC MD >>>>>>>>. >. MTCPF?PTCQRGLE ?. F?@GR?R JMQQ @. CVMRGA QNCAGCQ A. AJGK?RC AF?LEC 4. Figure 47.16 8FGAF MD RFC DMJJMUGLE QR?RCKCLRQ GQ LMR QSNNMPRCB @W RFGQ EP?NF >. 5FCPC ?PC KMPC TSJLCP?@JC DGQFCQ RF?L APGRGA?JJW CLB?LECPCB ?LB CLB?LECPCB DGQFCQ AMK@GLCB. ?. 5FCPC ?PC KMPC APGRGA?JJW CLB?LECPCB ?KNFG@G?LQ RF?L TSJLCP?@JC, CLB?LECPCB ?LB APGRGA?JJW CLB?LECPCB PCNRGJCQ AMK@GLCB. @. 8GRFGL C?AF EPMSN, RFCPC ?PC KMPC APGRGA?JJW CLB?LECPCB QNCAGCQ RF?L TSJLCP?@JC QNCAGCQ. A. " EPC?RCP NCPACLR?EC MD @GPB QNCAGCQ ?PC APGRGA?JJW CLB?LECPCB RF?L KMJJSQI QNCAGCQ.
- 1$ 2 ,0 ./$*).
5. 8GRF ?L CVRGLARGML P?RC MD 100 &/.4: ?LB ?L CQRGK?RCB 10 KGJJGML QNCAGCQ, FMU K?LW CVRGLARGMLQ ?PC CVNCARCB RM MAASP GL ? ACLRSPW >. ?. @. A. 100 10, 000 100, 000 1, 000, 000 A. ? NCQR MD ? APMN NCQR 10. 1MJJGL?RGML GQ ?L CV?KNJC MD >>>>>>>>. >. ?. @. A. ? NMQQG@JC QMSPAC MD LCU BPSEQ AFCKGA?J BGTCPQGRW ?L CAMQWQRCK QCPTGAC APMN NCQR AMLRPMJ
6. "L ?B?NRGTC P?BG?RGML GQ>>>>>>>>. >. ? @SPQR MD QNCAG?RGML ?. ? FC?JRFW JCTCJ MD 67 P?BG?RGML @. ? FWNMRFCQGXCB A?SQC MD ? K?QQ CVRGLARGML A. CTGBCLAC MD ?L ?QRCPMGB GKN?AR 7. 5FC LSK@CP MD ASPPCLRJW BCQAPG@CB QNCAGCQ ML RFC NJ?LCR GQ ?@MSR >>>>>>>>. >. 17,000 ?. 150,000 @. 1.5 KGJJGML A. 10 KGJJGML 8. " K?QQ CVRGLARGML GQ BCDGLCB ?Q >>>>>>>>. >. ?. @. A. ? JMQQ MD 95 NCPACLR MD QNCAGCQ ?L ?QRCPMGB GKN?AR ? @MSLB?PW @CRUCCL ECMJMEGA?J NCPGMBQ ? JMQQ MD 50 NCPACLR MD QNCAGCQ
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
9. " QCAMLB?PW NJ?LR AMKNMSLB KGEFR @C SQCB DMP UFGAF MD RFC DMJJMUGLE >. ? LCU APMN T?PGCRW ?. ? LCU BPSE @. ? QMGJ LSRPGCLR
1426
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L?KC MD RFC JCEGQJ?RGML RF?R K?ICQ RFGQ GJJCE?J >. ?. @. A. 3CB -GQR .GEP?RMPW #GPB "AR $*5&4 &LB?LECPCB 4NCAGCQ "AR (&4")
@. *LRCPL?RGML?J 6LGML DMP RFC $MLQCPT?RGML MD /?RSPC (*6$/) A. ,WMRM 1PMRMAMJ 16. "@MSR UF?R NCPACLR?EC MD J?LB ML RFC NJ?LCR GQ QCR ?QGBC ?Q ? NPCQCPTC MD QMKC RWNC >. 1 NCPACLR ?. 6 NCPACLR @. 11 NCPACLR A. 15 NCPACLR
15. 8F?R U?Q RFC L?KC MD RFC DGPQR GLRCPL?RGML?J ?EPCCKCLR ML AJGK?RC AF?LEC >. 3CB -GQR ?. .MLRPC?J 1PMRMAMJ
C-$/$CA' /#$)&$)" ,0 ./$*).

17. %CQAPG@C RFC CTGBCLAC DMP RFC A?SQC MD RFC $PCR?ACMSQp1?JCMECLC (,p1E) K?QQ CVRGLARGML. 18. %CQAPG@C RFC RUM KCRFMBQ SQCB RM A?JASJ?RC AMLRCKNMP?PW CVRGLARGML P?RCQ. 19. &VNJ?GL FMU @GMBGTCPQGRW JMQQ A?L GKN?AR APMN BGTCPQGRW. 20. %CQAPG@C RUM RWNCQ MD AMKNMSLBQ DPMK JGTGLE RFGLEQ RF?R ?PC SQCB ?Q KCBGA?RGMLQ. 21. %CQAPG@C RFC KCAF?LGQKQ @W UFGAF FSK?L NMNSJ?RGML EPMURF ?LB PCQMSPAC SQC A?SQCQ GLAPC?QCB CVRGLARGML P?RCQ. 22. &VNJ?GL UF?R CVRGLARGML RFPC?RQ ? DPME JGTGLE ML ? KMSLR?GLQGBC GL $MQR? 3GA? KGEFR D?AC. 23. %CQAPG@C RUM AMLQGBCP?RGMLQ GL AMLQCPT?RGML NPCQCPTC BCQGEL. 24. %CQAPG@C UF?R F?NNCLQ RM ?L CAMQWQRCK UFCL ? ICWQRMLC QNCAGCQ GQ PCKMTCB.
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1427
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A1 R +<I@F;@: /89C< F= K?< C<D<EKJ
!@>LI< A1
1428
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A2 R "<FCF>@:8C /@D< CCF:B
!@>LI< A2
A3 R "<FCF>@:8C /@D< C?8IK
!@>LI< A3 (@OBAFQ: 2F@E>OA 3. -ROMEV, *O.)
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A4 R (<8JLI<D<EKJ 8E; K?< (<KI@: .PJK<D

(<8JLI<D<EKJ 8E; K?< (<KI@: .PJK<D
(<8JLI<D<EK 0E@K A99I<M@8K@FE (<KI@: HL@M8C<EK
1 KJ = 10}9 J 1 \J = 10}6 J 1 JJ = 0.001 J 1 @J = 0.01 J 1 J = 100 @J 1 J = 1000 JJ 1 HJ = 1000 J 1 \D = 10}6 D 1 JD = 10}3 D 1 D = 1000 JD 1 HD = 1000 D 1 \I = 10}6 I 1 JI = 10}3 I 1 I = 1000 JI 1 HI = 1000 I 1 @J2 = 100 JJ2 1 J2 = 10,000 @J2 1 E> = 10,000 J2 u 1 @J2 = 0.155 PNR>OB FK@E 1 J2 = 10.764 PNR>OB CBBQ 1 J2 = 1.196 PNR>OB V>OAP 1 E> = 2.471 >@OBP 1[# = 5/9 ^ ([& - 32) 1 JI = 0.034 CIRFA LRK@B 1 I = 1.057 NR>OQP 1 HI = 264.172 D>IILKP 1 D = 0.035 LRK@B 1 HD = 2.205 MLRKAP 1 JJ = 0.039 FK@E 1 @J = 0.394 FK@E 1 J = 39.37 FK@EBP 1 J = 3.28 CBBQ 1 J = 1.093 V>OAP 1 HJ = 0.621 JFIBP
AGGIFO@D8K< .K8E;8I; HL@M8C<EK
K>KLJBQBO KJ JF@OLJBQBO \J JFIIFJBQBO ,BKDQE JJ
@BKQFJBQBO @J JBQBO HFILJBQBO J HJ
JF@OLDO>J \D ->PP JFIIFDO>J DO>J HFILDO>J JF@OLIFQBO 6LIRJB JFIIFIFQBO IFQBO HFILIFQBO JD D HD \I JI I HI
PNR>OB 2 @BKQFJBQBO @J !OB> PNR>OB JBQBO EB@Q>OB 4BJMBO>QROB /89C< A1 #BIPFRP J2 E> [#
1430
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1431
A).2 - & 4
Chapter 1
1 Figure 1.6 1: $; 2: '; 3: "; 4: #; 5: %; 6: &. 5FC MPGEGL?J FWNMRFCQGQ GQ GLAMPPCAR, ?Q RFC AMDDCCK?ICP UMPIQ UFCL NJSEECB GLRM RFC MSRJCR. "JRCPL?RGTC FWNMRFCQCQ GLAJSBC RF?R RFC AMDDCC K?ICP KGEFR @C @PMICL MP RF?R RFC AMDDCC K?ICP U?QLsR RSPLCB ML. 2 Figure 1.7 1: GLBSARGTC; 2: BCBSARGTC; 3: BCBSARGTC; 4: GLBSARGTC. 3 Figure 1.16 $MKKSLGRGCQ CVGQR UGRFGL NMNSJ?RGMLQ UFGAF CVGQR UGRFGL CAMQWQRCKQ. 4 # 5 " 6 % 7 % 8 $ 9 " 10 $ 11 " 12 # 13 $ 14 % 15 % 16 "LQUCPQ UGJJ T?PW, @SR QFMSJB ?NNJW RFC QRCNQ MD RFC QAGCLRGDGA KCRFMB. 0LC NMQQG@GJGRW AMSJB @C ? A?P UFGAF BMCQLsR QR?PR. 5FC FWNMRFCQGQ AMSJB @C RF?R RFC A?P BMCQLs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s F?@GR?R, ?LB FMU NJ?LRQ PCQNMLB RM RFCGP CLTGPMLKCLRQ. 5MNGAQ RF?R D?JJ MSRQGBC MD @GMJMEW (RFC tQRSBW MD JGDCu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sQ AMKKSLGRW, RFC AMKKSLGRWsQ CAMQWQRCK, ?LB RFC CAMQWQRCKsQ N?PR GL RFC @GMQNFCPC. 8FCL QRSBWGLE ?L GLBGTGBS?J MPE?LGQK, ? @GMJMEGQR AMSJB CV?KGLC RFC ACJJ ?LB GRQ MPE?LCJJCQ, RFC RGQQSCQ RF?R RFC ACJJQ K?IC SN, RFC MPE?LQ ?LB RFCGP PCQNCARGTC MPE?L QWQRCKQ, ?LB RFC QSK RMR?JqRFC MPE?LGQK GRQCJD.
Chapter 2
1 Figure 2.3 $?P@ML-12 F?Q QGV LCSRPMLQ. $?P@ML-13 F?Q QCTCL LCSRPMLQ. 2 Figure 2.7 &JCKCLRQ GL EPMSN 1 LCCB RM JMQC MLC CJCARPML RM ?AFGCTC ? QR?@JC CJCARPML AMLDGESP?RGML. &JCKCLRQ GL EPMSNQ 14 ?LB 17 LCCB RM E?GL DMSP ?LB MLC CJCARPMLQ, PCQNCARGTCJW, RM ?AFGCTC ? QR?@JC AMLDGESP?RGML. 3 Figure 2.24 $ 4 " 5 % 6 $ 7 " 8 % 9 " 10 $ 11 # 12 % 13 " 14 *MLGA @MLBQ ?PC APC?RCB @CRUCCL GMLQ. 5FC CJCARPMLQ ?PC LMR QF?PCB @CRUCCL RFC ?RMKQ, @SR P?RFCP ?PC ?QQMAG?RCB KMPC UGRF MLC GML RF?L RFC MRFCP. *MLGA @MLBQ ?PC QRPMLE @MLBQ, @SR ?PC UC?ICP RF?L AMT?JCLR @MLBQ, KC?LGLE GR R?ICQ JCQQ CLCPEW RM @PC?I ?L GMLGA @MLB AMKN?PCB UGRF ? AMT?JCLR MLC. 15 )WBPMECL @MLBQ ?LB T?L BCP 8??JQ GLRCP?ARGMLQ DMPK UC?I ?QQMAG?RGMLQ @CRUCCL BGDDCPCLR KMJCASJCQ MP UGRFGL BGDDCPCLR PCEGMLQ MD RFC Q?KC KMJCASJC. 5FCW NPMTGBC RFC QRPSARSPC ?LB QF?NC LCACQQ?PW DMP NPMRCGLQ ?LB %/" UGRFGL ACJJQ QM RF?R RFCW DSLARGML NPMNCPJW. 16 #SDDCPQ ?@QMP@ RFC DPCC FWBPMECL GMLQ ?LB FWBPMVGBC GMLQ RF?R PCQSJR DPMK AFCKGA?J PC?ARGMLQ. #CA?SQC RFCW A?L @MLB RFCQC GMLQ, RFCW NPCTCLR GLAPC?QCQ MP BCAPC?QCQ GL N). "L CV?KNJC MD ? @SDDCP QWQRCK GQ RFC @GA?P@ML?RC QWQRCK GL RFC FSK?L @MBW. 5FGQ QWQRCK GQ ?@JC RM ?@QMP@ FWBPMECL ?LB FWBPMVGBC GMLQ RM NPCTCLR AF?LECQ GL N) ?LB ICCN ACJJQ DSLARGMLGLE NPMNCPJW. 17 4MKC GLQCARQ A?L U?JI ML U?RCP, ?JRFMSEF RFCW ?PC FC?TGCP (BCLQCP) RF?L U?RCP, @CA?SQC MD RFC QSPD?AC RCLQGML MD U?RCP. 4SPD?AC RCLQGML PCQSJRQ DPMK AMFCQGML, MP RFC ?RRP?ARGML @CRUCCL U?RCP KMJCASJCQ ?R RFC QSPD?AC MD RFC @MBW MD U?RCP (RFC JGOSGB-?GP/E?Q GLRCPD?AC). 18 $?P@ML GQ SLGOSC ?LB DMSLB GL ?JJ JGTGLE RFGLEQ @CA?SQC GR A?L DMPK SN RM DMSP AMT?JCLR @MLBQ @CRUCCL ?RMKQ MP KMJCASJCQ. 5FCQC A?L @C LMLNMJ?P MP NMJ?P AMT?JCLR @MLBQ, ?LB RFCW ?JJMU DMP RFC DMPK?RGML MD JMLE AF?GLQ MD A?P@ML KMJCASJCQ RF?R AMK@GLC RM DMPK NPMRCGLQ ?LB %/". 19 4?RSP?RCB RPGEJWACPGBCQ AMLR?GL LM BMS@JC @MLBQ @CRUCCL A?P@ML ?RMKQ; RFCW ?PC SQS?JJW QMJGB ?R PMMK RCKNCP?RSPC. 6LQ?RSP?RCB RPGEJWACPGBCQ AMLR?GL ?R JC?QR MLC BMS@JC @MLB @CRUCCL A?P@ML ?RMKQ ?LB ?PC SQS?JJW JGOSGB ?R PMMK RCKNCP?RSPC.
Chapter 3
1 Figure 3.5 (JSAMQC ?LB E?J?ARMQC ?PC ?JBMQCQ. 'PSARMQC GQ ? ICRMQC. 2 Figure 3.23 1MJ?P ?LB AF?PECB ?KGLM ?AGB PCQGBSCQ (RFC PCK?GLBCP ?DRCP NCNRGBC @MLB DMPK?RGML) ?PC KMPC JGICJW RM @C DMSLB ML RFC QSPD?AC MD QMJS@JC NPMRCGLQ UFCPC RFCW A?L GLRCP?AR UGRF U?RCP, ?LB LMLNMJ?P (C.E., ?KGLM ?AGB QGBC AF?GLQ) ?PC KMPC JGICJW RM @C DMSLB GL RFC GLRCPGMP UFCPC RFCW ?PC QCOSCQRCPCB DPMK U?RCP. *L KCK@P?LC NPMRCGLQ, LMLNMJ?P ?LB FWBPMNFM@GA ?KGLM ?AGB QGBC AF?GLQ ?QQMAG?RC UGRF RFC FWBPMNFM@GA R?GJQ MD NFMQNFMJGNGBQ, UFGJC NMJ?P ?LB AF?PECB ?KGLM ?AGB QGBC AF?GLQ GLRCP?AR UGRF RFC NMJ?P FC?B EPMSNQ MP UGRF RFC ?OSCMSQ QMJSRGML. )MUCTCP,
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RFCPC ?PC CVACNRGMLQ. 4MKCRGKCQ, NMQGRGTCJW ?LB LCE?RGTCJW AF?PECB ?KGLM ?AGB QGBC AF?GLQ GLRCP?AR UGRF MLC ?LMRFCP GL RFC GLRCPGMP MD ? NPMRCGL, ?LB NMJ?P MP AF?PECB ?KGLM ?AGB QGBC AF?GLQ RF?R GLRCP?AR UGRF ? JGE?LB A?L @C DMSLB GL RFC JGE?LB @GLBGLE NMAICR. 3 Figure 3.33 "BCLGLC GQ J?PECP RF?L AWRMQGLC ?LB UGJJ LMR @C ?@JC RM @?QC N?GP NPMNCPJW UGRF RFC ES?LGLC ML RFC MNNMQGLE QRP?LB. 5FGQ UGJJ A?SQC RFC %/" RM @SJEC. %/" PCN?GP CLXWKCQ K?W PCAMELGXC RFC @SJEC ?LB PCNJ?AC RFC GLAMPPCAR LSAJCMRGBC. 4 # 5 " 6 % 7 % 8 # 9 # 10 % 11 " 12 $ 13 # 14 $ 15 % 16 #GMJMEGA?J K?APMKMJCASJCQ ?PC MPE?LGA @CA?SQC RFCW AMLR?GL A?P@ML. 17 *L ? BCFWBP?RGML QWLRFCQGQ PC?ARGML, RFC FWBPMECL MD MLC KMLMKCP AMK@GLCQ UGRF RFC FWBPMVWJ EPMSN MD ?LMRFCP KMLMKCP, PCJC?QGLE ? KMJCASJC MD U?RCP. 5FGQ APC?RCQ ?L MNCLGLE GL RFC MSRCP QFCJJQ MD ?RMKQ GL RFC KMLMKCPQ, UFGAF A?L QF?PC CJCARPMLQ ?LB DMPK AMT?JCLR @MLBQ. 18 (JWAMECL ?LB QR?PAF ?PC NMJWQ?AAF?PGBCQ. 5FCW ?PC RFC QRMP?EC DMPK MD EJSAMQC. (JWAMECL GQ QRMPCB GL ?LGK?JQ GL RFC JGTCP ?LB GL KSQAJC ACJJQ, UFCPC?Q QR?PAF GQ QRMPCB GL RFC PMMRQ, QCCBQ, ?LB JC?TCQ MD NJ?LRQ. 4R?PAF F?Q RUM BGDDCPCLR DMPKQ, MLC SL@P?LAFCB (?KWJMQC) ?LB MLC @P?LAFCB (?KWJMNCARGL), UFCPC?Q EJWAMECL GQ ? QGLEJC RWNC MD ? FGEFJW @P?LAFCB KMJCASJC. 19 5FC D 1-4 EJWAMQGBGA JGLI?EC GL ACJJSJMQC A?LLMR @C @PMICL BMUL @W FSK?L BGECQRGTC CLXWKCQ. )CP@GTMPCQ QSAF ?Q AMUQ, @SDD?JMQ, ?LB FMPQCQ ?PC ?@JC RM BGECQR EP?QQ RF?R GQ PGAF GL ACJJSJMQC ?LB SQC GR ?Q ? DMMB QMSPAC @CA?SQC @?ARCPG? ?LB NPMRGQRQ GL RFCGP BGECQRGTC QWQRCKQ, CQNCAG?JJW GL RFC PSKCL, QCAPCRC RFC CLXWKC ACJJSJ?QC. $CJJSJ?QCQ A?L @PC?I BMUL ACJJSJMQC GLRM EJSAMQC KMLMKCPQ RF?R A?L @C SQCB ?Q ?L CLCPEW QMSPAC @W RFC ?LGK?J. 20 '?R QCPTCQ ?Q ? T?JS?@JC U?W DMP ?LGK?JQ RM QRMPC CLCPEW. *R A?L ?JQM NPMTGBC GLQSJ?RGML. 8?VCQ A?L NPMRCAR NJ?LR JC?TCQ ?LB K?KK?JG?L DSP DPMK ECRRGLE UCR. 1FMQNFMJGNGBQ ?LB QRCPMGBQ ?PC GKNMPR?LR AMKNMLCLRQ MD ?LGK?J ACJJ KCK@P?LCQ, ?Q UCJJ ?Q NJ?LR, DSLE?J, ?LB @?ARCPG?J KCK@P?LCQ. 21 5P?LQ D?RQ ?PC APC?RCB ?PRGDGAG?JJW UFCL FWBPMECL E?Q GQ @S@@JCB RFPMSEF MGJQ RM QMJGBGDW RFCK. 5FC BMS@JC @MLBQ MD RFC *0: AMLDMPK?RGML GL RFC FWBPMA?P@ML AF?GL K?W @C AMLTCPRCB RM BMS@JC @MLBQ GL RFC ;9(5: AMLDGESP?RGML. 4MKC PCQR?SP?LRQ ?PC @?LLGLE RP?LQ D?RQ @CA?SQC RFCW A?SQC FGEFCP JCTCJQ MD -%-, MP t@?BuAFMJCQRCPMJ. 22 " AF?LEC GL ECLC QCOSCLAC A?L JC?B RM ? BGDDCPCLR ?KGLM ?AGB @CGLE ?BBCB RM ? NMJWNCNRGBC AF?GL GLQRC?B MD RFC LMPK?J MLC. 5FGQ A?SQCQ ? AF?LEC GL NPMRCGL QRPSARSPC ?LB DSLARGML. 'MP CV?KNJC, GL QGAIJC ACJJ ?LCKG?, RFC FCKMEJM@GL D AF?GL F?Q ? QGLEJC ?KGLM ?AGB QS@QRGRSRGMLqRFC ?KGLM ?AGB EJSR?KGA ?AGB GL NMQGRGML QGV GQ QS@QRGRSRCB @W T?JGLC. #CA?SQC MD RFGQ AF?LEC, FCKMEJM@GL KMJCASJCQ DMPK ?EEPCE?RCQ, ?LB RFC BGQA-QF?NCB PCB @JMMB ACJJQ ?QQSKC ? APCQACLR QF?NC, UFGAF PCQSJRQ GL QCPGMSQ FC?JRF NPM@JCKQ. 23 5FC QCOSCLAC ?LB LSK@CP MD ?KGLM ?AGBQ GL ? NMJWNCNRGBC AF?GL GQ GRQ NPGK?PW QRPSARSPC. 5FC JMA?J DMJBGLE MD RFC NMJWNCNRGBC GL QMKC PCEGMLQ GQ RFC QCAMLB?PW QRPSARSPC MD RFC NPMRCGL. 5FC RFPCC-BGKCLQGML?J QRPSARSPC MD ? NMJWNCNRGBC GQ ILMUL ?Q GRQ RCPRG?PW QRPSARSPC, APC?RCB GL N?PR @W AFCKGA?J GLRCP?ARGMLQ QSAF ?Q FWBPMECL @MLBQ @CRUCCL NMJ?P QGBC AF?GLQ, T?L BCP 8??JQ GLRCP?ARGMLQ, BGQSJDGBC JGLI?ECQ, ?LB FWBPMNFM@GA GLRCP?ARGMLQ. 4MKC NPMRCGLQ ?PC DMPKCB DPMK KSJRGNJC NMJWNCNRGBCQ, ?JQM ILMUL ?Q QS@SLGRQ, ?LB RFC GLRCP?ARGML MD RFCQC QS@SLGRQ DMPKQ RFC OS?RCPL?PW QRPSARSPC. 24 %/" F?Q ? BMS@JC-FCJGV QRPSARSPC. 5FC QSE?P ?LB RFC NFMQNF?RC ?PC ML RFC MSRQGBC MD RFC FCJGV ?LB RFC LGRPMECLMSQ @?QCQ ?PC GL RFC GLRCPGMP. 5FC KMLMKCPQ MD %/" ?PC LSAJCMRGBCQ AMLR?GLGLE BCMVWPG@MQC, MLC MD RFC DMSP LGRPMECLMSQ @?QCQ (", 5, ( ?LB $), ?LB ? NFMQNF?RC EPMSN. 3/" GQ SQS?JJW QGLEJC-QRP?LBCB ?LB GQ K?BC MD PG@MLSAJCMRGBCQ RF?R ?PC JGLICB @W NFMQNFMBGCQRCP JGLI?ECQ. " PG@MLSAJCMRGBC AMLR?GLQ PG@MQC (RFC NCLRMQC QSE?P), MLC MD RFC DMSP LGRPMECLMSQ @?QCQ (",6, (, ?LB $), ?LB RFC NFMQNF?RC EPMSN. 25 5FC DMSP RWNCQ MD 3/" ?PC KCQQCLECP 3/", PG@MQMK?J 3/", RP?LQDCP 3/", ?LB KGAPM3/". .CQQCLECP 3/" A?PPGCQ RFC GLDMPK?RGML DPMK RFC %/" RF?R AMLRPMJQ ?JJ ACJJSJ?P ?ARGTGRGCQ. 5FC K3/" @GLBQ RM RFC PG@MQMKCQ RF?R ?PC AMLQRPSARCB MD NPMRCGLQ ?LB P3/", ?LB R3/" RP?LQDCPQ RFC AMPPCAR ?KGLM ?AGB RM RFC QGRC MD NPMRCGL QWLRFCQGQ. KGAPM3/" PCESJ?RCQ RFC ?T?GJ?@GJGRW MD K3/" DMP RP?LQJ?RGML.
Chapter 4
1 Figure 4.7 4S@QR?LACQ A?L BGDDSQC KMPC OSGAIJW RFPMSEF QK?JJ ACJJQ. 4K?JJ ACJJQ F?TC LM LCCB DMP MPE?LCJJCQ ?LB RFCPCDMPC BM LMR LCCB RM CVNCLB CLCPEW ECRRGLE QS@QR?LACQ ?APMQQ MPE?LCJJC KCK@P?LCQ. -?PEC ACJJQ F?TC MPE?LCJJCQ RF?R A?L QCN?P?RC ACJJSJ?P NPMACQQCQ, CL?@JGLE RFCK RM @SGJB KMJCASJCQ RF?R ?PC KMPC AMKNJCV. 2 Figure 4.8 'PCC PG@MQMKCQ ?LB PMSEF CLBMNJ?QKGA PCRGASJSK (UFGAF AMLR?GLQ PG@MQMKCQ) UMSJB LMR @C ?@JC RM DMPK. 3 Figure 4.18 *R UMSJB CLB SN ML RFC MSRQGBC. "DRCP RFC TCQGAJC N?QQCQ RFPMSEF RFC (MJEG ?NN?P?RSQ ?LB DSQCQ UGRF RFC NJ?QK? KCK@P?LC, GR RSPLQ GLQGBC MSR. 4 $ 5 # 6 % 7 " 8 % 9 # 10 " 11 % 12 " 13 # 14 $ 15 $ 16 # 17 % 18 $ 19 $ 20 " JGEFR KGAPMQAMNC UMSJB @C GBC?J UFCL TGCUGLE ? QK?JJ JGTGLE MPE?LGQK, CQNCAG?JJW UFCL RFC ACJJ F?Q @CCL QR?GLCB RM PCTC?J BCR?GJQ. 21 " QA?LLGLE CJCARPML KGAPMQAMNC UMSJB @C GBC?J UFCL WMS U?LR RM TGCU RFC KGLSRC BCR?GJQ MD ? ACJJsQ QSPD?AC, @CA?SQC GRQ @C?K MD CJCARPMLQ KMTCQ @?AI ?LB DMPRF MTCP RFC QSPD?AC RM AMLTCW RFC GK?EC. 22 " RP?LQKGQQGML CJCARPML KGAPMQAMNC UMSJB @C GBC?J DMP TGCUGLE RFC ACJJsQ GLRCPL?J QRPSARSPCQ, @CA?SQC K?LW MD RFC GLRCPL?J QRPSARSPCQ F?TC KCK@P?LCQ RF?R ?PC LMR TGQG@JC @W RFC JGEFR KGAPMQAMNC. 23 5FC ?BT?LR?ECQ MD JGEFR KGAPMQAMNCQ ?PC RF?R RFCW ?PC C?QGJW M@R?GLCB, ?LB RFC JGEFR @C?K BMCQ LMR IGJJ RFC ACJJQ. )MUCTCP, RWNGA?J JGEFR KGAPMQAMNCQ ?PC QMKCUF?R JGKGRCB GL RFC ?KMSLR MD BCR?GJ RFCW A?L PCTC?J. &JCARPML KGAPMQAMNCQ ?PC GBC?J @CA?SQC WMS A?L TGCU GLRPGA?RC BCR?GJQ, @SR RFCW ?PC @SJIW ?LB AMQRJW, ?LB NPCN?P?RGML DMP RFC KGAPMQAMNGA CV?KGL?RGML IGJJQ RFC QNCAGKCL. 24 5FC ACJJ U?JJ UMSJB @C R?PECRCB @W ?LRG@GMRGAQ ?Q UCJJ ?Q RFC @?ARCPG?sQ ?@GJGRW RM PCNJGA?RC. 5FGQ UMSJB GLFG@GR RFC @?ARCPG?sQ ?@GJGRW RM PCNPMBSAC, ?LB GR UMSJB AMKNPMKGQC GRQ BCDCLQC KCAF?LGQKQ. 25 4MKC KGAPM@CQ ?PC @CLCDGAG?J. 'MP GLQR?LAC, E. *630 @?ARCPG? NMNSJ?RC RFC FSK?L ESR ?LB FCJN @PC?I BMUL DG@CP GL RFC BGCR. 4MKC DMMBQ QSAF ?Q WMESPR ?PC DMPKCB @W @?ARCPG?. 26 3G@MQMKCQ ?PC ?@SLB?LR GL KSQAJC ACJJQ ?Q UCJJ @CA?SQC KSQAJC ACJJQ ?PC AMLQRPSARCB MD RFC NPMRCGLQ K?BC @W RFC PG@MQMKCQ. 27 #MRF ?PC QGKGJ?P GL RF?R RFCW ?PC CLTCJMNCB GL ? BMS@JC KCK@P?LC, @MRF F?TC ?L GLRCPKCK@P?LC QN?AC, ?LB @MRF K?IC "51. #MRF KGRMAFMLBPG? ?LB AFJMPMNJ?QRQ F?TC %/", ?LB KGRMAFMLBPG? F?TC GLLCP DMJBQ A?JJCB APGQR?C ?LB ? K?RPGV,
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1433
UFGJC AFJMPMNJ?QRQ F?TC AFJMPMNFWJJ ?LB ?AACQQMPW NGEKCLRQ GL RFC RFWJ?IMGBQ RF?R DMPK QR?AIQ (EP?L?) ?LB ? QRPMK?. 28 t'MPK DMJJMUQ DSLARGMLu PCDCPQ RM RFC GBC? RF?R RFC DSLARGML MD ? @MBW N?PR BGAR?RCQ RFC DMPK MD RF?R @MBW N?PR. "Q ?L CV?KNJC, AMKN?PC WMSP ?PK RM ? @?RsQ UGLE. 8FGJC RFC @MLCQ MD RFC RUM AMPPCQNMLB, RFC N?PRQ QCPTC BGDDCPCLR DSLARGMLQ GL C?AF MPE?LGQK ?LB RFCGP DMPKQ F?TC ?B?NRCB RM DMJJMU RF?R DSLARGML. 29 4GLAC RFC CVRCPL?J QSPD?AC MD RFC LSAJC?P KCK@P?LC GQ AMLRGLSMSQ UGRF RFC PMSEF CLBMNJ?QKGA PCRGASJSK, UFGAF GQ N?PR MD RFC CLBMKCK@P?LC QWQRCK, RFCL GR GQ AMPPCAR RM Q?W RF?R GR GQ N?PR MD RFC QWQRCK. 30 $CLRPGMJCQ ?LB DJ?ECJJ? ?PC ?JGIC GL RF?R RFCW ?PC K?BC SN MD KGAPMRS@SJCQ. *L ACLRPGMJCQ, RUM PGLEQ MD LGLC KGAPMRS@SJC tRPGNJCRQu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
Chapter 5
1 Figure 5.12 /M, GR KSQR F?TC @CCL FWNMRMLGA ?Q ? FWNMRMLGA QMJSRGML UMSJB A?SQC U?RCP RM CLRCP RFC ACJJQ, RFCPC@W K?IGLE RFCK @SPQR. 2 Figure 5.16 $CJJQ RWNGA?JJW F?TC ? FGEF AMLACLRP?RGML MD NMR?QQGSK GL RFC AWRMNJ?QK ?LB ?PC @?RFCB GL ? FGEF AMLACLRP?RGML MD QMBGSK. *LHCARGML MD NMR?QQGSK BGQQGN?RCQ RFGQ CJCARPMAFCKGA?J EP?BGCLR. *L FC?PR KSQAJC, RFC QMBGSK/NMR?QQGSK NMRCLRG?J GQ PCQNMLQG@JC DMP RP?LQKGRRGLE RFC QGEL?J RF?R A?SQCQ RFC KSQAJC RM AMLRP?AR. 8FCL RFGQ NMRCLRG?J GQ BGQQGN?RCB, RFC QGEL?J A?LsR @C RP?LQKGRRCB, ?LB RFC FC?PR QRMNQ @C?RGLE. 1MR?QQGSK GLHCARGMLQ ?PC ?JQM SQCB RM QRMN RFC FC?PR DPMK @C?RGLE BSPGLE QSPECPW. 3 Figure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tFC?Bu MD RFC KMJCASJC A?L. 5FSQ, RFC FC?B MPGCLRQ RM U?RCP, ?LB RFC R?GJ RM MRFCP JGNGBQ. 18 )C?TW KMJCASJCQ KMTC KMPC QJMUJW RF?L JGEFRCP MLCQ. *R R?ICQ KMPC CLCPEW GL RFC KCBGSK RM KMTC RFCK ?JMLE. *LAPC?QGLE MP BCAPC?QGLE RCKNCP?RSPC GLAPC?QCQ MP BCAPC?QCQ RFC CLCPEW GL RFC KCBGSK, ?DDCARGLE KMJCASJ?P KMTCKCLR. 5FC BCLQCP ? QMJSRGML GQ, RFC F?PBCP GR GQ DMP KMJCASJCQ RM KMTC RFPMSEF GR, A?SQGLE BGDDSQGML RM QJMU BMUL BSC RM DPGARGML. -GTGLE ACJJQ PCOSGPC ? QRC?BW QSNNJW MD LSRPGCLRQ ?LB ? QRC?BW P?RC MD U?QRC PCKMT?J. *D RFC BGQR?LAC RFCQC QS@QR?LACQ LCCB RM RP?TCJ GQ RMM EPC?R, BGDDSQGML A?LLMR KMTC LSRPGCLRQ ?LB U?QRC K?RCPG?JQ CDDGAGCLRJW RM QSQR?GL JGDC. 19 8?RCP KMTCQ RFPMSEF ? KCK@P?LC GL MQKMQGQ @CA?SQC RFCPC GQ ? AMLACLRP?RGML EP?BGCLR ?APMQQ RFC KCK@P?LC MD QMJSRC ?LB QMJTCLR. 5FC QMJSRC A?LLMR CDDCARGTCJW KMTC RM @?J?LAC RFC AMLACLRP?RGML ML @MRF QGBCQ MD RFC KCK@P?LC, QM U?RCP KMTCQ RM ?AFGCTC RFGQ @?J?LAC. 20 *LHCARGML MD GQMRMLGA QMJSRGMLQ CLQSPCQ RF?R RFCPC UGJJ @C LM NCPRSP@?RGML MD RFC MQKMRGA @?J?LAC, ?LB LM U?RCP R?ICL DPMK RGQQSCQ MP ?BBCB RM RFCK DPMK RFC @JMMB. 21 5FC ACJJ F?PTCQRQ CLCPEW DPMK "51 NPMBSACB @W GRQ MUL KCR?@MJGQK RM NMUCP ?ARGTC RP?LQNMPR NPMACQQCQ, QSAF ?Q RFC ?ARGTGRW MD NSKNQ. 22 5FC QMBGSK-NMR?QQGSK NSKN DMPACQ MSR RFPCC (NMQGRGTC) /?+ GMLQ DMP CTCPW RUM (NMQGRGTC) ,+ GMLQ GR NSKNQ GL, RFSQ RFC ACJJ JMQCQ ? NMQGRGTC AF?PEC ?R CTCPW AWAJC MD RFC NSKN. 23 5FC NPMRCGLQ ?JJMU ? ACJJ RM QCJCAR UF?R AMKNMSLB UGJJ @C RP?LQNMPRCB, KCCRGLE RFC LCCBQ MD RFC ACJJ ?LB LMR @PGLEGLE GL ?LWRFGLE CJQC. 24 *MLQ ?PC AF?PECB, ?LB AMLQCOSCLRJW, RFCW ?PC FWBPMNFGJGA ?LB A?LLMR ?QQMAG?RC UGRF RFC JGNGB NMPRGML MD RFC KCK@P?LC. *MLQ KSQR @C RP?LQNMPRCB @W A?PPGCP NPMRCGLQ MP GML AF?LLCJQ.
Chapter 6
1 Figure 6.8 " AMKNMQR NGJC BCAMKNMQGLE GQ ?L CVCPEMLGA NPMACQQ; CLRF?JNW GLAPC?QCQ (CLCPEW GQ PCJC?QCB) ?LB CLRPMNW GLAPC?QCQ (J?PEC KMJCASJCQ ?PC @PMICL BMUL GLRM QK?JJCP MLCQ). " @?@W BCTCJMNGLE DPMK ? DCPRGJGXCB CEE GQ ?L CLBCPEMLGA NPMACQQ; CLRF?JNW BCAPC?QCQ (CLCPEW GQ ?@QMP@CB) ?LB CLRPMNW BCAPC?QCQ. 4?LB ?PR @CGLE BCQRPMWCB GQ ?L CVCPEMLGA NPMACQQ; RFCPC GQ LM AF?LEC GL CLRF?JNW, @SR CLRPMNW GLAPC?QCQ. " @?JJ PMJJGLE BMULFGJJ GQ ?L CVCPEMLGA NPMACQQ; CLRF?JNW BCAPC?QCQ (CLCPEW GQ PCJC?QCB), @SR RFCPC GQ LM AF?LEC GL CLRF?JNW. 2 Figure 6.10 /M. 8C A?L QRMPC AFCKGA?J CLCPEW @CA?SQC MD RFC LCCB RM MTCPAMKC RFC @?PPGCP RM GRQ @PC?IBMUL. 3 Figure 6.14 5FPCC QMBGSK GMLQ AMSJB @C KMTCB @W RFC FWBPMJWQGQ MD MLC "51 KMJCASJC. 5FC x( MD RFC AMSNJCB PC?ARGML KSQR @C LCE?RGTC. .MTCKCLR MD RFPCC QMBGSK GMLQ ?APMQQ RFC KCK@P?LC UGJJ R?IC 6.3 IA?J MD CLCPEW (2.1 IA?J `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5FGQ GQ ?L?@MJGQK. 17 &LCPEW GQ PCOSGPCB DMP ACJJSJ?P KMRGML, RFPMSEF @C?RGLE MD AGJG? MP DJ?ECJJ?, ?Q UCJJ ?Q FSK?L KMRGML, NPMBSACB @W KSQAJC AMLRP?ARGML. $CJJQ ?JQM LCCB CLCPEW RM NCPDMPK BGECQRGML, ?Q FSK?LQ PCOSGPC CLCPEW RM BGECQR DMMB. 18 " QNMLR?LCMSQ PC?ARGML GQ MLC RF?R F?Q ? LCE?RGTC x( ?LB RFSQ PCJC?QCQ CLCPEW. )MUCTCP, ? QNMLR?LCMSQ PC?ARGML LCCB LMR MAASP OSGAIJW MP QSBBCLJW JGIC ?L GLQR?LR?LCMSQ PC?ARGML. *R K?W MAASP MTCP JMLE NCPGMBQ BSC RM ? J?PEC CLCPEW MD ?ARGT?RGML, UFGAF NPCTCLRQ RFC PC?ARGML DPMK MAASPPGLE OSGAIJW. 19 5FC RP?LQGRGML QR?RC GQ ?JU?WQ FGEFCP GL CLCPEW RF?L RFC PC?AR?LRQ ?LB RFC NPMBSARQ MD ? PC?ARGML (RFCPCDMPC, ?@MTC), PCE?PBJCQQ MD UFCRFCP RFC PC?ARGML GQ CLBCPEMLGA MP CVCPEMLGA. 20 5FC ?LR D?PK F?B JMUCP CLRPMNW @CDMPC RFC C?PRFOS?IC @CA?SQC GR U?Q ? FGEFJW MPBCPCB QWQRCK. "DRCP RFC C?PRFOS?IC, RFC QWQRCK @CA?KC KSAF KMPC BGQMPBCPCB ?LB F?B FGEFCP CLRPMNW. 21 8FGJC AMMIGLE, DMMB GQ FC?RGLE SN ML RFC QRMTC, @SR LMR ?JJ MD RFC FC?R EMCQ RM AMMIGLE RFC DMMB, QMKC MD GR GQ JMQR ?Q FC?R CLCPEW RM RFC QSPPMSLBGLE ?GP, GLAPC?QGLE CLRPMNW. 8FGJC BPGTGLE, A?PQ @SPL E?QMJGLC RM PSL RFC CLEGLC ?LB KMTC RFC A?P. 5FGQ PC?ARGML GQ LMR AMKNJCRCJW CDDGAGCLR, ?Q QMKC CLCPEW BSPGLE RFGQ NPMACQQ GQ JMQR ?Q FC?R CLCPEW, UFGAF GQ UFW RFC FMMB ?LB RFC AMKNMLCLRQ SLBCPLC?RF GR FC?R SN UFGJC RFC CLEGLC GQ RSPLCB ML. 5FC RGPCQ ?JQM FC?R SN @CA?SQC MD DPGARGML UGRF RFC N?TCKCLR, UFGAF GQ ?BBGRGML?J CLCPEW JMQQ. 5FGQ CLCPEW RP?LQDCP, JGIC ?JJ MRFCPQ, ?JQM GLAPC?QCQ CLRPMNW. 22 5FC ?ARGT?RGML CLCPEW DMP FWBPMJWQGQ GQ TCPW JMU. /MR MLJW GQ "51 FWBPMJWQGQ ?L CVCPEMLGA NPMACQQ UGRF ? J?PEC yx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sQ LCCBQ, DCCB@?AI GLFG@GRGML CDDCARGTCJW A?SQCQ RFC ACJJ RM BCAPC?QC NPMBSARGML MD RF?R N?PRGASJ?P NPMBSAR. *L ECLCP?J, RFGQ PCBSACQ RFC NPMBSARGML MD QSNCPDJSMSQ NPMBSARQ ?LB AMLQCPTCQ CLCPEW, K?VGKGXGLE CLCPEW CDDGAGCLAW.
Chapter 7
1 Figure 7.11 "DRCP %/1 NMGQMLGLE, RFC CJCARPML RP?LQNMPR AF?GL A?L LM JMLECP DMPK ? NPMRML EP?BGCLR, ?LB "51 QWLRF?QC A?L LM JMLECP K?IC "51. %/1 GQ ?L CDDCARGTC BGCR BPSE @CA?SQC GR SLAMSNJCQ "51 QWLRFCQGQ; GL MRFCP UMPBQ, ?DRCP R?IGLE GR, ? NCPQML M@R?GLQ JCQQ CLCPEW MSR MD RFC DMMB FC MP QFC C?RQ. *LRCPCQRGLEJW, MLC MD RFC UMPQR QGBC CDDCARQ MD RFGQ BPSE GQ FWNCPRFCPKG?, MP MTCPFC?RGLE MD RFC @MBW. 4GLAC "51 A?LLMR @C DMPKCB, RFC CLCPEW DPMK CJCARPML RP?LQNMPR GQ JMQR ?Q FC?R. 2 Figure 7.12 "DRCP AW?LGBC NMGQMLGLE, RFC CJCARPML RP?LQNMPR AF?GL A?L LM JMLECP NSKN CJCARPMLQ GLRM RFC GLRCPKCK@P?LC QN?AC. 5FC N) MD RFC GLRCPKCK@P?LC QN?AC UMSJB GLAPC?QC, RFC N) EP?BGCLR UMSJB BCAPC?QC, ?LB "51 QWLRFCQGQ UMSJB QRMN. 3 Figure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MLC N?RFU?W K?W MAASP GL ?LMRFCP, ?LB RFCW A?L KMTC DPMK MLC N?RFU?W RM ?LMRFCP DJSGBJW GL PCQNMLQC RM RFC LCCBQ MD RFC ACJJ. 26 $GRP?RC A?L GLFG@GR NFMQNFMDPSARMIGL?QC @W DCCB@?AI PCESJ?RGML. 27 /CE?RGTC DCCB@?AI KCAF?LGQKQ ?ARS?JJW AMLRPMJ ? NPMACQQ; GR A?L RSPL GR MDD, UFCPC?Q NMQGRGTC DCCB@?AI ?AACJCP?RCQ RFC NPMACQQ, ?JJMUGLE RFC ACJJ LM AMLRPMJ MTCP GR. /CE?RGTC DCCB@?AI L?RSP?JJW K?GLR?GLQ FMKCMQR?QGQ, UFCPC?Q NMQGRGTC DCCB@?AI BPGTCQ RFC QWQRCK ?U?W DPMK COSGJG@PGSK.
Chapter 8
1 Figure 8.6 -CTCJQ MD A?P@ML BGMVGBC (? LCACQQ?PW NFMRMQWLRFCRGA QS@QRP?RC) UGJJ GKKCBG?RCJW D?JJ. "Q ? PCQSJR, RFC P?RC MD NFMRMQWLRFCQGQ UGJJ @C GLFG@GRCB. 2 Figure 8.15 ". 3 Figure 8.18 % 4 " 5 $ 6 # 7 # 8 " 9 # 10 $ 11 # 12 % 13 $ 14 $ 15 " 16 5FC MSRAMKC MD JGEFR PC?ARGMLQ GL NFMRMQWLRFCQGQ GQ RFC AMLTCPQGML MD QMJ?P CLCPEW GLRM AFCKGA?J CLCPEW RF?R RFC AFJMPMNJ?QRQ A?L SQC RM BM UMPI (KMQRJW ?L?@MJGA NPMBSARGML MD A?P@MFWBP?RCQ DPMK A?P@ML BGMVGBC). 17 #CA?SQC JGMLQ C?R ?LGK?JQ RF?R C?R NJ?LRQ. 18 5FC CLCPEW A?PPGCPQ RF?R KMTC DPMK RFC JGEFR-BCNCLBCLR PC?ARGML RM RFC JGEFR-GLBCNCLBCLR MLC ?PC tDSJJu @CA?SQC RFCW @PGLE CLCPEW. "DRCP RFC CLCPEW GQ PCJC?QCB, RFC tCKNRWu CLCPEW A?PPGCPQ PCRSPL RM RFC JGEFR-BCNCLBCLR PC?ARGML RM M@R?GL KMPC CLCPEW. 5FCPC GQ LMR KSAF ?ARS?J KMTCKCLR GLTMJTCB. #MRF "51 ?LB /"%1) ?PC NPMBSACB GL RFC QRPMK? UFCPC RFCW ?PC ?JQM SQCB ?LB PCAMLTCPRCB GLRM "%1, 1G, ?LB /"%1+. 19 " NFMRML MD JGEFR FGRQ ?L ?LRCLL? KMJCASJC GL NFMRMQWQRCK **, ?LB RFC CLCPEW PCJC?QCB @W GR RP?TCJQ RFPMSEF MRFCP ?LRCLL? KMJCASJCQ RM RFC PC?ARGML ACLRCP. 5FC CLCPEW A?SQCQ ?L CJCARPML RM JC?TC ? KMJCASJC MD AFJMPMNFWJJ ( RM ? NPGK?PW CJCARPML ?AACNRMP NPMRCGL. 5FC CJCARPML RP?TCJQ RFPMSEF RFC CJCARPML RP?LQNMPR AF?GL ?LB GQ ?AACNRCB @W ? NGEKCLR KMJCASJC GL NFMRMQWQRCK *. 20 #MRF MD RFCQC KMJCASJCQ A?PPW CLCPEW; GL RFC A?QC MD /"%1), GR F?Q PCBSAGLE NMUCP RF?R GQ SQCB RM DSCJ RFC NPMACQQ MD K?IGLE A?P@MFWBP?RC KMJCASJCQ GL JGEFR-GLBCNCLBCLR PC?ARGMLQ. 21 #CA?SQC 3S#1, RFC KMJCASJC LCCBCB ?R RFC QR?PR MD RFC AWAJC, GQ PCECLCP?RCB DPMK (31. 22 /MLC MD RFC AWAJC AMSJB R?IC NJ?AC, @CA?SQC 3S#GQ$0 GQ CQQCLRG?J GL DGVGLE A?P@ML BGMVGBC. 4NCAGDGA?JJW, 3S#GQ$0 A?R?JWXCQ RFC PC?ARGML @CRUCCL A?P@ML BGMVGBC ?LB 3S#1 ?R RFC QR?PR MD RFC AWAJC. 23 #CA?SQC (31 F?Q RFPCC A?P@ML ?RMKQ, ?LB C?AF RSPL MD RFC AWAJC R?ICQ GL MLC A?P@ML ?RMK GL RFC DMPK MD A?P@ML BGMVGBC.
Chapter 9
1 Figure 9.8 $. 5FC BMULQRPC?K ACJJSJ?P PCQNMLQC UMSJB @C GLFG@GRCB. 2 Figure 9.10 &3, UMSJB @CAMKC NCPK?LCLRJW ?ARGT?RCB, PCQSJRGLE GL ACJJ NPMJGDCP?RGML, KGEP?RGML, ?BFCQGML, ?LB RFC EPMURF MD LCU @JMMB TCQQCJQ. "NMNRMQGQ UMSJB @C GLFG@GRCB. 3 Figure 9.17 $. 4 Figure 9.18 . (<9,<: NPMBSACQ ? @GMDGJK @CA?SQC RFC FGEFCP ACJJ BCLQGRW GL RFC @GMDGJK NCPKGRQ RFC DMPK?RGML MD ? BCLQC QSPD?AC RF?R FCJNQ NPMRCAR RFC @?ARCPG? DPMK ?LRG@GMRGAQ. 5 # 6 $ 7 # 8 " 9 % 10 $ 11 # 12 # 13 % 14 $ 15 $ 16 % 17 *LRP?ACJJSJ?P QGEL?JGLE MAASPQ UGRFGL ? ACJJ, ?LB GLRCPACJJSJ?P QGEL?JGLE MAASPQ @CRUCCL ACJJQ. 18 5FC QCAPCRCB JGE?LBQ ?PC OSGAIJW PCKMTCB @W BCEP?B?RGML MP PC?@QMPNRGML GLRM RFC ACJJ QM RF?R RFCW A?LLMR RP?TCJ D?P. 19 *LRCPL?J PCACNRMPQ ?PC JMA?RCB GLQGBC RFC ACJJ, ?LB RFCGP JGE?LBQ CLRCP RFC ACJJ RM @GLB RFC PCACNRMP. 5FC AMKNJCV DMPKCB @W RFC GLRCPL?J PCACNRMP ?LB RFC JGE?LB RFCL CLRCPQ RFC LSAJCSQ ?LB BGPCARJW ?DDCARQ NPMRCGL NPMBSARGML @W @GLBGLE RM RFC AFPMKMQMK?J %/" ?LB GLGRG?RGLE RFC K?IGLE MD K3/" RF?R AMBCQ DMP NPMRCGLQ. $CJJQSPD?AC PCACNRMPQ, FMUCTCP, ?PC CK@CBBCB GL RFC NJ?QK? KCK@P?LC, ?LB RFCGP JGE?LBQ BM LMR CLRCP RFC ACJJ. #GLBGLE MD RFC JGE?LB RM RFC ACJJ-QSPD?AC PCACNRMP GLGRG?RCQ ? ACJJ QGEL?JGLE A?QA?BC ?LB BMCQ LMR BGPCARJW GLDJSCLAC RFC K?IGLE MD NPMRCGLQ; FMUCTCP, GR K?W GLTMJTC RFC ?ARGT?RGML MD GLRP?ACJJSJ?P NPMRCGLQ. 20 "L GML AF?LLCJ PCACNRMP MNCLCB SN ? NMPC GL RFC KCK@P?LC, UFGAF ?JJMUCB RFC GMLGA BWC RM KMTC GLRM RFC ACJJ. 21 %GDDCPCLR ACJJQ NPMBSAC BGDDCPCLR NPMRCGLQ, GLAJSBGLE ACJJ-QSPD?AC PCACNRMPQ ?LB QGEL?JGLE N?RFU?W AMKNMLCLRQ. 5FCPCDMPC, RFCW PCQNMLB RM BGDDCPCLR JGE?LBQ, ?LB RFC QCAMLB KCQQCLECPQ ?ARGT?RC BGDDCPCLR N?RFU?WQ. 4GEL?J GLRCEP?RGML A?L ?JQM AF?LEC RFC CLB PCQSJR MD QGEL?JGLE. 22 5FC @GLBGLE MD RFC JGE?LB RM RFC CVRP?ACJJSJ?P BMK?GL UMSJB ?ARGT?RC RFC N?RFU?W LMPK?JJW ?ARGT?RCB @W RFC PCACNRMP BML?RGLE RFC GLRP?ACJJSJ?P BMK?GL. 23 *D ? IGL?QC GQ KSR?RCB QM RF?R GR GQ ?JU?WQ ?ARGT?RCB, GR UGJJ AMLRGLSMSQJW QGEL?J RFPMSEF RFC N?RFU?W ?LB JC?B RM SLAMLRPMJJCB EPMURF ?LB NMQQG@JW A?LACP. *D ? IGL?QC GQ KSR?RCB QM RF?R GR A?LLMR DSLARGML, RFC ACJJ UGJJ LMR PCQNMLB RM JGE?LB @GLBGLE. 24 3CACNRMPQ ML RFC ACJJ QSPD?AC KSQR @C GL AMLR?AR UGRF RFC CVRP?ACJJSJ?P K?RPGV GL MPBCP RM PCACGTC NMQGRGTC QGEL?JQ RF?R ?JJMU RFC ACJJ RM JGTC. *D RFC PCACNRMPQ ?PC LMR ?ARGT?RCB @W @GLBGLE, RFC ACJJ UGJJ SLBCPEM ?NMNRMQGQ. 5FGQ CLQSPCQ RF?R ACJJQ ?PC GL RFC AMPPCAR NJ?AC GL RFC @MBW ?LB FCJNQ RM NPCTCLR GLT?QGTC ACJJ EPMURF ?Q MAASPQ GL KCR?QR?QGQ GL A?LACP. 25 :C?QRQ ?PC CSI?PWMRCQ ?LB F?TC K?LW MD RFC Q?KC QWQRCKQ RF?R FSK?LQ BM; FMUCTCP, RFCW ?PC QGLEJC-ACJJCB, QM RFCW ?PC C?QW RM EPMU, EPMU P?NGBJW, F?TC ? QFMPR ECLCP?RGML RGKC, ?LB ?PC KSAF QGKNJCP RF?L FSK?LQ. 26 .SJRGACJJSJ?P MPE?LGQKQ KSQR AMMPBGL?RC K?LW BGDDCPCLR CTCLRQ GL BGDDCPCLR ACJJ RWNCQ RF?R K?W @C TCPW BGQR?LR DPMK C?AF MRFCP. 4GLEJC-ACJJCB MPE?LGQKQ ?PC MLJW AMLACPLCB UGRF RFCGP GKKCBG?RC CLTGPMLKCLR ?LB RFC NPCQCLAC MD MRFCP ACJJQ GL RFC ?PC?.
Chapter 10
1 Figure 10.6 %. 5FC IGLCRMAFMPC @CAMKCQ ?RR?AFCB RM RFC KGRMRGA QNGLBJC. 4GQRCP AFPMK?RGBQ JGLC SN ?R RFC KCR?NF?QC NJ?RC. $MFCQGL NPMRCGLQ @PC?I BMUL ?LB RFC QGQRCP AFPMK?RGBQ QCN?P?RC. 5FC LSAJCSQ PCDMPKQ ?LB RFC ACJJ BGTGBCQ. 2 Figure 10.13 3@ ?LB MRFCP LCE?RGTC PCESJ?RMPW NPMRCGLQ AMLRPMJ ACJJ BGTGQGML ?LB RFCPCDMPC NPCTCLR RFC DMPK?RGML MD RSKMPQ. .SR?RGMLQ RF?R NPCTCLR RFCQC NPMRCGLQ DPMK A?PPWGLE MSR RFCGP DSLARGML A?L
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PCQSJR GL A?LACP. 3 Figure 10.14 %. &6 @GLBGLE K?PIQ N53 DMP BCEP?B?RGML. 4 $ 5 # 6 % 7 % 8 # 9 % 10 # 11 # 12 % 13 $ 14 " 15 " 16 " 17 $ 18 % 19 % 20 $ 21 " 22 $ 23 $ 24 % 25 % 26 $ 27 # 28 )SK?L QMK?RGA ACJJQ F?TC 46 AFPMKMQMKCQ: 22 N?GPQ ?LB 2 QCV AFPMKMQMKCQ RF?R K?W MP K?W LMR DMPK ? N?GP. 5FGQ GQ RFC 25 MP BGNJMGB AMLBGRGML. )SK?L E?KCRCQ F?TC 23 AFPMKMQMKCQ, MLC C?AF MD 23 SLGOSC AFPMKMQMKCQ, MLC MD UFGAF GQ ? QCV AFPMKMQMKC. 5FGQ GQ RFC 5 MP F?NJMGB AMLBGRGML. 29 5FC ECLMKC AMLQGQRQ MD RFC QSK RMR?J MD ?L MPE?LGQKs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tGKKMPR?JGXCB.u 41 " NPMRM-MLAMECLC GQ ? QCEKCLR MD %/" RF?R AMBCQ DMP MLC MD RFC NMQGRGTC ACJJ AWAJC PCESJ?RMPQ. *D RF?R ECLC @CAMKCQ KSR?RCB QM RF?R GR NPMBSACQ ? FWNCP?ARGT?RCB NPMRCGL NPMBSAR, GR GQ AMLQGBCPCB ?L MLAMECLC. " RSKMP QSNNPCQQMP ECLC GQ ? QCEKCLR MD %/" RF?R AMBCQ DMP MLC MD RFC LCE?RGTC ACJJ AWAJC PCESJ?RMPQ. *D RF?R ECLC @CAMKCQ KSR?RCB QM RF?R RFC NPMRCGL NPMBSAR @CAMKCQ JCQQ ?ARGTC, RFC ACJJ AWAJC UGJJ PSL SLAFCAICB. " QGLEJC MLAMECLC A?L GLGRG?RC ?@LMPK?J ACJJ BGTGQGMLQ; FMUCTCP, RSKMP QSNNPCQQMPQ JMQC RFCGP CDDCARGTCLCQQ MLJW UFCL @MRF AMNGCQ MD RFC ECLC ?PC B?K?ECB. 42 3CESJ?RMPW KCAF?LGQKQ RF?R KGEFR @C JMQR GLAJSBC KMLGRMPGLE MD RFC OS?JGRW MD RFC ECLMKGA %/", PCAPSGRGLE MD PCN?GP CLXWKCQ, ?LB RFC RPGEECPGLE MD ?NMNRMQGQ. 43 *D ? ACJJ F?Q B?K?ECB %/", RFC JGICJGFMMB MD NPMBSAGLE D?SJRW NPMRCGLQ GQ FGEFCP. 5FC B?SEFRCP ACJJQ MD QSAF ? B?K?ECB N?PCLR ACJJ UMSJB ?JQM NPMBSAC D?SJRW NPMRCGLQ RF?R KGEFR CTCLRS?JJW @CAMKC A?LACPMSQ. *D N53 PCAMELGXCQ RFGQ B?K?EC ?LB RPGEECPQ RFC ACJJ RM QCJD-BCQRPSAR, RFC B?K?ECB %/" GQ BCEP?BCB ?LB PCAWAJCB. /M DSPRFCP F?PK AMKCQ RM RFC MPE?LGQK. "LMRFCP FC?JRFW ACJJ GQ RPGEECPCB RM BGTGBC GLQRC?B. 44 5FC AMKKML AMKNMLCLRQ MD CSI?PWMRGA ACJJ BGTGQGML ?LB @GL?PW DGQQGML ?PC %/" BSNJGA?RGML, QCEPCE?RGML MD BSNJGA?RCB AFPMKMQMKCQ, ?LB BGTGQGML MD RFC AWRMNJ?QKGA AMLRCLRQ. 45 "Q RFC AFPMKMQMKC GQ @CGLE BSNJGA?RCB, C?AF MPGEGL KMTCQ ?U?W DPMK RFC QR?PRGLE NMGLR MD PCNJGA?RGML. 5FC AFPMKMQMKCQ ?PC ?RR?AFCB RM RFC ACJJ KCK@P?LC TG? NPMRCGLQ; RFC EPMURF MD RFC KCK@P?LC ?Q RFC ACJJ CJMLE?RCQ ?GBQ GL RFCGP KMTCKCLR.
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Chapter 11
1 Figure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s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
Chapter 12
1 Figure 12.5 :MS A?LLMR @C QSPC GD RFC NJ?LR GQ FMKMXWEMSQ MP FCRCPMXWEMSQ ?Q RFC B?R? QCR GQ RMM QK?JJ: @W P?LBMK AF?LAC, ?JJ RFPCC NJ?LRQ KGEFR F?TC ?AOSGPCB MLJW RFC BMKGL?LR ECLC CTCL GD RFC PCACQQGTC MLC GQ NPCQCLR. *D RFC PMSLB NC? N?PCLR GQ FCRCPMXWEMSQ, RFCPC GQ ? MLC-CGEFRF NPM@?@GJGRW RF?R ? P?LBMK Q?KNJC MD RFPCC NPMECLW NC?Q UGJJ ?JJ @C PMSLB. 2 Figure 12.6 *LBGTGBS?J 1 F?Q RFC ECLMRWNC ((. *LBGTGBS?J 2 F?Q RFC ECLMRWNC A(. *LBGTGBS?J 3 F?Q RFC ECLMRWNC A(. 3 Figure 12.12 )?JD MD RFC DCK?JC MDDQNPGLE UMSJB @C FCRCPMXWEMSQ (9$9>) UGRF PCB CWCQ, ?LB F?JD UMSJB @C FMKMXWEMSQ PCACQQGTC (9>9>) UGRF UFGRC CWCQ. )?JD MD RFC K?JC MDDQNPGLE UMSJB @C FCKGXWEMSQ BMKGL?LR (9$:) UGRFC PCB WCQ, ?LB F?JD UMSJB @C FCKGXWEMSQ PCACQQGTC (9>:) UGRF UFGRC CWCQ. 4 Figure 12.16 5FC NMQQG@JC ECLMRWNCQ ?PC 1N::, 1N:W, NN::, ?LB NN:W. 5FC DMPKCP RUM ECLMRWNCQ UMSJB PCQSJR GL NJ?LRQ UGRF NSPNJC DJMUCPQ ?LB WCJJMU NC?Q, UFGJC RFC J?RRCP RUM ECLMRWNCQ UMSJB PCQSJR GL NJ?LRQ UGRF UFGRC DJMUCPQ UGRF WCJJMU NC?Q, DMP ? 1:1 P?RGM MD C?AF NFCLMRWNC. :MS MLJW LCCB ? 2 ` 2 1SLLCRR QOS?PC (DMSP QOS?PCQ RMR?J) RM BM RFGQ ?L?JWQGQ @CA?SQC RUM MD RFC ?JJCJCQ ?PC FMKMXWEMSQ. 5 " 6 # 7 # 8 $ 9 " 10 $ 11 % 12 % 13 $ 14 " 15 # 16 % 17 5FC E?PBCL NC? GQ QCQQGJC ?LB F?Q DJMUCPQ RF?R AJMQC RGEFRJW BSPGLE QCJD-NMJJGL?RGML. 5FCQC DC?RSPCQ FCJN RM NPCTCLR ?AAGBCLR?J MP SLGLRCLRGML?J DCPRGJGX?RGMLQ RF?R AMSJB F?TC BGKGLGQFCB RFC ?AASP?AW MD .CLBCJsQ B?R?. 18 5UM QCRQ MD 10 N?PCLRQ UMSJB @C SQCB. *L RFC DGPQR APMQQ, NMJJCL UMSJB @C RP?LQDCPPCB DPMK ? RPSC-@PCCBGLE R?JJ NJ?LR RM RFC QRGEK? MD ? RPSC-@PCCBGLE BU?PD NJ?LR. *L RFC QCAMLB APMQQ, NMJJCL UMSJB @C RP?LQDCPPCB DPMK ? RPSC-@PCCBGLE BU?PD NJ?LR RM RFC QRGEK? MD ? RPSC-@PCCBGLE R?JJ NJ?LR. 'MP C?AF APMQQ, '1 ?LB '2 MDDQNPGLE UMSJB @C ?L?JWXCB RM BCRCPKGLC GD MDDQNPGLE RP?GRQ UCPC ?DDCARCB ?AAMPBGLE RM UFGAF N?PCLR BML?RCB C?AF RP?GR. 19 #CA?SQC ?VG?J GQ BMKGL?LR, RFC ECLC UMSJB @C BCQGEL?RCB ?Q A. '1 UMSJB @C ?JJ FCRCPMXWEMSQ A( UGRF ?VG?J NFCLMRWNC. '2 UMSJB F?TC NMQQG@JC ECLMRWNCQ MD AA, A(, ?LB ((; RFCQC UMSJB AMPPCQNMLB RM ?VG?J, ?VG?J, ?LB RCPKGL?J NFCLMRWNCQ, PCQNCARGTCJW. 20 5FC 1SLLCRR QOS?PC UMSJB @C 2 ` 2 ?LB UGJJ F?TC ! ?LB ! ?JMLE RFC RMN, ?LB ! ?LB ; ?JMLE RFC JCDR QGBC. $JMAIUGQC DPMK RFC RMN JCDR, RFC ECLMRWNCQ JGQRCB UGRFGL RFC @MVCQ UGJJ @C !;, !;,
1438
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;;, ?LB ;;. 5FC NFCLMRWNGA P?RGM UGJJ @C 1 R?JJ:1 BU?PD. 21 /M, K?JCQ A?L MLJW CVNPCQQ AMJMP @JGLBLCQQ. 5FCW A?LLMR A?PPW GR @CA?SQC ?L GLBGTGBS?J LCCBQ RUM 9 AFPMKMQMKCQ RM @C ? A?PPGCP. 22 $MLQGBCPGLE C?AF ECLC QCN?P?RCJW, RFC APMQQ ?R A UGJJ NPMBSAC MDDQNPGLE MD UFGAF F?JD ?PC AA ?LB F?JD ?PC A(; B UGJJ NPMBSAC ?JJ B); C UGJJ NPMBSAC F?JD C* ?LB F?JD **. 1PMNMPRGMLQ RFCL ?PC (1/2) ` (1) ` (1/2), MP 1/4 AAB)C*; AMLRGLSGLE DMP RFC MRFCP NMQQG@GJGRGCQ WGCJBQ 1/4 AAB)**, 1/4 A(B)C*, ?LB 1/4 A(B)**. 5FC NPMNMPRGMLQ RFCPCDMPC ?PC 1:1:1:1. 23 &NGQR?QGQ BCQAPG@CQ ?L ?LR?EMLGQRGA GLRCP?ARGML @CRUCCL ECLCQ UFCPCGL MLC ECLC K?QIQ MP GLRCPDCPCQ UGRF RFC CVNPCQQGML MD ?LMRFCP. 5FC ECLC RF?R GQ GLRCPDCPGLE GQ PCDCPPCB RM ?Q CNGQR?RGA, ?Q GD GR GQ tQR?LBGLE SNMLu RFC MRFCP (FWNMQR?RGA) ECLC RM @JMAI GRQ CVNPCQQGML. 24 5FC APMQQ A?L @C PCNPCQCLRCB ?Q ? 4 ` 4 1SLLCRR QOS?PC, UGRF RFC DMJJMUGLE E?KCRCQ DMP C?AF N?PCLR: $&, $@, >&, ?LB >@. 'MP ?JJ 12 MD RFC MDDQNPGLE RF?R CVNPCQQ ? BMKGL?LR $ ECLC, RFC MDDQNPGLE UGJJ @C UFGRC. 5FC RFPCC MDDQNPGLE RF?R ?PC FMKMXWEMSQ PCACQQGTC DMP > @SR CVNPCQQ ? BMKGL?LR & ECLC UGJJ @C WCJJMU. 5FC PCK?GLGLE >>@@ MDDQNPGLE UGJJ @C EPCCL.
Chapter 13
1 Figure 13.3 /M. 5FC NPCBGARCB DPCOSCLAW MD PCAMK@GL?LR MDDQNPGLE P?LECQ DPMK 0% (DMP JGLICB RP?GRQ) RM 50% (DMP SLJGLICB RP?GRQ). 2 Figure 13.4 % 3 Figure 13.6 #. 4 " 5 $ 6 $ 7 " 8 # 9 $ 10 $ 11 " 12 % 13 " 14 5FC $FPMKMQMK?J 5FCMPW MD *LFCPGR?LAC NPMNMQCB RF?R ECLCQ PCQGBC ML AFPMKMQMKCQ. 5FC SLBCPQR?LBGLE RF?R AFPMKMQMKCQ ?PC JGLC?P ?PP?WQ MD ECLCQ CVNJ?GLCB JGLI?EC, ?LB APMQQGLE MTCP CVNJ?GLCB PCAMK@GL?RGML. 15 &V?AR BG?EP?K QRWJC UGJJ T?PW; BG?EP?K QFMSJB JMMI JGIC Figure 13.6.
Chapter 14
1 Figure 14.10 $MKN?PRKCLR?JGX?RGML CL?@JCQ ? CSI?PWMRGA ACJJ RM BGTGBC NPMACQQCQ GLRM BGQAPCRC QRCNQ QM GR A?L @SGJB KMPC AMKNJCV NPMRCGL ?LB 3/" NPMBSARQ. #SR RFCPC GQ ?L ?BT?LR?EC RM F?TGLE ? QGLEJC AMKN?PRKCLR ?Q UCJJ: 3/" ?LB NPMRCGL QWLRFCQGQ MAASPQ KSAF KMPC OSGAIJW GL ? NPMI?PWMRGA ACJJ. 2 Figure 14.14 %/" JGE?QC, ?Q RFGQ CLXWKC HMGLQ RMECRFCP 0I?X?IG DP?EKCLRQ. 3 Figure 14.21 *D RFPCC LSAJCMRGBCQ ?PC ?BBCB, MLC ?BBGRGML?J ?KGLM ?AGB UGJJ @C GLAMPNMP?RCB GLRM RFC NPMRCGL AF?GL, @SR RFC PC?BGLE DP?KC UMLR QFGDR. 4 $ 5 % 6 % 7 % 8 # 9 $ 10 % 11 # 12 " 13 % 14 $ 15 # 16 -GTC 3 ACJJQ ?AOSGPCB ECLCRGA GLDMPK?RGML DPMK RFC FC?R-IGJJCB 4 ACJJQ RF?R tRP?LQDMPKCBu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sQ CVNCPGKCLRQ UGRF E. *630 EPMUL GL 15/ BCBSACB RFGQ DGLBGLE. 21 "R ?L MPGEGL MD PCNJGA?RGML, RUM PCNJGA?RGML DMPIQ ?PC DMPKCB RF?R ?PC CVRCLBCB GL RUM BGPCARGMLQ. 0L RFC J?EEGLE QRP?LB, 0I?X?IG DP?EKCLRQ ?PC DMPKCB GL ? BGQAMLRGLSMSQ K?LLCP. 22 4FMPR %/" DP?EKCLRQ ?PC DMPKCB ML RFC J?EEGLE QRP?LB QWLRFCQGXCB GL ? BGPCARGML ?U?W DPMK RFC PCNJGA?RGML DMPI. 5FCQC ?PC QWLRFCQGXCB @W %/" NMJ. 23 1333 QCAMLBQ MP 22.2 KGLSRCQ. 24 "R RFC PCNJGA?RGML DMPI, RFC CTCLRQ R?IGLE NJ?AC ?PC FCJGA?QC ?ARGML, @GLBGLE MD QGLEJC-QRP?LB @GLBGLE NPMRCGLQ, NPGKCP QWLRFCQGQ, ?LB QWLRFCQGQ MD LCU QRP?LBQ. *D RFCPC GQ ? KSR?RCB FCJGA?QC ECLC, RFC PCNJGA?RGML DMPI UGJJ LMR @C CVRCLBCB. 25 1PGKCP NPMTGBCQ ? 3'-0) EPMSN DMP %/" NMJ RM QR?PR ?BBGLE LSAJCMRGBCQ. 5FCPC UMSJB @C LM PC?ARGML GL RFC RS@C UGRFMSR ? NPGKCP, ?LB LM @?LBQ UMSJB @C TGQG@JC ML RFC CJCARPMNFMPCQGQ. 26 5CJMKCP?QC F?Q ?L GL@SGJR 3/" RCKNJ?RC RF?R CVRCLBQ RFC 3' CLB, QM NPGKCP GQ QWLRFCQGXCB ?LB CVRCLBCB. 5FSQ, RFC CLBQ ?PC NPMRCARCB. 27 .SR?RGMLQ ?PC LMR PCN?GPCB, ?Q GL RFC A?QC MD VCPMBCPK? NGEKCLRMQ?. (CLC DSLARGML K?W @C ?DDCARCB MP GR K?W LMR @C CVNPCQQCB.
Chapter 15
1 Figure 15.11 /M. 1PMI?PWMRCQ SQC BGDDCPCLR NPMKMRCPQ RF?L CSI?PWMRCQ. 2 Figure 15.13 .SR?RGMLQ GL RFC QNJGACMQMKC PCAMELGRGML QCOSCLAC ?R C?AF CLB MD RFC GLRPML, MP GL RFC NPMRCGLQ ?LB 3/"Q RF?R K?IC SN RFC QNJGACMQMKC, K?W GKN?GP QNJGAGLE. .SR?RGMLQ K?W ?JQM ?BB LCU QNJGACMQMKC PCAMELGRGML QGRCQ. 4NJGAGLE CPPMPQ AMSJB JC?B RM GLRPMLQ @CGLE PCR?GLCB GL QNJGACB 3/", CVMLQ @CGLE CVAGQCB, MP AF?LECQ GL RFC JMA?RGML MD RFC QNJGAC QGRC. 3 Figure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1439
RFC CLB MD RFC ECLC, RFC NMJWKCP?QC QR?JJQ ?R ? PSL MD ( LSAJCMRGBCQ ML RFC %/" RCKNJ?RC. 5FC PFM NPMRCGL AMJJGBCQ UGRF RFC NMJWKCP?QC ?LB PCJC?QCQ K3/" DPMK RFC RP?LQAPGNRGML @S@@JC. 3FM-GLBCNCLBCLR RCPKGL?RGML GQ AMLRPMJJCB @W QNCAGDGA QCOSCLACQ GL RFC %/" RCKNJ?RC QRP?LB. "Q RFC NMJWKCP?QC LC?PQ RFC CLB MD RFC ECLC @CGLE RP?LQAPG@CB, GR CLAMSLRCPQ ? PCEGML PGAF GL $p( LSAJCMRGBCQ. 5FGQ APC?RCQ ?L K3/" F?GPNGL RF?R A?SQCQ RFC NMJWKCP?QC RM QR?JJ PGEFR ?Q GR @CEGLQ RM RP?LQAPG@C ? PCEGML PGAF GL "p5 LSAJCMRGBCQ. #CA?SQC "p6 @MLBQ ?PC JCQQ RFCPKMQR?@JC, RFC AMPC CLXWKC D?JJQ ?U?W. 18 5FC K3/" UMSJB @C: 5'-"6(($$((66"66""($"-3'. 5FC NPMRCGL UMSJB @C: ."(:. &TCL RFMSEF RFCPC ?PC QGV AMBMLQ, RFC DGDRF AMBML AMPPCQNMLBQ RM ? QRMN, QM RFC QGVRF AMBML UMSJB LMR @C RP?LQJ?RCB. 19 /SAJCMRGBC AF?LECQ GL RFC RFGPB NMQGRGML MD AMBMLQ K?W LMR AF?LEC RFC ?KGLM ?AGB ?LB UMSJB F?TC LM CDDCAR ML RFC NPMRCGL. 0RFCP LSAJCMRGBC AF?LECQ RF?R AF?LEC GKNMPR?LR ?KGLM ?AGBQ MP APC?RC MP BCJCRC QR?PR MP QRMN AMBMLQ UMSJB F?TC QCTCPC CDDCARQ ML RFC ?KGLM ?AGB QCOSCLAC MD RFC NPMRCGL.
Chapter 16
1 Figure 16.5 5PWNRMNF?L GQ ?L ?KGLM ?AGB CQQCLRG?J DMP K?IGLE NPMRCGLQ, QM RFC ACJJ ?JU?WQ LCCBQ RM F?TC QMKC ML F?LB. )MUCTCP, GD NJCLRW MD RPWNRMNF?L GQ NPCQCLR, GR GQ U?QRCDSJ RM K?IC KMPC, ?LB RFC CVNPCQQGML MD RFC ;97 PCACNRMP GQ PCNPCQQCB. -?ARMQC, ? QSE?P DMSLB GL KGJI, GQ LMR ?JU?WQ ?T?GJ?@JC. *R K?ICQ LM QCLQC RM K?IC RFC CLXWKCQ LCACQQ?PW RM BGECQR ?L CLCPEW QMSPAC RF?R GQ LMR ?T?GJ?@JC, QM RFC 3(* MNCPML GQ MLJW RSPLCB ML UFCL J?ARMQC GQ NPCQCLR. 2 Figure 16.7 5FC LSAJCMQMKCQ UMSJB N?AI KMPC RGEFRJW RMECRFCP. 3 Figure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
1440
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Chapter 17
1 Figure 17.6 #. 5FC CVNCPGKCLR UMSJB PCQSJR GL @JSC AMJMLGCQ MLJW. 2 Figure 17.8 %MJJW U?Q ? 'GLL%MPQCR QFCCN @CA?SQC CTCL RFMSEF RFC MPGEGL?J ACJJ A?KC DPMK ? 4AMRRGQF @J?AID?AC QFCCN ?LB RFC QSPPME?RC KMRFCP U?Q ? 4AMRRGQF @J?AID?AC, RFC %/" A?KC DPMK ? 'GLL-%MPQCR. 3 Figure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
Chapter 18
1 Figure 18.14 -MQQ MD ECLCRGA K?RCPG?J GQ ?JKMQR ?JU?WQ JCRF?J, QM MDDQNPGLE UGRF 25+1 AFPMKMQMKCQ ?PC KMPC JGICJW RM QSPTGTC. 2 Figure 18.22 'SQGML GQ KMQR JGICJW RM MAASP @CA?SQC RFC RUM QNCAGCQ UGJJ GLRCP?AR KMPC ?LB QGKGJ?P RP?GRQ GL DMMB ?AOSGQGRGML UGJJ @C QCJCARCB. 3 Figure 18.23 "LQUCP # 4 # 5 % 6 % 7 % 8 " 9 # 10 # 11 $ 12 $ 13 $ 14 % 15 " 16 $ 17 5FC NJ?LRQ RF?R A?L @CQR SQC RFC PCQMSPACQ MD RFC ?PC?, GLAJSBGLE AMKNCRGLE UGRF MRFCP GLBGTGBS?JQ DMP RFMQC PCQMSPACQ UGJJ NPMBSAC KMPC QCCBQ RFCKQCJTCQ ?LB RFMQC RP?GRQ RF?R ?JJMUCB RFCK RM @CRRCP SQC RFC PCQMSPACQ UGJJ GLAPC?QC GL RFC NMNSJ?RGML MD RFC LCVR ECLCP?RGML. 18 7CQRGEG?J QRPSARSPCQ ?PC AMLQGBCPCB CTGBCLAC DMP CTMJSRGML @CA?SQC KMQR QRPSARSPCQ BM LMR CVGQR GL ?L MPE?LGQK UGRFMSR QCPTGLE QMKC DSLARGML CGRFCP NPCQCLRJW MP GL RFC N?QR. " TCQRGEG?J QRPSARSPC GLBGA?RCQ ? N?QR DMPK MP DSLARGML RF?R F?Q QGLAC AF?LECB, @SR RFC QRPSARSPC PCK?GLQ NPCQCLR @CA?SQC GR F?B ? DSLARGML GL RFC ?LACQRMP. 19 *L QAGCLAC, ? RFCMPW GQ ? RFMPMSEFJW RCQRCB ?LB TCPGDGCB QCR MD CVNJ?L?RGMLQ DMP ? @MBW MD M@QCPT?RGMLQ MD L?RSPC. *R GQ RFC QRPMLECQR DMPK MD ILMUJCBEC GL QAGCLAC. *L AMLRP?QR, ? RFCMPW GL AMKKML TCPL?ASJ?P A?L KC?L ? ESCQQ MP QNCASJ?RGML ?@MSR QMKCRFGLE, KC?LGLE RF?R RFC ILMUJCBEC GKNJGCB @W RFC RFCMPW GQ TCPW UC?I. 20 5FC QR?RCKCLR GKNJGCQ RF?R RFCPC GQ ? EM?J RM CTMJSRGML ?LB RF?R RFC KMLICW PCNPCQCLRQ EPC?RCP NPMEPCQQ RM RF?R EM?J RF?L RFC KMSQC. #MRF QNCAGCQ ?PC JGICJW RM @C UCJJ ?B?NRCB RM RFCGP N?PRGASJ?P CLTGPMLKCLRQ, UFGAF GQ RFC MSRAMKC MD L?RSP?J QCJCARGML. 21 0PE?LGQKQ MD MLC QNCAGCQ A?L ?PPGTC RM ?L GQJ?LB RMECRFCP ?LB RFCL BGQNCPQC RFPMSEFMSR RFC AF?GL, C?AF QCRRJGLE GLRM BGDDCPCLR LGAFCQ ?LB CVNJMGRGLE BGDDCPCLR DMMB PCQMSPACQ RM PCBSAC AMKNCRGRGML. 22 *R GQ JGICJW RFC RUM QNCAGCQ UMSJB QR?PR RM PCNPMBSAC UGRF C?AF MRFCP. %CNCLBGLE ML RFC TG?@GJGRW MD RFCGP MDDQNPGLE, RFCW K?W DSQC @?AI GLRM MLC QNCAGCQ. 23 5FC DMPK?RGML MD E?KCRCQ UGRF LCU 5 LSK@CPQ A?L MAASP GL MLC ECLCP?RGML. "DRCP ? AMSNJC MD ECLCP?RGMLQ, CLMSEF MD RFCQC LCU FW@PGBQ A?L DMPK RM PCNPMBSAC RMECRFCP ?Q ? LCU QNCAGCQ. 24 #MRF KMBCJQ AMLRGLSC RM AMLDMPK RM RFC PSJCQ MD L?RSP?J QCJCARGML, ?LB RFC GLDJSCLACQ MD ECLC DJMU, ECLCRGA BPGDR, ?LB KSR?RGML. 25 *D RFC FW@PGB MDDQNPGLE ?PC ?Q DGR MP KMPC DGR RF?L RFC N?PCLRQ, PCNPMBSARGML UMSJB JGICJW AMLRGLSC @CRUCCL @MRF QNCAGCQ ?LB RFC FW@PGBQ, CTCLRS?JJW @PGLEGLE ?JJ MPE?LGQKQ SLBCP RFC SK@PCJJ? MD MLC QNCAGCQ.
Chapter 19
1 Figure 19.2 5FC CVNCARCB BGQRPG@SRGML GQ 320 77, 1607T, ?LB 20 TT NJ?LRQ. 1J?LRQ UGRF 77 MP 7T ECLMRWNCQ UMSJB F?TC TGMJCR DJMUCPQ, ?LB NJ?LRQ UGRF RFC TT ECLMRWNC UMSJB F?TC UFGRC DJMUCPQ, QM ? RMR?J MD 480 NJ?LRQ UMSJB @C CVNCARCB RM F?TC TGMJCR DJMUCPQ, ?LB 20 NJ?LRQ UMSJB F?TC UFGRC DJMUCPQ. 2 Figure 19.4 (CLCRGA BPGDR GQ JGICJW RM MAASP KMPC P?NGBJW ML ?L GQJ?LB UFCPC QK?JJCP NMNSJ?RGMLQ ?PC CVNCARCB RM MAASP. 3
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1441
Figure 19.8 .MRFQ F?TC QFGDRCB RM ? JGEFRCP AMJMP. 4 $ 5 " 6 % 7 % 8 $ 9 # 10 " 11 $ 12 % 13 $ 14 " 15 % 16 N = (8*2 + 4)/48 = .42; O = (12*2 + 4)/48 = .58; N2 = .17; 2NO = .48; O2 = .34 17 5FC )?PBW-8CGL@CPE NPGLAGNJC MD COSGJG@PGSK GQ SQCB RM BCQAPG@C RFC ECLCRGA K?ICSN MD ? NMNSJ?RGML. 5FC RFCMPW QR?RCQ RF?R ? NMNSJ?RGMLsQ ?JJCJC ?LB ECLMRWNC DPCOSCLAGCQ ?PC GLFCPCLRJW QR?@JC: SLJCQQ QMKC IGLB MD CTMJSRGML?PW DMPAC GQ ?ARGLE SNML RFC NMNSJ?RGML, ECLCP?RGML ?DRCP ECLCP?RGML MD RFC NMNSJ?RGML UMSJB A?PPW RFC Q?KC ECLCQ, ?LB GLBGTGBS?JQ UMSJB, ?Q ? UFMJC, JMMI CQQCLRG?JJW RFC Q?KC. 18 3CB GQ PCACQQGTC QM O2 = 200/ 800 = 0.25; O = 0.5; N = 1-O = 0.5; N2 = 0.25; 2NO = 0.5. :MS UMSJB CVNCAR 200 FMKMXWEMSQ @JSC DJMUCPQ, 400 FCRCPMXWEMSQ @JSC DJMUCPQ, ?LB 200 PCB DJMUCPQ. 19 " FSPPGA?LC IGJJQ ? J?PEC NCPACLR?EC MD ? NMNSJ?RGML MD Q?LB-BUCJJGLE APSQR?AC?LQqMLJW ? DCU GLBGTGBS?JQ QSPTGTC. 5FC ?JJCJCQ A?PPGCB @W RFMQC QSPTGTGLE GLBGTGBS?JQ UMSJB PCNPCQCLR RFC CLRGPC NMNSJ?RGMLsQ ECLC NMMJ. *D RFMQC QSPTGTGLE GLBGTGBS?JQ ?PC LMR PCNPCQCLR?RGTC MD RFC MPGEGL?J NMNSJ?RGML, RFC NMQR-FSPPGA?LC ECLC NMMJ UGJJ BGDDCP DPMK RFC MPGEGL?J ECLC NMMJ. 20 5FC RFCMPW MD L?RSP?J QCJCARGML QRCKQ DPMK RFC M@QCPT?RGML RF?R QMKC GLBGTGBS?JQ GL ? NMNSJ?RGML QSPTGTC JMLECP ?LB F?TC KMPC MDDQNPGLE RF?L MRFCPQ: RFSQ, KMPC MD RFCGP ECLCQ ?PC N?QQCB RM RFC LCVR ECLCP?RGML. 'MP CV?KNJC, ? @GE, NMUCPDSJ K?JC EMPGJJ? GQ KSAF KMPC JGICJW RF?L ? QK?JJCP, UC?ICP MLC RM @CAMKC RFC NMNSJ?RGMLsQ QGJTCP@?AI: RFC N?AIsQ JC?BCP UFM K?RCQ D?P KMPC RF?L RFC MRFCP K?JCQ MD RFC EPMSN. 5FCPCDMPC, RFC N?AI JC?BCP UGJJ D?RFCP KMPC MDDQNPGLE UFM QF?PC F?JD MD FGQ ECLCQ ?LB ?PC JGICJW RM EPMU @GEECP ?LB QRPMLECP JGIC RFCGP D?RFCP. 0TCP RGKC, RFC ECLCQ DMP @GEECP QGXC UGJJ GLAPC?QC GL DPCOSCLAW GL RFC NMNSJ?RGML, ?LB RFC ?TCP?EC @MBW QGXC, ?Q ? PCQSJR, EPMU J?PECP ML ?TCP?EC. 21 " AJGLC GQ ? RWNC MD ECMEP?NFGA T?PG?RGML RF?R GQ QCCL GL NMNSJ?RGMLQ MD ? EGTCL QNCAGCQ RF?R T?PW EP?BS?JJW ?APMQQ ?L CAMJMEGA?J EP?BGCLR. 'MP CV?KNJC, U?PK-@JMMBCB ?LGK?JQ RCLB RM F?TC J?PECP @MBGCQ GL RFC AMMJCP AJGK?RCQ AJMQCP RM RFC C?PRFsQ NMJCQ, ?JJMUGLE RFCK RM @CRRCP AMLQCPTC FC?R. 5FGQ GQ AMLQGBCPCB ? J?RGRSBGL?J AJGLC. 'JMUCPGLE NJ?LRQ RCLB RM @JMMK ?R BGDDCPCLR RGKCQ BCNCLBGLE ML UFCPC RFCW ?PC ?JMLE RFC QJMNC MD ? KMSLR?GL. 5FGQ GQ ILMUL ?Q ?L ?JRGRSBGL?J AJGLC. 22 5FC NC?AMAIsQ R?GJ GQ ? EMMB CV?KNJC MD RFC F?LBGA?N NPGLAGNJC. 5FC R?GJ, UFGAF K?ICQ RFC K?JCQ KMPC TGQG@JC RM NPCB?RMPQ ?LB JCQQ ?@JC RM CQA?NC, GQ AJC?PJW ? BGQ?BT?LR?EC RM RFC @GPBsQ QSPTGT?J. #SR @CA?SQC GR GQ ? BGQ?BT?LR?EC, MLJW RFC KMQR DGR K?JCQ QFMSJB @C ?@JC RM QSPTGTC UGRF GR. 5FSQ, RFC R?GJ QCPTCQ ?Q ?L FMLCQR QGEL?J MD OS?JGRW RM RFC DCK?JCQ MD RFC NMNSJ?RGML; RFCPCDMPC, RFC K?JC UGJJ C?PL KMPC K?RGLEQ ?LB EPC?RCP PCNPMBSARGTC QSAACQQ. 23 5FCPC ?PC QCTCP?J U?WQ CTMJSRGML A?L ?DDCAR NMNSJ?RGML T?PG?RGML: QR?@GJGXGLE QCJCARGML, BGPCARGML?J QCJCARGML, BGTCPQGDWGLE QCJCARGML, DPCOSCLAW-BCNCLBCLR QCJCARGML, ?LB QCVS?J QCJCARGML. "Q RFCQC GLDJSCLAC RFC ?JJCJC DPCOSCLAGCQ GL ? NMNSJ?RGML, GLBGTGBS?JQ A?L CGRFCP @CAMKC KMPC MP JCQQ PCJ?RCB, ?LB RFC NFCLMRWNCQ BGQNJ?WCB A?L @CAMKC KMPC QGKGJ?P MP KMPC BGQN?P?RC.
Chapter 20
1 Figure 20.6 $?RQ ?LB BMEQ ?PC N?PR MD RFC Q?KC EPMSN ?R DGTC JCTCJQ: @MRF ?PC GL RFC BMK?GL &SI?PW?, RFC IGLEBMK "LGK?JG?, RFC NFWJSK $FMPB?R?, RFC AJ?QQ .?KK?JG?, ?LB RFC MPBCP $?PLGTMP?. 2 Figure 20.10 3?@@GRQ ?LB FSK?LQ @CJMLE GL RFC AJ?BC RF?R GLAJSBCQ ?LGK?JQ UGRF F?GP. 5FC ?KLGMRGA CEE CTMJTCB @CDMPC F?GP @CA?SQC RFC "KLGMR? AJ?BC GQ J?PECP RF?L RFC AJ?BC RF?R CLAMKN?QQCQ ?LGK?JQ UGRF F?GP. 3 Figure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
Chapter 21
1 Figure 21.4 % 2 Figure 21.8 5FC FMQR ACJJ A?L AMLRGLSC RM K?IC LCU TGPSQ N?PRGAJCQ. 3 Figure 21.10 $ 4 # 5 % 6 % 7 % 8 # 9 $ 10 % 11 " 12 % 13 $ 14 % 15 $ 16 " 17 7GPSQCQ N?QQ RFPMSEF DGJRCPQ RF?R CJGKGL?RCB ?JJ @?ARCPG? RF?R UCPC TGQG@JC GL RFC JGEFR KGAPMQAMNCQ ?R RFC RGKC. "Q RFC @?ARCPG?-DPCC DGJRP?RC AMSJB QRGJJ A?SQC GLDCARGMLQ UFCL EGTCL RM ? FC?JRFW MPE?LGQK, RFGQ M@QCPT?RGML BCKMLQRP?RCB RFC CVGQRCLAC MD TCPW QK?JJ GLDCARGMSQ ?ECLRQ. 5FCQC ?ECLRQ UCPC J?RCP QFMUL RM @C SLPCJ?RCB RM @?ARCPG? ?LB UCPC AJ?QQGDGCB ?Q TGPSQCQ. 18 5FC TGPSQ A?LsR ?RR?AF RM BME ACJJQ, @CA?SQC BME ACJJQ BM LMR CVNPCQQ RFC PCACNRMPQ DMP RFC TGPSQ ?LB/ MP RFCPC GQ LM ACJJ UGRFGL RFC BME RF?R GQ NCPKGQQGTC DMP TGP?J PCNJGA?RGML. 19 3CTCPQC RP?LQAPGNR?QC GQ LCCBCB RM K?IC KMPC )*7-1 TGPSQCQ, QM R?PECRGLE RFC PCTCPQC RP?LQAPGNR?QC CLXWKC K?W @C ? U?W RM GLFG@GR RFC PCNJGA?RGML
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
Chapter 22
1 Figure 22.8 5FC CVRP?ACJJSJ?P K?RPGV ?LB MSRCP J?WCP MD ACJJQ NPMRCARQ RFC GLLCP @?ARCPG?. 5FC AJMQC NPMVGKGRW MD ACJJQ ?JQM D?AGJGR?RCQ J?RCP?J ECLC RP?LQDCP, ? NPMACQQ @W UFGAF ECLCQ QSAF ?Q ?LRG@GMRGA PCQGQR?LAC ECLCQ ?PC RP?LQDCPPCB DPMK MLC @?ARCPGSK RM ?LMRFCP. "LB CTCL GD J?RCP?J ECLC RP?LQDCP BMCQ LMR MAASP, MLC @?ARCPGSK RF?R NPMBSACQ ?L CVM-CLXWKC RF?R BCQRPMWQ ?LRG@GMRGA K?W Q?TC LCGEF@MPGLE @?ARCPG?. 2 Figure 22.15 " 3 Figure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
Chapter 23
1 Figure 23.5 "JJ CSI?PWMRGA ACJJQ F?TC KGRMAFMLBPG?, @SR LMR ?JJ CSI?PWMRGA ACJJQ F?TC AFJMPMNJ?QRQ. 2 Figure 23.15 $ 3 Figure 23.18 $ 4 % 5 $ 6 $ 7 % 8 % 9 # 10 # 11 $ 12 $ 13 $ 14 " 15 % 16 " 17 # 18 &SI?PWMRGA ACJJQ ?PMQC RFPMSEF CLBMQWK@GMRGA CTCLRQ RF?R E?TC PGQC RM RFC CLCPEW-NPMBSAGLE MPE?LCJJCQ UGRFGL RFC CSI?PWMRGA ACJJQ QSAF ?Q KGRMAFMLBPG? ?LB AFJMPMNJ?QRQ. 5FC LSAJC?P ECLMKC MD CSI?PWMRCQ GQ PCJ?RCB KMQR AJMQCJW RM RFC "PAF?C?, QM GR K?W F?TC @CCL ?L C?PJW ?PAF?C?L RF?R CLESJDCB ? @?ARCPG?J ACJJ RF?R CTMJTCB GLRM ? KGRMAFMLBPGML. .GRMAFMLBPG? ?NNC?P RM F?TC MPGEGL?RCB DPMK ?L ?JNF?NPMRCM@?ARCPGSK, UFCPC?Q AFJMPMNJ?QRQ MPGEGL?RCB ?Q ? AW?LM@?ARCPGSK. 5FCPC GQ ?JQM CTGBCLAC MD QCAMLB?PW CLBMQWK@GMRGA CTCLRQ. 0RFCP ACJJ AMKNMLCLRQ K?W ?JQM F?TC PCQSJRCB DPMK CLBMQWK@GMRGA CTCLRQ. 19 5FC ?@GJGRW RM NCPDMPK QCVS?J PCNPMBSARGML ?JJMUQ NPMRGQRQ RM PCAMK@GLC RFCGP ECLCQ ?LB NPMBSAC LCU T?PG?RGMLQ MD NPMECLW RF?R K?W @C @CRRCP QSGRCB RM RFC LCU CLTGPMLKCLR. *L AMLRP?QR, ?QCVS?J PCNPMBSARGML ECLCP?RCQ NPMECLW RF?R ?PC AJMLCQ MD RFC N?PCLR. 20 "Q ?L GLRCQRGL?J N?P?QGRC, G0(9+0( AWQRQ UMSJB @C CVNMQCB RM JMU N) GL RFC QRMK?AF ?AGBQ MD GRQ FMQR. 5M QSPTGTC RFGQ CLTGPMLKCLR ?LB PC?AF RFC GLRCQRGLC, RFC AWQRQ UMSJB F?TC RM @C PCQGQR?LR RM ?AGBGA AMLBGRGMLQ. 21 6LJGIC "3=(, NPMRGQRQ GL RFC ECLSQ C(<3,97( ?ARS?JJW ?PC J?PEC, KSJRGLSAJC?RC, QGLEJC ACJJQ. #CA?SQC RFCQC MPE?LGQKQ SLBCPEM KGRMQGQ UGRFMSR AWRMIGLCQGQ ?LB J?AI AWRMNJ?QKGA BGTGQGMLQ, RFCW A?LLMR @C AMLQGBCPCB RPSJW KSJRGACJJSJ?P. 22 #W BCDGLGRGML, ?L M@JGE?RC Q?NPM@C J?AIQ RFC ?@GJGRW RM NCPDMPK NFMRMQWLRFCQGQ, QM GR A?LLMR BGPCARJW M@R?GL LSRPGRGML @W QC?PAFGLE DMP JGEFR. *LQRC?B, ? AFCKMR?ARGA KCAF?LGQK RF?R QCLQCQ RFC MBMPQ PCJC?QCB BSPGLE BCA?W KGEFR @C ? KMPC CDDCARGTC QCLQGLE MPE?L DMP ? Q?NPM@C. 23 P3(:46+0<4 N?P?QGRCQ GLDCAR FSK?LQ ?LB A?SQC K?J?PG?. )MUCTCP, RFCW KSQR AMKNJCRC N?PR MD RFCGP JGDC AWAJC UGRFGL A567/,3,: KMQOSGRMCQ, ?LB RFCW A?L MLJW GLDCAR FSK?LQ TG? RFC @GRC UMSLB MD ? KMQOSGRM. *D RFC KMQOSGRM NMNSJ?RGML GQ BCAPC?QCB, RFCL DCUCP P3(:46+0<4 UMSJB @C ?@JC RM BCTCJMN ?LB GLDCAR FSK?LQ, RFCPC@W PCBSAGLE RFC GLAGBCLAC MD FSK?L GLDCARGMLQ UGRF RFGQ N?P?QGRC. 24 5FC RPWN?LMQMKCQ
!00%.$)8 ! X !00%.$)8
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RF?R A?SQC RFGQ BGQC?QC ?PC A?N?@JC MD CVNPCQQGLE ? EJWAMNPMRCGL AM?R UGRF ? BGDDCPCLR KMJCASJ?P QRPSARSPC UGRF C?AF ECLCP?RGML. #CA?SQC RFC GKKSLC QWQRCK KSQR PCQNMLB RM QNCAGDGA ?LRGECLQ RM P?GQC ? KC?LGLEDSJ BCDCLQC, RFC AF?LEGLE L?RSPC MD RPWN?LMQMKC ?LRGECLQ NPCTCLRQ RFC GKKSLC QWQRCK DPMK CTCP AJC?PGLE RFGQ GLDCARGML. .?QQGTC RPWN?LMQMKC GLDCARGML CTCLRS?JJW JC?BQ RM FMQR MPE?L D?GJSPC ?LB BC?RF.
Chapter 24
1 Figure 24.13 " 2 Figure 24.16 % 3 Figure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sQ ?AFGCTCKCLRQ RM NSPGDW PCJG?@JC QRP?GLQ MD @?ICPsQ WC?QR RM NPMBSAC @PC?B AMLQGQRCLRJW.
Chapter 25
1 Figure 25.5 #. 2 Figure 25.14 $. 3 Figure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qRFCW DMPK J?PEC JC?TCQ ?LB @P?LAFGLE PMMRQ.
Chapter 26
1 Figure 26.8 #. 5FC BGNJMGB XWEMRC DMPKQ ?DRCP RFC NMJJCL RS@C F?Q DGLGQFCB DMPKGLE, QM RF?R RFC K?JC ECLCP?RGTC LSAJCG A?L DSQC UGRF RFC DCK?JC E?KCRMNFWRC. 2 Figure 26.15 8GRFMSR ? KCE?QNMP?LEGSK, ?L CEE UMSJB LMR DMPK; UGRFMSR ? KGAPMQNMP?LEGSK, NMJJCL UMSJB LMR DMPK. 3 % 4 " 5 $ 6 " 7 " 8 % 9 # 10 " 11 $ 12 " 13 # 14 # 15 $ 16 " 17 % 18 % 19 #MRF NMJJGL?RGML ?LB FCP@GTMPW AMLRPG@SRCB RM BGTCPQGRW, UGRF NJ?LRQ LCCBGLE RM ?RRP?AR QMKC GLQCARQ ?LB PCNCJ MRFCPQ. 20 4CCBQ ?LB NMJJCL ?JJMUCB NJ?LRQ RM PCNPMBSAC GL ?@QCLAC MD U?RCP. 5FGQ ?JJMUCB RFCK RM CVN?LB RFCGP P?LEC MLRM BPW J?LB ?LB RM QSPTGTC BPMSEFR AMLBGRGMLQ. 21 5FC RPCCQ ?PC ?B?NRCB RM ?PGB UC?RFCP, ?LB BM LMR JMQC ?Q KSAF U?RCP BSC RM RP?LQNGP?RGML ?Q LML-AMLGDCPQ. 22 5FC DMSP KMBCPL-B?W NFWJ? MD EWKLMQNCPKQ ?PC $MLGDCPMNFWR?, $WA?BMNFWR?, (GLEIMNFWR?, ?LB (LCRMNFWR?. 23 5FC PCQCK@J?LAC @CRUCCL AWA?BQ ?LB N?JK RPCCQ GQ MLJW QSNCPDGAG?J. $WA?BQ ?PC EWKLMQNCPKQ ?LB BM LMR @C?P DJMUCPQ MP DPSGR. $WA?BQ NPMBSAC AMLCQ: J?PEC, DCK?JC AMLCQ RF?R NPMBSAC L?ICB QCCBQ, ?LB QK?JJCP K?JC AMLCQ ML QCN?P?RC NJ?LRQ. 1?JKQ BM LMR. 24 "LEGMQNCPKQ ?PC QSAACQQDSJ @CA?SQC MD DJMUCPQ ?LB DPSGR. 5FCQC QRPSARSPCQ NPMRCAR PCNPMBSARGML DPMK T?PG?@GJGRW GL RFC CLTGPMLKCLR. 25 6QGLE ?LGK?J NMJJGL?RMPQ NPMKMRCQ APMQQ-NMJJGL?RGML ?LB GLAPC?QCQ ECLCRGA BGTCPQGRW. 5FC
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MBBQ RF?R RFC NMJJCL UGJJ PC?AF ?LMRFCP DJMUCP ?PC EPC?RJW GLAPC?QCB AMKN?PCB UGRF RFC P?LBMKLCQQ MD UGLB NMJJGL?RGML. 26 #GMBGTCPQGRW GQ RFC T?PG?RGML GL ?JJ DMPKQ MD JGDC. *R A?L PCDCP RM T?PG?RGML UGRFGL ? QNCAGCQ, UGRFGL ?L CAMQWQRCK, MP ML ?L CLRGPC NJ?LCR. *R GQ GKNMPR?LR @CA?SQC GR CLQSPCQ ? PCQMSPAC DMP LCU DMMB APMNQ ?LB KCBGAGLCQ. 1J?LR JGDC @?J?LACQ RFC CAMQWQRCKQ, NPMRCARQ U?RCPQFCBQ, KGRGE?RCQ CPMQGML, KMBCP?RCQ AJGK?RC, ?LB NPMTGBCQ QFCJRCP DMP K?LW ?LGK?J QNCAGCQ.
Chapter 27
1 Figure 27.5 5FC ?LGK?J KGEFR BCTCJMN RUM FC?BQ ?LB LM R?GJ. 2 Figure 27.6 $ 3 Figure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s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
Chapter 28
1 Figure 28.3 # 2 Figure 28.20 % 3 Figure 28.36 $ 4 # 5 % 6 % 7 $ 8 # 9 " 10 $ 11 # 12 % 13 $ 14 " 15 # 16 % 17 $ 18 1GL?AMAWRCQ ?PC CNGRFCJG?J-JGIC ACJJQ, DMPK RFC MSRCPKMQR J?WCP MD QNMLECQ, ?LB CLAJMQC ? HCJJW-JGIC QS@QR?LAC A?JJCB KCQMFWJ. *L QMKC QNMLECQ, NMPMAWRCQ DMPK MQRG?, QGLEJC RS@C-QF?NCB ACJJQ RF?R ?AR ?Q T?JTCQ RM PCESJ?RC RFC DJMU MD U?RCP GLRM RFC QNMLEMAMCJ. $FM?LMAWRCQ (tAMJJ?P ACJJQu) ?PC NPCQCLR ?R T?PGMSQ JMA?RGMLQ, BCNCLBGLE ML RFC RWNC MD QNMLEC, @SR RFCW ?JU?WQ JGLC QMKC QN?AC RFPMSEF UFGAF U?RCP DJMUQ ?LB ?PC SQCB GL DCCBGLE. 19 5FC QNMLECQ BP?U U?RCP A?PPWGLE DMMB N?PRGAJCQ GLRM RFC QNMLEMAMCJ SQGLE RFC @C?RGLE MD DJ?ECJJ? ML RFC AFM?LMAWRCQ. 5FC DMMB N?PRGAJCQ ?PC A?SEFR @W RFC AMJJ?P MD RFC AFM?LMAWRC ?LB ?PC @PMSEFR GLRM RFC ACJJ @W NF?EMAWRMQGQ. %GECQRGML MD RFC DMMB N?PRGAJC R?ICQ NJ?AC GLQGBC RFC ACJJ. 5FC BGDDCPCLAC @CRUCCL RFGQ ?LB RFC KCAF?LGQKQ MD MRFCP ?LGK?JQ GQ RF?R BGECQRGML R?ICQ NJ?AC UGRFGL ACJJQ P?RFCP RF?L MSRQGBC MD ACJJQ. *R KC?LQ RF?R RFC MPE?LGQK A?L DCCB MLJW ML N?PRGAJCQ QK?JJCP RF?L RFC ACJJQ RFCKQCJTCQ. 20 /CK?RMAWQRQ ?PC tQRGLEGLE ACJJQu BCQGELCB RM N?P?JWXC NPCW. 5FC LCK?RMAWQRQ AMLR?GL ? LCSPMRMVGL RF?R PCLBCPQ NPCW GKKM@GJC. 21 1MPGDCP?LQ BM LMR NMQQCQQ RPSC RGQQSCQ, UFGJC ALGB?PG?LQ BM F?TC RGQQSCQ. #CA?SQC MD RFGQ BGDDCPCLAC, NMPGDCP?LQ BM LMR F?TC ? LCPTMSQ QWQRCK MP KSQAJCQ DMP JMAMKMRGML, UFGAF ALGB?PG?LQ F?TC. 22 .MJJSQIQ F?TC ? J?PEC KSQASJ?P DMMR RF?R K?W @C KMBGDGCB GL T?PGMSQ U?WQ, QSAF
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?Q GLRM RCLR?AJCQ, @SR GR DSLARGMLQ GL JMAMKMRGML. 5FCW F?TC ? K?LRJC, ? QRPSARSPC MD RGQQSC RF?R AMTCPQ ?LB CLAJMQCQ RFC BMPQ?J NMPRGML MD RFC ?LGK?J, ?LB QCAPCRCQ RFC QFCJJ UFCL GR GQ NPCQCLR. 5FC K?LRJC CLAJMQCQ RFC K?LRJC A?TGRW, UFGAF FMSQCQ RFC EGJJQ (UFCL NPCQCLR), CVAPCRMPW NMPCQ, ?LSQ, ?LB EML?BMNMPCQ. 5FC AMCJMK MD KMJJSQIQ GQ PCQRPGARCB RM RFC PCEGML ?PMSLB RFC QWQRCKGA FC?PR. 5FC K?GL @MBW A?TGRW GQ ? FCKMAMCJ. .?LW KMJJSQIQ F?TC ? P?BSJ? LC?P RFC KMSRF RF?R GQ SQCB DMP QAP?NGLE DMMB. 23 .MJJSQIQ F?TC ? QFCJJ, CTCL GD GR GQ ? PCBSACB QFCJJ. /CKCPRGLCQ BM LMR F?TC ? QFCJJ. /CKCPRGLCQ F?TC ? NPM@MQAGQ; KMJJSQIQ BM LMR. /CKCPRGLCQ F?TC ? AJMQCB AGPASJ?RMPW QWQRCK, UFCPC?Q .MJJSQIQ F?TC ?L MNCL AGPASJ?RMPW QWQRCK. 24 *R GQ ? RPSC ?LGK?J UGRF ?R JC?QR PSBGKCLRQ MD RFC NFWQGMJMEGA?J QWQRCKQqDCCBGLE, LCPTMSQ, KSQAJC, ?LB PCNPMBSARGTCqDMSLB GL tFGEFCP ?LGK?JQu JGIC KGAC ?LB FSK?LQ. *R GQ QM QK?JJ RF?R J?PEC LSK@CPQ A?L @C P?GQCB GL 1CRPG BGQFCQ. *R PCNPMBSACQ P?NGBJW. *R GQ RP?LQN?PCLR QM RF?R CTCPW ACJJ GL RFC JGTGLE ?LGK?J A?L @C QCCL SLBCP RFC KGAPMQAMNC. #CDMPC GR BGCQ (?DRCP 2p3 UCCIQ), GR QFMUQ QGELQ MD ?EGLE ?LB RFSQ K?W NPMTGBC ECLCP?J AJSCQ ?Q RM RFC ?EGLE NPMACQQ. 25 5FCPC ?PC LCK?RMBCQ UGRF QCN?P?RC QCVCQ ?LB FCPK?NFPMBGRCQ GL ?BBGRGML RM QNCAGCQ RF?R PCNPMBSAC N?PRFCLMECLRGA?JJW. 5FC LCK?RMBC C(,569/()+0;0: ,3,.(5: F?Q ? QCJD-DCPRGJGXGLE FCPK?NFPMBGRC QCV ?LB ? NSPC K?JC QCV. 26 5FC "PRFPMNMB? GLAJSBC RFC )CV?NMB?, UFGAF ?PC K?LBG@SJ?RCQ UGRF QGV JCEQ, RFC .WPG?NMB?, UFGAF ?PC K?LBG@SJ?RCQ UGRF K?LW JCEQ ?LB GLAJSBC RFC ACLRGNCBCQ ?LB KGJJGNCBCQ, RFC $PSQR?AC?, UFGAF ?PC KMQRJW K?PGLC K?LBG@SJ?RCQ, ?LB RFC $FCJGACP?R?, UFGAF GLAJSBC RFC QNGBCPQ ?LB QAMPNGMLQ ?LB RFCGP IGL. 27 "PRFPMNMBQ F?TC ?L CVMQICJCRML, UFGAF GQ KGQQGLE GL ?LLCJGBQ. "PRFPMNMB QCEKCLR?RGML GQ KMPC QNCAG?JGXCB UGRF K?HMP MPE?LQ AMLACLRP?RCB GL @MBW R?EK?. "LLCJGB QCEKCLR?RGML GQ SQS?JJW KMPC SLGDMPK UGRF RFC GLRCQRGLC CVRCLBGLE RFPMSEF KMQR QCEKCLRQ. 28 5FC "QRCPMGBC? ?PC RFC QC? QR?PQ, RFC &AFGLMGBC? ?PC RFC QC? SPAFGLQ ?LB Q?LB BMJJ?PQ, RFC 0NFGSPMGBC? ?PC RFC @PGRRJC QR?PQ, RFC $PGLMGBC? ?PC RFC QC? JGJGCQ ?LB DC?RFCP QR?PQ, RFC )MJMRFSPMGBC? ?PC RFC QC? ASASK@CPQ.
Chapter 29
1 Figure 29.3 " 2 Figure 29.20 % 3 Figure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
Chapter 30
1 Figure 30.7 " ?LB #. 5FC AMPRCV, NGRF, ?LB CNGBCPKGQ ?PC K?BC MD N?PCLAFWK? ACJJQ. 2 Figure 30.32 :CQ, WMS A?L COS?JGXC RFC U?RCP JCTCJ @W ?BBGLE RFC QMJSRC RM RFC JCDR QGBC MD RFC RS@C QSAF RF?R U?RCP KMTCQ
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RMU?PB RFC JCDR SLRGJ RFC U?RCP JCTCJQ ?PC COS?J. 3 Figure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
Chapter 31
1 Figure 31.5 5FC ?GP AMLRCLR MD RFC QMGJ BCAPC?QCQ. 2 Figure 31.6 5FC " FMPGXML GQ RFC RMNQMGJ, ?LB RFC # FMPGXML GQ QS@QMGJ. 3 Figure 31.9 4MW@C?LQ ?PC ?@JC RM DGV LGRPMECL GL RFCGP PMMRQ, UFGAF ?PC LMR F?PTCQRCB ?R RFC CLB MD RFC EPMUGLE QC?QML. 5FC @CJMUEPMSLB LGRPMECL A?L @C SQCB GL RFC LCVR QC?QML @W RFC AMPL. 4 $ 5 # 6 " 7 # 8 % 9 " 10 # 11 # 12 # 13 " 14 " 15 " 16 %CDGAGCLAGCQ GL RFCQC LSRPGCLRQ AMSJB PCQSJR GL QRSLRCB EPMURF, QJMU EPMURF, ?LB AFJMPMQGQ. 17 T?L )CJKMLR QFMUCB RF?R NJ?LRQ BM LMR AMLQSKC QMGJ, UFGAF GQ AMPPCAR. )C ?JQM RFMSEFR RF?R NJ?LR EPMURF ?LB GLAPC?QCB UCGEFR PCQSJRCB DPMK RFC GLR?IC MD U?RCP, ? AMLAJSQGML RF?R F?Q QGLAC @CCL BGQNPMTCL. 18 "LQUCPQ K?W T?PW. &QQCLRG?J K?APMLSRPGCLRQ GLAJSBC A?P@ML, FWBPMECL, MVWECL, LGRPMECL, NFMQNFMPSQ, NMR?QQGSK, A?JAGSK, K?ELCQGSK, ?LB QSJDSP. &QQCLRG?J KGAPMLSRPGCLRQ GLAJSBC GPML, K?LE?LCQC, @MPML, KMJW@BCLSK, AMNNCP, XGLA, AFJMPGLC, LGAICJ, AM@?JR, QMBGSK, ?LB QGJGAML. 19 " KGLCP?J QMGJ DMPKQ DPMK RFC UC?RFCPGLE MD PMAIQ; GR GQ GLMPE?LGA K?RCPG?J. "L MPE?LGA QMGJ GQ DMPKCB DPMK QCBGKCLR?RGML; GR KMQRJW AMLQGQRQ MD FSKSQ. 20 1?PCLR K?RCPG?J, AJGK?RC, RMNMEP?NFW, @GMJMEGA?J D?ARMPQ, ?LB RGKC ?DDCAR QMGJ DMPK?RGML. 1?PCLR K?RCPG?J GQ RFC K?RCPG?J GL UFGAF QMGJQ DMPK. $JGK?RC BCQAPG@CQ FMU RCKNCP?RSPC, KMGQRSPC, ?LB UGLB A?SQC BGDDCPCLR N?RRCPLQ MD UC?RFCPGLE, GLDJSCLAGLE RFC AF?P?ARCPGQRGAQ MD RFC QMGJ. 5MNMEP?NFW ?DDCARQ RFC AF?P?ARCPGQRGAQ ?LB DCPRGJGRW MD ? QMGJ. #GMJMEGA?J D?ARMPQ GLAJSBC RFC NPCQCLAC MD JGTGLE MPE?LGQKQ RF?R EPC?RJW ?DDCAR QMGJ DMPK?RGML. 1PMACQQCQ QSAF ?Q DPCCXGLE ?LB RF?UGLE K?W NPMBSAC AP?AIQ GL PMAIQ; NJ?LR PMMRQ A?L NCLCRP?RC RFCQC APCTGACQ ?LB NPMBSAC KMPC DP?EKCLR?RGML. 5GKC ?DDCARQ QMGJ @CA?SQC QMGJ BCTCJMNQ MTCP JMLE NCPGMBQ. 21 5MNMEP?NFW ?DDCARQ U?RCP PSLMDD, UFGAF QRPGNQ ?U?W N?PCLR K?RCPG?J ?LB ?DDCARQ NJ?LR EPMURF. 4RCCNQ QMGJQ ?PC KMPC NPMLC RM CPMQGML ?LB K?W @C RFGLLCP RF?L QMGJQ RF?R ?PC ML JCTCJ QSPD?ACQ. 22 #CA?SQC GR GQ L?RSP?J ?LB BMCQ LMR PCOSGPC SQC MD ? LMLPCLCU?@JC PCQMSPAC, QSAF ?Q L?RSP?J E?Q. 23 1FMRMQWLRFCQGQ F?PTCQRQ ?LB QRMPCQ CLCPEW, UFCPC?Q @GMJMEGA?J LGRPMECL DGV?RGML PCOSGPCQ CLCPEW. 24 " LMBSJC PCQSJRQ DPMK RFC QWK@GMQGQ @CRUCCL ? NJ?LR ?LB @?ARCPGSK. 8GRFGL LMBSJCQ, RFC NPMACQQ MD LGRPMECL DGV?RGML ?JJMUQ RFC NJ?LR RM M@R?GL LGRPMECL DPMK RFC ?GP.
Chapter 32
1 Figure 32.3 1MJJCL (MP QNCPK); A?PNCJJ?RC; QR?KGL?RC. 2 Figure 32.8 #: 5FC NMJJCL RS@C UGJJ DMPK @SR UGJJ LMR @C ESGBCB RMU?PB RFC CEE. 3 Figure 32.20 # 4 # 5 % 6 " 7 " 8 # 9 # 10 % 11 $ 12 " 13 $ 14 % 15 $ 16 *LQGBC RFC DJMUCP ?PC RFC PCNPMBSARGTC MPE?LQ MD RFC NJ?LR. 5FC QR?KCL GQ RFC K?JC PCNPMBSARGTC MPE?L. 1MJJCL GQ NPMBSACB GL RFC QR?KCL. 5FC A?PNCJ GQ RFC DCK?JC PCNPMBSARGTC MPE?L. 5FC MT?PW GQ RFC QUMJJCL
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1447
@?QC MD RFC A?PNCJ UFCPC MTSJCQ ?PC DMSLB. /MR ?JJ DJMUCPQ F?TC CTCPW MLC MD RFC DMSP N?PRQ. 17 1J?LRQ F?TC RUM BGQRGLAR NF?QCQ GL RFCGP JGDCAWAJC: RFC E?KCRMNFWRC QR?EC ?LB RFC QNMPMNFWRC QR?EC. *L RFC E?KCRMNFWRC QR?EC, UFCL PCNPMBSARGTC ACJJQ SLBCPEM KCGMQGQ ?LB NPMBSAC F?NJMGB ACJJQ A?JJCB QNMPCQ, RFC E?KCRMNFWRC QR?EC @CEGLQ. 4NMPCQ BGTGBC @W ACJJ BGTGQGML RM DMPK NJ?LR QRPSARSPCQ MD ?L CLRGPCJW LCU NJ?LR. 5FC ACJJQ GL RFCQC QRPSARSPCQ MP NJ?LRQ ?PC F?NJMGB. 4MKC MD RFCQC ACJJQ SLBCPEM ACJJ BGTGQGML ?LB DMPK QCV ACJJQ. 'CPRGJGX?RGML, RFC HMGLGLE MD F?NJMGB QCV ACJJQ, @CEGLQ RFC QNMPMNFWRC QR?EC. $CJJQ DMPKCB GL RFGQ QR?EC F?TC RFC BGNJMGB LSK@CP MD AFPMKMQMKCQ. .CGMQGQ GL QMKC MD RFCQC ACJJQ DMPKQ QNMPCQ, ?LB RFC AWAJC @CEGLQ ?E?GL: ? NPMACQQ ILMUL ?Q ?JRCPL?RGML MD ECLCP?RGMLQ. 18 " RWNGA?J DJMUCP F?Q DMSP K?GL N?PRQ, MP UFMPJQ: RFC A?JWV, AMPMJJ?, ?LBPMCAGSK, ?LB EWLMCAGSK. 5FC MSRCPKMQR UFMPJ MD RFC DJMUCP F?Q EPCCL, JC?DW QRPSARSPCQ ILMUL ?Q QCN?JQ, UFGAF ?PC AMJJCARGTCJW A?JJCB RFC A?JWV. *R FCJNQ RM NPMRCAR RFC SLMNCLCB @SB. 5FC QCAMLB UFMPJ GQ K?BC SN MD @PGEFRJW AMJMPCB NCR?JQ RF?R ?PC ILMUL AMJJCARGTCJW ?Q RFC AMPMJJ?. 5FC RFGPB UFMPJ GQ RFC K?JC PCNPMBSARGTC QRPSARSPC ILMUL ?Q RFC ?LBPMCAGSK. 5FC ?LBPMCAGSK F?Q QR?KCLQ, UFGAF F?TC ?LRFCPQ ML ? QR?JI MP DGJ?KCLR. 1MJJCL EP?GLQ ?PC @MPLC ML RFC ?LRFCPQ. 5FC EWLMCAGSK GQ RFC DCK?JC PCNPMBSARGTC QRPSARSPC. 5FC A?PNCJ GQ RFC GLBGTGBS?J QRPSARSPC MD RFC EWLMCAGSK ?LB F?Q ? QRGEK?, RFC QR?JI MP QRWJC, ?LB RFC MT?PW. 19 *D ?JJ DMSP UFMPJQ MD ? DJMUCP ?PC NPCQCLR, GR GQ ? AMKNJCRC DJMUCP. *D ?LW MD RFC DMSP N?PRQ GQ KGQQGLE, GR GQ ILMUL ?Q GLAMKNJCRC. 'JMUCPQ RF?R AMLR?GL @MRF ?L ?LBPMCAGSK ?LB EWLMCAGSK ?PC A?JJCB ?LBPMEWLMSQ MP FCPK?NFPMBGRCQ. 5FMQC RF?R AMLR?GL MLJW ?L ?LBPMCAGSK ?PC ILMUL ?Q QR?KGL?RC DJMUCPQ, ?LB RFMQC RF?R F?TC MLJW A?PNCJQ ?PC ILMUL ?Q A?PNCJJ?RC. *D @MRF K?JC ?LB DCK?JC DJMUCPQ ?PC @MPLC ML RFC Q?KC NJ?LR, GR GQ A?JJCB KMLMCAGMSQ, UFGJC NJ?LRQ UGRF K?JC ?LB DCK?JC DJMUCPQ ML QCN?P?RC NJ?LRQ ?PC RCPKCB BGMCAGMSQ. 20 .?LW QCCBQ CLRCP ? NCPGMB MD GL?ARGTGRW MP CVRPCKCJW JMU KCR?@MJGA ?ARGTGRW, ? NPMACQQ ILMUL ?Q BMPK?LAW. %MPK?LAW ?JJMUQ QCCBQ RM RGBC MTCP SLD?TMP?@JC AMLBGRGMLQ ?LB ECPKGL?RC ML PCRSPL RM D?TMP?@JC AMLBGRGMLQ. '?TMP?@JC AMLBGRGMLQ AMSJB @C ?Q BGTCPQC ?Q KMGQRSPC, JGEFR, AMJB, DGPC, MP AFCKGA?J RPC?RKCLRQ. "DRCP FC?TW P?GLQ, K?LW LCU QCCBJGLEQ CKCPEC. 'MPCQR DGPCQ ?JQM JC?B RM RFC CKCPECLAC MD LCU QCCBJGLEQ. 21 4MKC DPSGRQ F?TC @SGJR-GL KCAF?LGQKQ RF?R ?JJMU RFCK RM BGQNCPQC QCCBQ @W RFCKQCJTCQ, @SR MRFCPQ PCOSGPC RFC ?QQGQR?LAC MD ?ECLRQ JGIC UGLB, U?RCP, ?LB ?LGK?JQ. 'PSGR RF?R ?PC BGQNCPQCB @W RFC UGLB ?PC JGEFR GL UCGEFR ?LB MDRCL F?TC UGLE-JGIC ?NNCLB?ECQ RF?R ?JJMU RFCK RM @C A?PPGCB @W RFC UGLB; MRFCP F?TC QRPSARSPCQ PCQCK@JGLE ? N?P?AFSRC RF?R ICCN RFCK ?DJM?R GL RFC UGLB. 4MKC DPSGRQ, QSAF ?Q RFMQC MD B?LBCJGMLQ, F?TC F?GPW, UCGEFRJCQQ QRPSARSPCQ RF?R ?JJMU RFCK RM DJM?R GL RFC UGLB. 'PSGRQ BGQNCPQCB @W U?RCP ?PC JGEFR ?LB @SMW?LR, EGTGLE RFCK RFC ?@GJGRW RM DJM?R; AMAMLSRQ ?PC MLC CV?KNJC. "LGK?JQ ?LB @GPBQ C?R DPSGRQ ?LB BGQNCPQC RFCGP QCCBQ @W JC?TGLE BPMNNGLEQ ?R BGQR?LR JMA?RGMLQ. 0RFCP ?LGK?JQ @SPW DPSGR RF?R K?W J?RCP ECPKGL?RC. 4MKC DPSGRQ QRGAI RM ?LGK?JQs @MBGCQ ?LB ?PC A?PPGCB RM LCU JMA?RGMLQ. 1CMNJC ?JQM AMLRPG@SRC RM QCCB BGQNCPQ?J UFCL RFCW A?PPW DPSGRQ RM LCU NJ?ACQ. 22 "QCVS?J PCNPMBSARGML BMCQ LMR PCOSGPC RFC CVNCLBGRSPC MD RFC NJ?LRs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
Chapter 33
1 Figure 33.11 " 2 Figure 33.21 #MRF NPMACQQCQ ?PC RFC PCQSJR MD LCE?RGTC DCCB@?AI JMMNQ. /CE?RGTC DCCB@?AI JMMNQ, UFGAF RCLB RM ICCN ? QWQRCK ?R COSGJG@PGSK, ?PC KMPC AMKKML RF?L NMQGRGTC DCCB@?AI JMMNQ. 3 Figure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sQ DSLARGMLGLE ?LB @MBW RCKNCP?RSPC. 4K?JJCP @MBGCB ?LGK?JQ F?TC ? PCJ?RGTCJW J?PEC QSPD?AC ?PC? AMKN?PCB RM ? KSAF J?PECP ?LGK?J. 5FC J?PEC ?LGK?Js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
1448
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RM ? AF?LEC GL RFC GLRCPL?J MP CVRCPL?J CLTGPMLKCLR PCOSGPCQ ? AF?LEC GL RFC BGPCARGML MD RFC QRGKSJSQ. " LCE?RGTC DCCB@?AI JMMN ?AAMKNJGQFCQ RFGQ, UFGJC ? NMQGRGTC DCCB@?AI JMMN UMSJB AMLRGLSC RFC QRGKSJSQ ?LB PCQSJR GL F?PK RM RFC ?LGK?J. 29 .?KK?JG?L CLXWKCQ GLAPC?QC ?ARGTGRW RM RFC NMGLR MD BCL?RSP?RGML, GLAPC?QGLE RFC AFCKGA?J ?ARGTGRW MD RFC ACJJQ GLTMJTCB. #?ARCPG?J CLXWKCQ F?TC ? QNCAGDGA RCKNCP?RSPC DMP RFCGP KMQR CDDGAGCLR ?ARGTGRW ?LB ?PC GLFG@GRCB ?R CGRFCP FGEFCP MP JMUCP RCKNCP?RSPCQ. 'CTCP PCQSJRQ GL ?L GLAPC?QC GL RFC BCQRPSARGML MD RFC GLT?BGLE @?ARCPG? @W GLAPC?QGLE RFC CDDCARGTCLCQQ MD @MBW BCDCLQCQ ?LB ?L GLFG@GRGLE @?ARCPG?J KCR?@MJGQK. 30 %G?@CRCQ GQ MDRCL ?QQMAG?RCB UGRF ? J?AI GL NPMBSARGML MD GLQSJGL. 8GRFMSR GLQSJGL, @JMMB EJSAMQC JCTCJQ EM SN ?DRCP ? KC?J, @SR LCTCP EM @?AI BMUL RM LMPK?J JCTCJQ.
Chapter 34
1 Figure 34.11 # 2 Figure 34.12 $ 3 Figure 34.19 $ 4 % 5 # 6 $ 7 $ 8 " 9 % 10 " 11 $ 12 " 13 # 14 $ 15 # 16 "LGK?JQ UGRF ? NMJWE?QRPGA BGECQRGTC QWQRCK F?TC ? KSJRG-AF?K@CPCB QRMK?AF. 5FC DMSP AMKN?PRKCLRQ MD RFC QRMK?AF ?PC A?JJCB RFC PSKCL, PCRGASJSK, MK?QSK, ?LB ?@MK?QSK. 5FCQC AF?K@CPQ AMLR?GL K?LW KGAPM@CQ RF?R @PC?I BMUL RFC ACJJSJMQC ?LB DCPKCLR RFC GLECQRCB DMMB. 5FC ?@MK?QSK GQ RFC tRPSCu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qQSAF ?Q NMR?QQGSK, QMBGSK, ?LB A?JAGSKq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
Chapter 35
1 Figure 35.3 # 2 Figure 35.11 1MR?QQGSK AF?LLCJ @JMAICPQ QJMU RFC PCNMJ?PGX?RGML NF?QC, @SR F?TC LM CDDCAR ML BCNMJ?PGX?RGML. 3 Figure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GLAPC?QCB @JMMB DJMU RM @P?GL PCEGMLQ. 24 5FC QNGL?J AMPB RP?LQKGRQ QCLQMPW GLDMPK?RGML DPMK RFC @MBW RM RFC @P?GL ?LB KMRMP AMKK?LBQ DPMK RFC @P?GL RM RFC @MBW RFPMSEF GRQ AMLLCARGMLQ UGRF NCPGNFCP?J LCPTCQ. *R ?JQM AMLRPMJQ KMRMP PCDJCVCQ. 25 5FC QWKN?RFCRGA LCPTMSQ QWQRCK NPCN?PCQ RFC @MBW DMP tDGEFR MP DJGEFR,u UFCPC?Q RFC N?P?QWKN?RFCRGA LCPTMSQ QWQRCK ?JJMUQ RFC @MBW RM tPCQR ?LB BGECQR.u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sQ BGQC?QC GLAJSBC BGQPSNRGTC KCKMPW JMQQ, AMLDSQGML ?@MSR RGKC MP NJ?AC, BGDDGASJRGCQ NJ?LLGLE MP CVCASRGLE R?QIQ, NMMP HSBEKCLR, ?LB NCPQML?JGRW AF?LECQ. 28 1MQQG@JC RPC?RKCLRQ DMP N?RGCLRQ UGRF K?HMP BCNPCQQGML GLAJSBC NQWAFMRFCP?NW ?LB NPCQAPGNRGML KCBGA?RGMLQ. ."0 GLFG@GRMP BPSEQ GLFG@GR RFC @PC?IBMUL MD ACPR?GL LCSPMRP?LQKGRRCPQ (GLAJSBGLE BMN?KGLC, QCPMRMLGL, LMPCNGLCNFPGLC) GL RFC QWL?NRGA AJCDR. 443* KCBGA?RGMLQ GLFG@GR RFC PCSNR?IC MD QCPMRMLGL GLRM RFC NPCQWL?NRGA LCSPML.
Chapter 36
1 Figure 36.5 % 2 Figure 36.14 # 3 Figure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s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
Chapter 37
1 Figure 37.5 1PMRCGLQ SLDMJB, MP BCL?RSPC, ?R FGEFCP RCKNCP?RSPCQ. 2 Figure 37.11 # 3 Figure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p80 NCPACLR MD GRQ QCAPCRGMLQ. &NGLCNFPGLC ?LB LMPCNGLCNFPGLC GLAPC?QC FC?PR P?RC, @PC?RFGLE P?RC, A?PBG?A KSQAJC AMLRP?ARGMLQ, ?LB @JMMB EJSAMQC JCTCJQ. 5FCW ?JQM ?AACJCP?RC RFC @PC?IBMUL MD EJSAMQC GL QICJCR?J KSQAJCQ ?LB QRMPCB D?RQ GL ?BGNMQC RGQQSC. 5FC PCJC?QC MD CNGLCNFPGLC ?LB LMPCNGLCNFPGLC GQ QRGKSJ?RCB @W LCSP?J GKNSJQCQ DPMK RFC QWKN?RFCRGA LCPTMSQ QWQRCK. 5FCQC LCSP?J GKNSJQCQ MPGEGL?RC DPMK RFC FWNMRF?J?KSQ GL PCQNMLQC RM QRPCQQ RM NPCN?PC RFC @MBW DMP RFC DGEFR-MP-DJGEFR PCQNMLQC.
Chapter 38
1 Figure 38.19# 2 Figure 38.37 # 3 Figure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sR LCCB ?LW DPMK RFC MSRQGBC.
Chapter 39
1 Figure 39.7 # 2 Figure 39.13 $ 3 Figure 39.20 5FC @JMMB N) UGJJ BPMN ?LB FCKMEJM@GL ?DDGLGRW DMP MVWECL UGJJ BCAPC?QC. 4 " 5 $ 6 # 7 % 8 % 9 % 10 # 11 # 12 % 13 " 14 $ 15 % 16 5FC K?GL
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@PMLAFSQ GQ RFC AMLBSGR GL RFC JSLE RF?R DSLLCJQ ?GP RM RFC ?GPU?WQ UFCPC E?Q CVAF?LEC MAASPQ. 5FC K?GL @PMLAFSQ ?RR?AFCQ RFC JSLEQ RM RFC TCPW CLB MD RFC RP?AFC? UFCPC GR @GDSPA?RCQ. 5FC RP?AFC? GQ RFC A?PRGJ?EGLMSQ QRPSARSPC RF?R CVRCLBQ DPMK RFC NF?PWLV RM RFC NPGK?PW @PMLAFG. *R QCPTCQ RM DSLLCJ ?GP RM RFC JSLEQ. 5FC ?JTCMJG ?PC RFC QGRCQ MD E?Q CVAF?LEC; RFCW ?PC JMA?RCB ?R RFC RCPKGL?J PCEGMLQ MD RFC JSLE ?LB ?PC ?RR?AFCB RM RFC PCQNGP?RMPW @PMLAFGMJCQ. 5FC ?AGLSQ GQ RFC QRPSARSPC GL RFC JSLE UFCPC E?Q CVAF?LEC MAASPQ. 17 5FC Q?A-JGIC QRPSARSPC MD RFC ?JTCMJG GLAPC?QCQ RFCGP QSPD?AC ?PC?. *L ?BBGRGML, RFC ?JTCMJG ?PC K?BC MD RFGL-U?JJCB N?PCLAFWK?J ACJJQ. 5FCQC DC?RSPCQ ?JJMU E?QCQ RM C?QGJW BGDDSQC ?APMQQ RFC ACJJQ. 18 '&71/'7$ KC?QSPCQ RFC DMPACB CVNGP?RMPW TMJSKC GL MLC QCAMLB GL PCJ?RGML RM RFC RMR?J DMPACB TGR?J A?N?AGRW (RFC RMR?J ?KMSLR MD ?GP RF?R GQ CVF?JCB DPMK RFC JSLE DPMK ? K?VGK?J GLF?J?RGML). 5FGQ P?RGM AF?LECQ UGRF ?JRCP?RGMLQ GL JSLE DSLARGML RF?R ?PGQC DPMK BGQC?QCQ QSAF ?Q DG@PMQGQ, ?QRFK?, ?LB $01%. 19 *D ?JJ RFC ?GP GL RFC JSLE UCPC CVF?JCB, RFCL MNCLGLE RFC ?JTCMJG DMP RFC LCVR GLQNGP?RGML UMSJB @C TCPW BGDDGASJR. 5FGQ GQ @CA?SQC RFC RGQQSCQ UMSJB QRGAI RMECRFCP. 20 0VWECL KMTCQ DPMK RFC JSLE RM RFC @JMMBQRPC?K RM RFC RGQQSCQ ?AAMPBGLE RM RFC NPCQQSPC EP?BGCLR. 5FGQ GQ KC?QSPCB ?Q RFC N?PRG?J NPCQQSPC MD MVWECL. *D RFC ?KMSLR MD MVWECL BPMNQ GL RFC GLQNGPCB ?GP, RFCPC UMSJB @C PCBSACB N?PRG?J NPCQQSPC. 5FGQ UMSJB BCAPC?QC RFC BPGTGLE DMPAC RF?R KMTCQ RFC MVWECL GLRM RFC @JMMB ?LB GLRM RFC RGQQSCQ. P O GQ ?JQM PCBSACB ?R FGEF CJCT?RGMLQ: P O ?R FGEF CJCT?RGMLQ GQ JMUCP RF?L ?R QC?
2 2
JCTCJ @CA?SQC RFC RMR?J ?RKMQNFCPGA NPCQQSPC GQ JCQQ RF?L ?RKMQNFCPGA NPCQQSPC ?R QC? JCTCJ. 21 " BMARMP A?L BCRCAR ? PCQRPGARGTC BGQC?QC SQGLE QNGPMKCRPW. #W BCRCARGLE RFC P?RC ?R UFGAF ?GP A?L @C CVNCJJCB DPMK RFC JSLE, ? BG?ELMQGQ MD DG@PMQGQ MP ?LMRFCP PCQRPGARGTC BGQC?QC A?L @C K?BC. 22 *LAPC?QCB ?GPU?W PCQGQR?LAC GLAPC?QCQ RFC TMJSKC ?LB NPCQQSPC GL RFC JSLE; RFCPCDMPC, RFC GLRP?NJCSP?J NPCQQSPC UMSJB @C JCQQ LCE?RGTC ?LB @PC?RFGLE UMSJB @C KMPC BGDDGASJR. 23 " NSLARSPC RM RFC RFMP?AGA A?TGRW UMSJB COS?JGXC RFC NPCQQSPC GLQGBC RFC RFMP?AGA A?TGRW RM RFC MSRQGBC CLTGPMLKCLR. 'MP RFC JSLE RM DSLARGML NPMNCPJW, RFC GLRP?NJCSP?J NPCQQSPC KSQR @C LCE?RGTC. 5FGQ GQ A?SQCB @W RFC AMLRP?ARGML MD RFC BG?NFP?EK NSJJGLE RFC JSLEQ BMUL ?LB BP?UGLE ?GP GLRM RFC JSLEQ. 24 5FC JSLE GQ N?PRGASJ?PJW QSQACNRG@JC RM AF?LECQ GL RFC K?ELGRSBC ?LB BGPCARGML MD EP?TGR?RGML?J DMPACQ. 8FCL QMKCMLC GQ QR?LBGLE MP QGRRGLE SNPGEFR, RFC NJCSP?J NPCQQSPC EP?BGCLR JC?BQ RM GLAPC?QCB TCLRGJ?RGML DSPRFCP BMUL GL RFC JSLE. 25 8GRFMSR A?P@MLGA ?LFWBP?QC, A?P@ML BGMVGBC UMSJB LMR @C FWBPMJWXCB GLRM A?P@MLGA ?AGB MP @GA?P@ML?RC. 5FCPCDMPC, TCPW JGRRJC A?P@ML BGMVGBC (MLJW 15 NCPACLR) UMSJB @C RP?LQNMPRCB GL RFC @JMMB ?U?W DPMK RFC RGQQSCQ. 26 $?P@ML KMLMVGBC F?Q ? FGEFCP ?DDGLGRW DMP FCKMEJM@GL RF?L MVWECL. 5FGQ KC?LQ RF?R A?P@ML KMLMVGBC UGJJ NPCDCPCLRG?JJW @GLB RM FCKMEJM@GL MTCP MVWECL. "BKGLGQRP?RGML MD 100 NCPACLR MVWECL GQ ?L CDDCARGTC RFCP?NW @CA?SQC ?R RF?R AMLACLRP?RGML, MVWECL UGJJ BGQNJ?AC RFC A?P@ML KMLMVGBC DPMK RFC FCKMEJM@GL.
Chapter 40
1 Figure 40.10 $ 2 Figure 40.11 # 3 Figure 40.17 #JMMB GL RFC JCEQ GQ D?PRFCQR ?U?W DPMK RFC FC?PR ?LB F?Q RM DJMU SN RM PC?AF GR. 4 " 5 % 6 $ 7 % 8 $ 9 # 10 $ 11 # 12 " 13 % 14 " 15 " 16 " AJMQCB AGPASJ?RMPW QWQRCK GQ ? AJMQCB-JMMN QWQRCK, GL UFGAF @JMMB GQ LMR DPCC GL ? A?TGRW. #JMMB GQ QCN?P?RC DPMK RFC @MBGJW GLRCPQRGRG?J DJSGB ?LB AMLR?GLCB UGRFGL @JMMB TCQQCJQ. *L RFGQ RWNC MD QWQRCK, @JMMB AGPASJ?RCQ SLGBGPCARGML?JJW DPMK RFC FC?PR ?PMSLB RFC QWQRCKGA AGPASJ?RMPW PMSRC, ?LB RFCL PCRSPLQ RM RFC FC?PR. 17 4WQRCKGA AGPASJ?RGML DJMUQ RFPMSEF RFC QWQRCKQ MD RFC @MBW. 5FC @JMMB DJMUQ ?U?W DPMK RFC FC?PR RM RFC @P?GL, JGTCP, IGBLCWQ, QRMK?AF, ?LB MRFCP MPE?LQ, RFC JGK@Q, ?LB RFC KSQAJCQ MD RFC @MBW; GR RFCL PCRSPLQ RM RFC FC?PR. 18 3CB @JMMB ACJJQ ?PC AM?RCB UGRF NPMRCGLQ A?JJCB ?LRGECLQ K?BC MD EJWAMJGNGBQ ?LB EJWAMNPMRCGLQ. 8FCL RWNC " ?LB RWNC # @JMMB ?PC KGVCB, RFC @JMMB ?EEJSRGL?RCQ @CA?SQC MD ?LRG@MBGCQ GL RFC NJ?QK? RF?R @GLB UGRF RFC MNNMQGLE ?LRGECL. 5WNC 0 @JMMB F?Q LM ?LRGECLQ. 5FC 3F @JMMB EPMSN F?Q CGRFCP RFC 3F ?LRGECL (3F+) MP LM 3F ?LRGECL (3Fp). 19 #JMMB GQ GKNMPR?LR DMP PCESJ?RGML MD RFC @MBWs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
Chapter 41
1 Figure 41.5 $ 2 Figure 41.6 " 3 Figure 41.8 -MMN BGSPCRGAQ BCAPC?QC RFC CVAPCRGML MD Q?JR GLRM RFC PCL?J KCBSJJ?, RFCPC@W PCBSAGLE GRQ MQKMJ?JGRW. "Q ? PCQSJR, JCQQ U?RCP GQ CVAPCRCB GLRM RFC KCBSJJ? @W RFC BCQACLBGLE JGK@, ?LB KMPC U?RCP GQ CVAPCRCB ?Q SPGLC. 4 # 5 # 6 " 7 $ 8 # 9 " 10 $ 11 % 12 % 13 " 14 $ 15 " 16 $ 17 " 18 &VAPCRGML ?JJMUQ ?L MPE?LGQK RM PGB GRQCJD MD U?QRC KMJCASJCQ RF?R AMSJB @C RMVGA
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GD ?JJMUCB RM ?AASKSJ?RC. *R ?JQM ?JJMUQ RFC MPE?LGQK RM ICCN RFC ?KMSLR MD U?RCP ?LB BGQQMJTCB QMJSRCQ GL @?J?LAC. 19 &JCARPMJWRC GMLQ MDRCL PCOSGPC QNCAG?J KCAF?LGQKQ RM APMQQ RFC QCKG-NCPKC?@JC KCK@P?LCQ GL RFC @MBW. "ARGTC RP?LQNMPR GQ RFC KMTCKCLR ?E?GLQR ? AMLACLRP?RGML EP?BGCLR. 20 5FC JMMN MD )CLJC GQ N?PR MD RFC PCL?J RS@SJC RF?R JMMNQ GLRM RFC PCL?J KCBSJJ?. *L RFC JMMN MD )CLJC, RFC DGJRP?RC CVAF?LECQ QMJSRCQ ?LB U?RCP UGRF RFC PCL?J KCBSJJ? ?LB RFC T?Q? PCAR? (RFC NCPGRS@SJ?P A?NGJJ?PW LCRUMPI). 5FC T?Q? PCAR? ?ARQ ?Q RFC AMSLRCPASPPCLR CVAF?LECP. 5FC IGBLCWQ K?GLR?GL RFC MQKMJ?JGRW MD RFC PCQR MD RFC @MBW ?R ? AMLQR?LR 300 K0QK @W AMLACLRP?RGLE RFC DGJRP?RC ?Q GR N?QQCQ RFPMSEF RFC JMMN MD )CLJC. 21 &VRCPL?JJW, RFC IGBLCWQ ?PC QSPPMSLBCB @W RFPCC J?WCPQ. 5FC MSRCPKMQR J?WCP GQ ? RMSEF AMLLCARGTC RGQQSC J?WCP A?JJCB RFC PCL?J D?QAG?. 5FC QCAMLB J?WCP GQ A?JJCB RFC NCPGPCL?J D?R A?NQSJC, UFGAF FCJNQ ?LAFMP RFC IGBLCWQ GL NJ?AC. 5FC RFGPB ?LB GLLCPKMQR J?WCP GQ RFC PCL?J A?NQSJC. *LRCPL?JJW, RFC IGBLCW F?Q RFPCC PCEGMLQq?L MSRCP AMPRCV, ? KCBSJJ? GL RFC KGBBJC, ?LB RFC PCL?J NCJTGQ GL RFC PCEGML A?JJCB RFC FGJSK MD RFC IGBLCW, UFGAF GQ RFC AMLA?TC N?PR MD RFC t@C?Lu QF?NC. 22 5FC PCKMT?J MD U?QRCQ, UFGAF AMSJB MRFCPUGQC @C RMVGA RM ?L MPE?LGQK, GQ CVRPCKCJW GKNMPR?LR DMP QSPTGT?J. )?TGLE MPE?LQ RF?R QNCAG?JGXC GL RFGQ NPMACQQ ?LB RF?R MNCP?RC QCN?P?RCJW DPMK MRFCP MPE?LQ NPMTGBCQ ? KC?QSPC MD Q?DCRW DMP RFC MPE?LGQK. 23 (1) .GAPMMPE?LGQKQ CLESJD DMMB @W CLBMAWRMQGQq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
Chapter 42
1 Figure 42.11 $ 2 Figure 42.14 .)$ PCACNRMPQ BGDDCP DPMK NCPQML RM NCPQML. 5FSQ, .)$ PCACNRMPQ ML ?L GLAMKN?RG@JC BMLMP ?PC AMLQGBCPCB tLML-QCJDu ?LB ?PC PCHCARCB @W RFC GKKSLC QWQRCK. 3 Figure 42.16 *D RFC @JMMB MD RFC KMRFCP ?LB DCRSQ KGVCQ, KCKMPW ACJJQ RF?R PCAMELGXC RFC 3F ?LRGECL A?L DMPK J?RC GL RFC DGPQR NPCEL?LAW. %SPGLE QS@QCOSCLR NPCEL?LAGCQ, RFCQC KCKMPW ACJJQ J?SLAF ?L GKKSLC ?RR?AI ML RFC DCR?J @JMMB ACJJQ. *LHCARGML MD ?LRG-3F ?LRG@MBW BSPGLE RFC DGPQR NPCEL?LAW NPCTCLRQ RFC GKKSLC PCQNMLQC DPMK MAASPPGLE. 4 % 5 # 6 " 7 $ 8 % 9 # 10 # 11 $ 12 % 13 " 14 $ 15 " 16 $ 17 # 18 % 19 # 20 $ 21 % 22 *D RFC .)$ * KMJCASJCQ CVNPCQQCB ML BMLMP ACJJQ BGDDCP DPMK RFC .)$ * KMJCASJCQ CVNPCQQCB ML PCAGNGCLR ACJJQ, /, ACJJQ K?W GBCLRGDW RFC BMLMP ACJJQ ?Q tLML-QCJDu ?LB NPMBSAC NCPDMPGL ?LB EP?LXWKCQ RM GLBSAC RFC BMLMP ACJJQ RM SLBCPEM ?NMNRMQGQ, UFGAF UMSJB BCQRPMW RFC RP?LQNJ?LRCB MPE?L. 23 5FC CLRGPC AMKNJCKCLR QWQRCK UMSJB NPM@?@JW @C ?DDCARCB CTCL UFCL MLJW ? DCU KCK@CPQ UCPC KSR?RCB QSAF RF?R RFCW AMSJB LM JMLECP @GLB. #CA?SQC RFC AMKNJCKCLR GLTMJTCQ RFC @GLBGLE MD ?ARGT?RCB NPMRCGLQ GL ? QNCAGDGA QCOSCLAC, UFCL MLC MP KMPC NPMRCGLQ GL RFC QCOSCLAC ?PC ?@QCLR, RFC QS@QCOSCLR NPMRCGLQ UMSJB @C GLA?N?@JC MD @GLBGLE RM CJGAGR RFC AMKNJCKCLRsQ N?RFMECL-BCQRPSARGTC CDDCARQ. 24 "L ?LRGECL GQ ? KMJCASJC RF?R PC?ARQ UGRF QMKC AMKNMLCLR MD RFC GKKSLC PCQNMLQC (?LRG@MBW, # ACJJ PCACNRMP, 5 ACJJ PCACNRMP). "L CNGRMNC GQ RFC PCEGML ML RFC ?LRGECL RFPMSEF UFGAF @GLBGLE UGRF RFC GKKSLC AMKNMLCLR ?ARS?JJW MAASPQ. 25 " L?bTC 5 MP # ACJJ GQ MLC RF?R F?Q LMR @CCL ?ARGT?RCB @W @GLBGLE RM RFC ?NNPMNPG?RC CNGRMNC. /?bTC 5 ?LB # ACJJQ A?LLMR NPMBSAC
!00%.$)8 ! X !00%.$)8
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PCQNMLQCQ. 26 5FC 5)1 PCQNMLQC GLTMJTCQ RFC QCAPCRGML MD AWRMIGLCQ RM QRGKSJ?RC K?APMNF?ECQ ?LB $5-Q ?LB GKNPMTC RFCGP BCQRPSARGML MD GLRP?ACJJSJ?P N?RFMECLQ ?LB RSKMP ACJJQ. *R GQ ?QQMAG?RCB UGRF GLDJ?KK?RGML. 5FC 5)2 PCQNMLQC GQ GLTMJTCB GL RFC QRGKSJ?RGML MD # ACJJQ GLRM NJ?QK? ACJJQ RF?R QWLRFCQGXC ?LB QCAPCRC ?LRG@MBGCQ. 27 5FC BGTCPQGRW MD 5$3Q ?JJMUQ RFC GKKSLC QWQRCK RM F?TC KGJJGMLQ MD BGDDCPCLR 5 ACJJQ, ?LB RFCPC@W RM @C QNCAGDGA GL BGQRGLESGQFGLE ?LRGECLQ. 5FGQ BGTCPQGRW ?PGQCQ DPMK KSR?RGML ?LB PCAMK@GL?RGML GL RFC ECLCQ RF?R CLAMBC RFC T?PG?@JC PCEGMLQ MD 5$3Q. 28 5 ACJJQ @GLB ?LRGECLQ RF?R F?TC @CCL BGECQRCB ?LB CK@CBBCB GL .)$ KMJCASJCQ @W "1$Q. *L AMLRP?QR, # ACJJQ DSLARGML RFCKQCJTCQ ?Q "1$Q RM @GLB GLR?AR, SLNPMACQQCB ?LRGECLQ. 29 6NML PCGLDCARGML, RFC KCKMPW ACJJQ UGJJ GKKCBG?RCJW BGDDCPCLRG?RC GLRM NJ?QK? ACJJQ ?LB $5-Q UGRFMSR GLNSR DPMK "1$Q MP 5) ACJJQ. *L AMLRP?QR, RFC ?B?NRGTC GKKSLC PCQNMLQC RM RFC GLGRG?J GLDCARGML PCOSGPCQ RGKC DMP L?bTC # ?LB 5 ACJJQ UGRF RFC ?NNPMNPG?RC ?LRGECL QNCAGDGAGRGCQ RM @C GBCLRGDGCB ?LB ?ARGT?RCB. 30 $PMQQ PC?ARGTGRW MD ?LRG@MBGCQ A?L @C @CLCDGAG?J UFCL GR ?JJMUQ ?L GLBGTGBS?J'Q GKKSLC QWQRCK RM PCQNMLB RM ?L ?PP?W MD QGKGJ?P N?RFMECLQ ?DRCP @CGLE CVNMQCB RM HSQR MLC MD RFCK. " NMRCLRG?J AMQR MD APMQQ PC?ARGTGRW GQ ?L ?LRG@MBW PCQNMLQC RM N?PRQ MD RFC @MBW (QCJD) GL ?BBGRGML RM RFC ?NNPMNPG?RC ?LRGECL.
Chapter 43
1 Figure 43.8 % 2 Figure 43.15 $ 3 Figure 43.17 # 4 " 5 # 6 % 7 " 8 " 9 $ 10 " 11 $ 12 $ 13 " 14 " 15 % 16 " 17 " 18 $ 19 # 20 % 21 # 22 " 23 % 24 # 25 " 26 # 27 # 28 % 29 " 30 % 31 4CVS?J PCNPMBSARGML NPMBSACQ ? LCU AMK@GL?RGML MD ECLCQ GL RFC MDDQNPGLE RF?R K?W @CRRCP CL?@JC RFCK RM QSPTGTC AF?LECQ GL RFC CLTGPMLKCLR ?LB ?QQGQR GL RFC QSPTGT?J MD RFC QNCAGCQ. 32 5FC NPCQCLAC MD RFC 8 AFPMKMQMKC GL @GPBQ BCRCPKGLCQ DCK?JCLCQQ ?LB RFC NPCQCLAC MD RFC : AFPMKMQMKC GL K?KK?JQ BCRCPKGLCQ K?JCLCQQ. 5FC ?@QCLAC MD RFMQC AFPMKMQMKCQ ?LB RFC FMKMECLCGRW MD RFC MDDQNPGLE (;; MP 99) JC?BQ RM RFC BCTCJMNKCLR MD RFC MRFCP QCV. 33 &VRCPL?J DCPRGJGX?RGML A?L APC?RC J?PEC LSK@CPQ MD MDDQNPGLE UGRFMSR PCOSGPGLE QNCAG?JGXCB BCJGTCPW MP PCNPMBSARGTC QSNNMPR MPE?LQ. 0DDQNPGLE BCTCJMN ?LB K?RSPC OSGAIJW AMKN?PCB RM GLRCPL?JJW DCPRGJGXGLE QNCAGCQ. " BGQ?BT?LR?EC GQ RF?R RFC MDDQNPGLE ?PC MSR GL RFC CLTGPMLKCLR ?LB NPCB?RGML A?L ?AAMSLR DMP J?PEC JMQQ MD MDDQNPGLE. 5FC CK@PWMQ ?PC QSQACNRG@JC RM AF?LECQ GL RFC CLTGPMLKCLR, UFGAF DSPRFCP BCNJCRCQ RFCGP LSK@CPQ. *LRCPL?JJW DCPRGJGXGLE QNCAGCQ AMLRPMJ RFCGP CLTGPMLKCLR ?LB NPMRCAR RFCGP MDDQNPGLE DPMK NPCB?RMPQ @SR KSQR F?TC QNCAG?JGXCB MPE?LQ RM AMKNJCRC RFCQC R?QIQ ?LB SQS?JJW NPMBSAC DCUCP CK@PWMQ. 34 1?GPCB CVRCPL?J DCPRGJGX?RGML ?JJMUQ RFC DCK?JC RM QCJCAR RFC K?JC DMP K?RGLE. *R ?JQM F?Q ? EPC?RCP AF?LAC MD DCPRGJGX?RGML R?IGLE NJ?AC, UFCPC?Q QN?ULGLE HSQR NSRQ ? J?PEC LSK@CP MD QNCPK ?LB CEEQ RMECRFCP ?LB P?LBMK GLRCP?ARGMLQ PCQSJR GL RFC DCPRGJGX?RGML. 35 *L NF?QC MLC (CVAGRCKCLR), T?QMBGJ?RGML JC?BQ RM T?QMAMLECQRGML ?LB CLJ?PECKCLR MD CPCARGJC RGQQSCQ. 7?EGL?J QCAPCRGMLQ ?PC PCJC?QCB RM JS@PGA?RC RFC T?EGL? BSPGLE GLRCPAMSPQC. *L NF?QC RUM (NJ?RC?S), QRGKSJ?RGML AMLRGLSCQ, RFC MSRCP RFGPB MD RFC T?EGL?J U?JJ CLJ?PECQ UGRF @JMMB, ?LB @PC?RFGLE ?LB FC?PR P?RC GLAPC?QC. *L NF?QC RFPCC (MPE?QK), PFWRFKGA, GLTMJSLR?PW AMLRP?ARGMLQ MD KSQAJCQ MAASP. *L RFC K?JC, PCNPMBSARGTC ?AACQQMPW EJ?LBQ ?LB RS@SJCQ AMLQRPGAR, BCNMQGRGLE QCKCL GL RFC SPCRFP?; RFCL, RFC SPCRFP? AMLRP?ARQ, CVNCJJGLE RFC QCKCL RFPMSEF RFC NCLGQ. *L UMKCL, RFC SRCPSQ ?LB T?EGL?J KSQAJCQ AMLRP?AR GL U?TCQ RF?R K?W J?QR QJGEFRJW JCQQ RF?L ? QCAMLB C?AF. *L NF?QC DMSP (PCQMJSRGML), RFC NPMACQQCQ JGQRCB GL RFC DGPQR RFPCC NF?QCQ PCTCPQC RFCKQCJTCQ ?LB PCRSPL RM RFCGP LMPK?J QR?RC. .CL CVNCPGCLAC ? PCDP?ARMPW NCPGMB GL UFGAF RFCW A?LLMR K?GLR?GL ?L CPCARGML MP CH?ASJ?RC DMP ? NCPGMB MD RGKC P?LEGLE DPMK KGLSRCQ RM FMSPQ. 8MKCL BM LMR CVNCPGCLAC ? PCDP?ARMPW NCPGMB. 36 4RCK ACJJQ ?PC J?GB BMUL GL RFC K?JC BSPGLE ECQR?RGML ?LB JGC BMPK?LR SLRGJ ?BMJCQACLAC. 4RCK ACJJQ GL RFC DCK?JC GLAPC?QC RM MLC RM RUM KGJJGML ?LB CLRCP RFC DGPQR KCGMRGA BGTGQGML ?LB ?PC ?PPCQRCB GL NPMNF?QC. "R ?BMJCQACLAC, QNCPK?RMECLCQGQ @CEGLQ ?LB AMLRGLSCQ SLRGJ BC?RF, NPMBSAGLE RFC K?VGKSK LSK@CP MD QNCPK UGRF C?AF KCGMRGA BGTGQGML. 0MECLCQGQ AMLRGLSCQ ?E?GL ?R ?BMJCQACLAC GL @?RAFCQ MD MMEMLG? UGRF C?AF KCLQRPS?J AWAJC. 5FCQC MMEMLG? DGLGQF RFC DGPQR KCGMRGA BGTGQGML, NPMBSAGLE ? NPGK?PW MMAWRC UGRF KMQR MD RFC AWRMNJ?QK ?LB GRQ AMLRCLRQ, ?LB ? QCAMLB ACJJ A?JJCB ? NMJ?P @MBW AMLR?GLGLE 23 AFPMKMQMKCQ. 5FC QCAMLB KCGMRGA BGTGQGML PCQSJRQ GL ? QCAMLB?PW MMAWRC ?LB ? QCAMLB MMAWRC. "R MTSJ?RGML, ? K?RSPC F?NJMGB CEE GQ PCJC?QCB. *D RFGQ CEE GQ DCPRGJGXCB, GR DGLGQFCQ RFC QCAMLB KCGMRGA BGTGQGML, GLAJSBGLE RFC AFPMKMQMKCQ BML?RCB @W RFC QNCPK GL RFC DGLGQFCB ACJJ. 5FGQ GQ ? BGNJMGB, DCPRGJGXCB CEE. 37 /CE?RGTC DCCB@?AI GL RFC K?JC QWQRCK GQ QSNNJGCB RFPMSEF RUM FMPKMLCQ: GLFG@GL ?LB RCQRMQRCPMLC. *LFG@GL GQ NPMBSACB @W 4CPRMJG ACJJQ UFCL RFC QNCPK AMSLR CVACCBQ QCR JGKGRQ. 5FC FMPKMLC GLFG@GRQ (L3) ?LB '4), BCAPC?QGLE RFC ?ARGTGRW MD RFC 4CPRMJG ACJJQ. *LAPC?QCB JCTCJQ MD RCQRMQRCPMLC ?DDCAR RFC PCJC?QC MD @MRF (L3) ?LB -), BCAPC?QGLE RFC ?ARGTGRW MD RFC -CWBGE ACJJQ, PCQSJRGLE GL BCAPC?QCB RCQRMQRCPMLC ?LB QNCPK NPMBSARGML. 38 -MU JCTCJQ MD NPMECQRCPMLC ?JJMU RFC FWNMRF?J?KSQ RM QCLB (L3) RM RFC ?LRCPGMP NGRSGR?PW ?LB A?SQC RFC PCJC?QC MD '4) ?LB -). '4) QRGKSJ?RCQ DMJJGAJCQ ML RFC MT?PW RM EPMU ?LB NPCN?PC RFC CEEQ DMP MTSJ?RGML. "Q RFC DMJJGAJCQ GLAPC?QC GL QGXC, RFCW @CEGL RM PCJC?QC CQRPMECL ?LB ? JMU JCTCJ MD NPMECQRCPMLC GLRM RFC @JMMB. 5FC JCTCJ MD CQRPMECL PGQCQ RM ? NC?I, A?SQGLE ? QNGIC GL RFC AMLACLRP?RGML MD -). 5FGQ A?SQCQ RFC KMQR K?RSPC DMJJGAJC RM PSNRSPC ?LB MTSJ?RGML MAASPQ. 39 5FC DGPQR RPGKCQRCP J?WQ BMUL RFC @?QGA QRPSARSPCQ MD RFC @MBW, GLAJSBGLE RFC JGK@ @SBQ, FC?PR, CWCQ, ?LB RFC JGTCP. 5FC QCAMLB RPGKCQRCP AMLRGLSCQ RFC BCTCJMNKCLR MD ?JJ MD RFC MPE?LQ ?LB QWQRCKQ CQR?@JGQFCB BSPGLE RFC DGPQR RPGKCQRCP. 5FC NJ?ACLR? R?ICQ MTCP RFC NPMBSARGML MD CQRPMECL ?LB FGEF JCTCJQ MD NPMECQRCPMLC ?LB F?LBJCQ RFC LSRPGCLR ?LB U?QRC PCOSGPCKCLRQ MD RFC DCRSQ. 5FC RFGPB RPGKCQRCP CVFG@GRQ RFC EPC?RCQR EPMURF MD RFC DCRSQ, ASJKGL?RGLE GL J?@MP ?LB BCJGTCPW. 40 4R?EC MLC MD J?@MP PCQSJRQ GL RFC RFGLLGLE MD RFC ACPTGV ?LB RFC BGJ?RGML MD RFC ACPTGA?J MNCLGLE. 4R?EC RUM BCJGTCPQ RFC @?@W, ?LB QR?EC RFPCC BCJGTCPQ RFC NJ?ACLR?. 41 .SJRGNJC QNCPK A?L DSQC UGRF RFC CEE, PCQSJRGLE GL NMJWQNCPKW. 5FC PCQSJRGLE CK@PWM GQ LMR ECLCRGA?JJW TG?@JC ?LB BGCQ UGRFGL ? DCU B?WQ. 42 .?KK?JG?L CEEQ BM LMR LCCB ? JMR MD WMJI @CA?SQC RFC BCTCJMNGLE DCRSQ M@R?GLQ LSRPGCLRQ DPMK RFC KMRFCP. 0RFCP QNCAGCQ, GL UFGAF RFC DCRSQ
1454
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BCTCJMNQ MSRQGBC MD RFC KMRFCPs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
Chapter 44
1 Figure 44.10 5PMNGA?J J?ICQ BMLsR DPCCXC, QM RFCW BMLsR SLBCPEM QNPGLE RSPLMTCP GL RFC Q?KC U?W RCKNCP?RC J?ICQ BM. )MUCTCP, QRP?RGDGA?RGML BMCQ MAASP, ?Q UCJJ ?Q QC?QML?J RSPLMTCP. 2 Figure 44.12 $. #MPC?J DMPCQRQ ?PC LMR BMKGL?RCB @W BCAGBSMSQ RPCCQ. 3 Figure 44.21 $. 1FMRMQWLRFCRGA MPE?LGQKQ UMSJB @C DMSLB GL RFC NFMRGA, ?@WQQ?J, LCPGRGA, ?LB MAC?LGA XMLCQ. 4 # 5 % 6 % 7 $ 8 % 9 $ 10 % 11 # 12 $ 13 # 14 &AMJMEGQRQ UMPIGLE GL MPE?LGQK?J MP NMNSJ?RGML CAMJMEW KGEFR ?QI QGKGJ?P OSCQRGMLQ ?@MSR FMU RFC @GMRGA ?LB ?@GMRGA AMLBGRGMLQ ?DDCAR N?PRGASJ?P MPE?LGQKQ ?LB, RFSQ, KGEFR DGLB AMJJ?@MP?RGML RM @C KSRS?JJW @CLCDGAG?J. -CTCJQ MD CAMJMEW QSAF ?Q AMKKSLGRW CAMJMEW MP CAMQWQRCK CAMJMEW KGEFR NMQC EPC?RCP AF?JJCLECQ DMP AMJJ?@MP?RGML @CA?SQC RFCQC ?PC?Q ?PC TCPW @PM?B ?LB K?W GLAJSBC K?LW BGDDCPCLR CLTGPMLKCLR?J AMKNMLCLRQ. 15 *R GQ @CLCDGAG?J RM AMLQGBCP ? NMNSJ?RGML RM @C ?JJ MD RFC GLBGTGBS?JQ JGTGLE GL RFC Q?KC ?PC? ?R RFC Q?KC RGKC @CA?SQC GR ?JJMUQ RFC CAMJMEGQR RM GBCLRGDW ?LB QRSBW ?JJ MD RFC ?@GMRGA ?LB @GMRGA D?ARMPQ RF?R K?W ?DDCAR RFC KCK@CPQ MD RFC NMNSJ?RGML. )MUCTCP, RFGQ BCDGLGRGML MD ? NMNSJ?RGML AMSJB @C AMLQGBCPCB ? BP?U@?AI GD GR NPMFG@GRQ RFC CAMJMEGQR DPMK QRSBWGLE ? NMNSJ?RGMLs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
Chapter 45
1 Figure 45.2 4K?JJCP ?LGK?JQ PCOSGPC JCQQ DMMB ?LB MRFCP PCQMSPACQ, QM RFC CLTGPMLKCLR A?L QSNNMPR KMPC MD RFCK. 2 Figure 45.10b " 3 Figure 45.16 4R?EC 4 PCNPCQCLRQ ? NMNSJ?RGML RF?R GQ BCAPC?QGLE. 4 $ 5 % 6 " 7 " 8 # 9 % 10 " 11 $ 12 # 13 " 14 # 15 % 16 % 17 # 18 # 19 % 20 # 21 $ 22 % 23 $ 24 # 25 5FC PCQC?PAFCP UMSJB K?PI ? ACPR?GL LSK@CP MD NCLESGLQ UGRF ? R?E, PCJC?QC RFCK @?AI GLRM RFC NMNSJ?RGML, ?LB, ?R ? J?RCP RGKC, PCA?NRSPC NCLESGLQ RM QCC UF?R NCPACLR?EC MD RFC PCA?NRSPCB NCLESGLQ U?Q R?EECB. 5FGQ NCPACLR?EC UMSJB ?JJMU ?L CQRGK?RGML MD RFC QGXC MD RFC NCLESGL NMNSJ?RGML. 26 1?PCLR?J A?PC GQ LMR DC?QG@JC DMP MPE?LGQKQ F?TGLE K?LW MDDQNPGLE @CA?SQC RFCW BM LMR F?TC RFC CLCPEW ?T?GJ?@JC RM R?IC A?PC MD MDDQNPGLE. .MQR MD RFCGP CLCPEW @SBECR GQ SQCB GL RFC DMPK?RGML MD QCCBQ MP MDDQNPGLE, QM RFCPC GQ JGRRJC JCDR DMP N?PCLR?J A?PC. "JQM, RFC QFCCP LSK@CP MD MDDQNPGLE UMSJB K?IC GLBGTGBS?J N?PCLR?J A?PC GKNMQQG@JC. 27 *L RFC DGPQR N?PR MD RFC ASPTC, UFCL DCU GLBGTGBS?JQ MD RFC QNCAGCQ ?PC NPCQCLR ?LB PCQMSPACQ ?PC NJCLRGDSJ, EPMURF GQ CVNMLCLRG?J, QGKGJ?P RM ? +-QF?NCB ASPTC. -?RCP, EPMURF QJMUQ BSC RM RFC QNCAGCQ SQGLE SN PCQMSPACQ. 'GL?JJW, RFC NMNSJ?RGML JCTCJQ MDD ?R RFC A?PPWGLE A?N?AGRW MD RFC CLTGPMLKCLR, ?LB GR GQ PCJ?RGTCJW QR?@JC MTCP RGKC. 28 *D ? L?RSP?J BGQ?QRCP QSAF ?Q ? DGPC F?NNCLCB GL RFC UGLRCP, UFCL NMNSJ?RGMLQ ?PC JMU, GR UMSJB F?TC ? EPC?RCP CDDCAR ML RFC MTCP?JJ NMNSJ?RGML ?LB GRQ PCAMTCPW RF?L GD RFC Q?KC BGQ?QRCP MAASPPCB BSPGLE RFC QSKKCP,
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UFCL NMNSJ?RGML JCTCJQ ?PC FGEF. 29 3?NGBJW EPMUGLE AMSLRPGCQ F?TC ? J?PEC QCEKCLR MD RFC NMNSJ?RGML ?R ? PCNPMBSARGTC ?EC MP WMSLECP. 4JMUCP EPMUGLE NMNSJ?RGMLQ F?TC ? JMUCP NCPACLR?EC MD RFCQC GLBGTGBS?JQ, ?LB AMSLRPGCQ UGRF XCPM NMNSJ?RGML EPMURF F?TC ?L CTCL JMUCP NCPACLR?EC. 0L RFC MRFCP F?LB, ? FGEF NPMNMPRGML MD MJBCP GLBGTGBS?JQ GQ QCCL KMQRJW GL AMSLRPGCQ UGRF XCPM EPMURF, ?LB ? JMU NPMNMPRGML GQ KMQR AMKKML GL P?NGBJW EPMUGLE AMSLRPGCQ. 30 5FC AMKNCRGRGTC CVAJSQGML NPGLAGNJC QR?RCQ RF?R LM RUM QNCAGCQ AMKNCRGLE DMP RFC Q?KC PCQMSPACQ ?R RFC Q?KC RGKC ?LB NJ?AC A?L AMCVGQR MTCP RGKC. 5FSQ, MLC MD RFC AMKNCRGLE QNCAGCQ UGJJ CTCLRS?JJW BMKGL?RC. 0L RFC MRFCP F?LB, GD RFC QNCAGCQ CTMJTC QSAF RF?R RFCW SQC PCQMSPACQ DPMK BGDDCPCLR N?PRQ MD RFC F?@GR?R MP ?R BGDDCPCLR RGKCQ MD B?W, RFC RUM QNCAGCQ A?L CVGQR RMECRFCP GLBCDGLGRCJW. 31 %MEQ Q?JGT?RCB GL PCQNMLQC RM DMMB. 5FGQ U?Q RFC SLAMLBGRGMLCB QRGKSJSQ ?LB PCQNMLQC. %MEQ CVNMQCB RM DMMB F?B ? @CJJ PSLE PCNC?RCBJW ?R RFC Q?KC RGKC, CTCLRS?JJW JC?PLGLE RM ?QQMAG?RC RFC @CJJ UGRF DMMB. 0TCP RGKC, RFC BMEQ UMSJB Q?JGT?RC UFCL RFC @CJJ U?Q PSLE, CTCL GL RFC ?@QCLAC MD DMMB. 5FSQ, RFC @CJJ @CA?KC RFC AMLBGRGMLCB QRGKSJSQ, ?LB RFC Q?JGT?RGML GL PCQNMLQC RM RFC @CJJ @CA?KC RFC AMLBGRGMLCB PCQNMLQC.
Chapter 46
1 Figure 46.8 "AAMPBGLE RM RFC DGPQR J?U MD RFCPKMBWL?KGAQ, CLCPEW A?L LCGRFCP @C APC?RCB LMP BCQRPMWCB. &TCLRS?JJW, ?JJ CLCPEW AMLQSKCB @W JGTGLE QWQRCKQ GQ JMQR ?Q FC?R MP SQCB DMP PCQNGP?RGML, ?LB RFC RMR?J CLCPEW MSRNSR MD RFC QWQRCK KSQR COS?J RFC CLCPEW RF?R UCLR GLRM GR. 2 Figure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igure 46.17 $: /GRPGDGA?RGML @W @?ARCPG? AMLTCPRQ LGRP?RCQ (/03y) RM LGRPGRCQ (/02y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sQ U?RCP QSNNJGCQ ?PC RFPC?RCLCB @W NMJJSRGML ?LB CVF?SQRGML. 5FC CDDMPR RM QSNNJW DPCQF BPGLIGLE U?RCP RM RFC NJ?LCRsQ CTCP-CVN?LBGLE FSK?L NMNSJ?RGML GQ QCCL ?Q ? K?HMP AF?JJCLEC GL RFGQ ACLRSPW.
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Chapter 47
1 Figure 47.6 ". "L ?@SLB?LAC MD DCPL QNMPCQ DPMK QCTCP?J QNCAGCQ U?Q DMSLB @CJMU RFC ,-1E @MSLB?PW, @SR LMLC U?Q DMSLB ?@MTC. 2 Figure 47.9 5FC EPMSLB GQ NCPK?LCLRJW DPMXCL QM RFC QCCBQ UGJJ ICCN CTCL GD RFC CJCARPGAGRW D?GJQ. 3 # 4 Figure 47.16 $ 5 $ 6 " 7 $ 8 % 9 # 10 $ 11 % 12 $ 13 $ 14 $ 15 % 16 $ 17 5FC FWNMRFCQGXCB A?SQC MD RFC ,p1E CVRGLARGML CTCLR GQ ?L ?QRCPMGB GKN?AR. 5FC DGPQR NGCAC MD CTGBCLAC MD RFC GKN?AR GQ ? QNGIC GL GPGBGSK (?L CJCKCLR RF?R GQ P?PC ML &?PRF, @SR AMKKML GL KCRCMPQ) GL RFC ECMJMEGA?J J?WCPQ RF?R K?PI RFC ,p1E RP?LQGRGML. 5FC QCAMLB NGCAC MD CTGBCLAC GQ ?L GKN?AR AP?RCP MDD RFC :SA?Ra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1460
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1461
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1462
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1471
KBRO>I QR?B, 1275, 1278 KBROL?FLILDV, 32, 35 KBROLABDBKBO>QFSB AFPLOABO, 1026 .BROLABDBKBO>QFSB AFPLOABOP, 1018 KBROLK, 1026 KBROLKP, 991 KBROLQO>KPJFQQBO, 268 KBROLQO>KPJFQQBOP, 247 KBRQOLK, 41, 67 KBRQOLMEFI, 1217, 1242 KBUQ-DBKBO>QFLK PBNRBK@FKD, 463, 471 .FQOFCF@>QFLK, 590 KFQOFCF@>QFLK, 604 KFQOLDBK CFU>QFLK, 590, 604 KFQOLDBK>PB, 893, 897 KL?IB D>P, 67 KL?IB D>PBP, 46 KL@F@BMQFLK, 1039, 1058 KLAB, 877 .LABP, 842 KLABP LC 2>KSFBO, 991, 1026 KLARIB, 604 KLARIBP, 598, 893, 897 KLK-BIB@QOLIVQB, 1190, 1208 KLK-BKALPMBOJF@ AF@LQ, 929 KLK-BKALPMBOJF@ AF@LQP, 919 KLK-OBKBT>?IB OBPLRO@B, 1385, 1393 KLK-S>P@RI>O MI>KQ, 695 KLK-S>P@RI>O MI>KQP, 679 KLKAFPGRK@QFLK, 360, 367 KLKM>OBKQ>I (OB@LJ?FK>KQ) QVMB, 367 KLKM>OBKQ>I QVMBP, 355 KLKMLI>O @LS>IBKQ ?LKA, 68 .LKMLI>O @LS>IBKQ ?LKAP, 51 KLKO>KALJ J>QFKD, 510, 517 KLKPBKPB @LALK, 418 KLKPBKPB @LALKP, 400 KLKQBJMI>QB PQO>KA, 402, 418 KLOBMFKBMEOFKB, 1016, 1026, 1078, 1089 KLOQEBOK ?ILQQFKD, 452, 471 KLQL@ELOA, 798, 834 KR@IB>O BKSBILMB, 118, 137 KR@IBF@ >@FA, 103 .R@IBF@ >@FAP, 98 KR@IBLFA, 113, 137 KR@IBLIRP, 119, 137 KR@IBLMI>PJ, 118, 137 KR@IBLPLJB, 275, 295 KR@IBLQFAB, 103
KR@IBLQFAB BU@FPFLK OBM>FO, 389, 392 KR@IBLQFABP, 98 KR@IBRP, 40, 68, 117, 137 KR@IBRP-CFOPQ, 537 KR@IBRP-CFOPQ EVMLQEBPFP, 541 KRQOFBKQ, 604 KRQOFBKQP, 574, 588
*
/ ELOFWLK, 889, 897 L?IFD>QB >BOL?BP, 645, 669 L?IFD>QB >K>BOL?BP, 645, 669 L?PQOR@QFSB AFPB>PB, 1158 /?PQOR@QFSB AFPB>PBP, 1151 L@@FMFQ>I, 1011 L@@FMFQ>I IL?B, 1026 /@B>K RMTBIIFKD, 1290 L@B>K RMTBIIFKD, 1314 L@B>KF@ WLKB, 1303, 1314 /@Q>JBO ?LU, 419 L@Q>JBO ?LUBP, 406 L@QBQ ORIB, 45, 68 LALO>KQ, 1058 /ALO>KQP, 1040 /H>W>HF CO>DJBKQ, 392 /H>W>HF CO>DJBKQP, 384 LIC>@QFLK, 1040, 1058 LIC>@QLOV ?RI?, 1044, 1058 LIC>@QLOV BMFQEBIFRJ, 1040, 1058 LIC>@QLOV OB@BMQLO, 1040, 1058 LIFDLABKAOL@VQB, 1026 /IFDLABKAOL@VQBP, 995 LIFDLP>@@E>OFK, 877 /IFDLP>@@E>OFKP, 874 /JBD>, 85 LJBD> C>Q, 104 LJKFSLOB, 985 /JKFSLOBP, 961 LK@LDBKB, 295 LK@LDBKBP, 289 LK@LDBKF@ SFORP, 569 LK@LDBKF@ SFORPBP, 557 LK@LIVQF@ SFORP, 569 /K@LIVQF@ SFORPBP, 564 LKB-@EFIA MLIF@V, 1337, 1362 LLDBKBPFP, 1258, 1278 LMBK @FO@RI>QLOV PVPQBJ, 1164, 1184 LMBO>KQ @LKAFQFLKFKD, 1357, 1362 LMBO>QLO, 427, 442 LMBOLK, 443 LMBOLKP, 426
/MMLPFQFLK, 1113 LMMLPFQFLK, 1129 LMPLKFW>QFLK, 1219, 1242 LO?FQ>I, 68 LO?FQ>IP, 45 LOABO, 524, 541 LOD>K, 35 LOD>K LC #LOQF, 1047, 1058 LOD>K PVPQBJ, 27, 35 LOD>KBIIB, 35, 137 LOD>KBIIBP, 27, 115 LOD>KF@ @LJMLRKA, 884, 897 LOD>KF@ JLIB@RIB, 68 LOD>KF@ JLIB@RIBP, 59 LOD>KF@ PLFI, 897 LOD>KF@ PLFIP, 887 LOD>KFPJ, 35 /OD>KFPJP, 28 LOD>KLDBKBPFP, 730, 746, 1274, 1278 /OD>KP, 27 LOFDFK, 291, 295 /OKFQELOEVK@EFA>B, 823, 834 LP@RIRJ, 752, 793 LPJL@LKCLOJBO, 1208 LPJL@LKCLOJBOP, 1191 /PJLI>OFQV, 154 LPJLI>OFQV, 166 LPJLMEFIB, 604 LPJLOB@BMQLO, 1089 LPJLOB@BMQLOP, 1069 /PJLOBDRI>QFLK, 1190 LPJLOBDRI>QFLK, 1208 LPJLOBDRI>QLO, 1208 LPJLOBDRI>QLOV, 1191 /PJLPFP, 153 LPJLPFP, 166 LPJLQF@ ?>I>K@B, 1190, 1208 LPJLQF@ MOBPPROB, 1190, 1208 LPPBLRP QFPPRB, 1104, 1129 LPPF@IB, 1058 LPPF@IBP, 1045 /PPFCF@>QFLK, 1107 LPPFCF@>QFLK, 1129 /PQBF@EQEVBP, 805, 834 LPQBL?I>PQ, 1129 /PQBL?I>PQP, 1107 LPQBL@I>PQ, 1129 /PQBL@I>PQP, 1107 LPQBL@VQB, 1129 /PQBL@VQBP, 1107 LPQBLK, 955, 1129 LPQBLKP, 945 /PQBLKP, 1106 /PQBLMOLDBKFQLO @BIIP, 1107 LPQF>, 752, 1164
1472
).$%8
LPQFRJ, 793, 1184 LPQO>@LABOJ, 834 LPQO>@LABOJP, 802 LRQBO B>O, 1045, 1058 LS>I TFKALT, 1046, 1058 LS>OF>K @V@IB, 1261, 1278 LS>OV, 710, 724 LSFAR@Q, 1278 LSFAR@QP, 1257 LSFDBO, 793 LSFDBOP, 787 LSFM>OFQV, 1252, 1278 LSLSFM>OFQV, 1252, 1278 LSRI>QB @LKB, 724 LSRI>QB @LKBP, 705 LSRI>QFLK, 1262, 1278 LSRIB, 701, 724 LUFA>QFSB MELPMELOVI>QFLK, 199, 218 LUVDBK AFPPL@F>QFLK @ROSB, 1154, 1158 LUVDBK-@>OOVFKD @>M>@FQV, 1154, 1158 LUVQL@FK, 1072, 1089
+
00, 321, 348 M21, 288, 295 M53, 288, 295 0680, 234, 241 0700, 235, 241 0>@FKF>K @LOMRP@IB, 1058 0>@FKF>K @LOMRP@IBP, 1038 M>FOTFPB-BKA PBNRBK@FKD, 463 0>IBLDK>QE>B, 820, 834 0>IBLKQLILDV, 33 M>IBLKQLILDV, 35 M>IJ>QBIV @LJMLRKA IB>C, 855, 877 P(5, 825, 834 M>K@OB>P, 969, 985, 1083, 1089 M>KABJF@, 591, 604 M>MFII>, 1058 M>MFII>B, 1042 M>O>@BKQOF@, 365, 367 M>O>@OFKB PFDK>I, 268 M>O>@OFKB PFDK>IP, 247 M>O>CLIIF@RI>O @BII, 1089 M>O>CLIIF@RI>O @BIIP, 1082 M>O>MLAF>, 777 M>O>MLAFRJ, 793 M>O>PFQB, 1344, 1362 M>O>PFQF@ MI>KQ, 894, 897 0>O>PFQFPJ, 662 M>O>PFQFPJ, 669
M>O>PVJM>QEBQF@ KBOSLRP PVPQBJ, 1017, 1026 M>O>QEVOLFA DI>KA, 1089 M>O>QEVOLFA DI>KAP, 1082 M>O>QEVOLFA ELOJLKB (04(), 1075, 1089 0>O>WL>, 738, 746 M>OBK@EVJ> @BII, 877 0>OBK@EVJ> @BIIP, 843 M>OBKQ J>QBOF>I, 889, 897 0>OBKQ>I QVMBP, 355 M>OBKQ>I QVMBP, 367 M>OFBQ>I, 1011 M>OFBQ>I IL?B, 1026 0>OHFKPLKwP AFPB>PB, 1020, 1026 0>OQEBKLDBKBPFP, 1250 M>OQEBKLDBKBPFP, 1278 0>OQF>I MOBPPROB, 1142 M>OQF>I MOBPPROB, 1158 M>OQF@RI>QB J>QQBO, 1141, 1158 M>PPFSB FJJRKFQV, 1235, 1242 0>PPFSB QO>KPMLOQ, 149 M>PPFSB QO>KPMLOQ, 166 M>QBII>, 1102, 1129 M>QELDBK, 569, 1243 M>QELDBK->PPL@F>QBA JLIB@RI>O M>QQBOK (0!-0), 1242 M>QELDBK->PPL@F>QBA JLIB@RI>O M>QQBOKP (0!-0P), 1214 M>QELDBKP, 564, 1213 M>QQBOK OB@LDKFQFLK OB@BMQLO (022), 1243 M>QQBOK OB@LDKFQFLK OB@BMQLOP (022P), 1214 MB>Q JLPP, 694, 695 MB@QLO>I DFOAIB, 1100, 1129 MBAFDOBB >K>IVPFP, 329 MBAFM>IM, 793 MBAFM>IMP, 787 MBBO-OBSFBTBA J>KRP@OFMQ, 35 0BBO-OBSFBTBA J>KRP@OFMQP, 22 MBI>DF@ OB>IJ, 1301, 1314 MBIIF@IB, 638 MBIIF@IBP, 616 MBISF@ DFOAIB, 1101, 1129 MBKFP, 1255, 1278 0BMPFK, 967 MBMPFK, 985 MBMPFKLDBK, 967, 985 MBMQFAB ?LKA, 91, 104 MBMQFAB ELOJLKB, 1089 MBMQFAB ELOJLKBP, 1065 MBMQFALDIV@>K, 585, 604
MBMQFAVI QO>KPCBO>PB, 415, 419 0BO@BMQFLK, 1033 MBO@BMQFLK, 1058 MBOCLOFK, 1219, 1243 MBOF>KQE, 710, 724, 902, 929 MBOF@>OAFRJ, 1175, 1184 MBOF@>OM, 921, 929 MBOF@BKQOF@, 365, 367 MBOF@V@IB, 852, 877 MBOFABOJ, 848, 877 MBOFLAF@ Q>?IB, 44, 68 MBOFMEBO>I MOLQBFK, 166 0BOFMEBO>I MOLQBFKP, 146 MBOFMEBO>I OBPFPQ>K@B, 1182, 1184 MBOFOBK>I C>Q @>MPRIB, 1193, 1208 MBOFPQ>IPFP, 966, 985 MBOFPQLJB, 686, 695 MBOFQR?RI>O @>MFII>OV KBQTLOH, 1195, 1208 MBOJ>COLPQ, 1300, 1314 MBOJ>KBKQ QFPPRB, 841, 877 MBOJFPPFSB, 553, 569 MBOLUFPLJB, 137 0BOLUFPLJBP, 120 MBQ>I, 724 0BQ>IP, 710 MBQFLIB, 854, 877 0BQOLJVWLKQFA>B, 803, 834 M( M>MBO, 68 M( P@>IB, 57, 68 ME>DB QEBO>MV, 564, 569 ME>DLIVPLPLJB, 616, 638 ME>I>KDB, 1129 ME>I>KDBP, 1101 0E>OJ>@LDBKLJF@P, 466 ME>OJ>@LDBKLJF@P, 471 ME>OVKDB>I PIFQ, 834 0E>OVKDB>I PIFQP, 799 ME>OVKU, 1139, 1158 MEBKLQVMB, 326, 348 MEBOLJLKB, 1042, 1058 0EILBJ, 687 MEILBJ, 695 MELPME>Q>PB, 268 MELPME>Q>PBP, 262 MELPMEL>KEVAOFAB ?LKA, 190 MELPMEL>KEVAOFAB ?LKAP, 181 MELPMELAFBPQBO, 99, 104 MELPMELAFBPQBO>PB, 262, 268, 1068 MELPMELAFBPQBO>PB (0$%), 1089 MELPMELIFMFA, 104 0ELPMELIFMFAP, 86 0ELPMELOVI>QFLK, 198
).$%8
1473
MELPMELOVI>QFLK, 218 MELQF@ WLKB, 1301, 1314 MELQL>@Q, 234, 241 MELQL>RQLQOLME, 241 MELQL>RQLQOLMEP, 224 0ELQLJLOMELDBKBPFP, 868 MELQLJLOMELDBKBPFP, 877 MELQLK, 233, 241 0ELQLMBOFLAFPJ, 868 MELQLMBOFLAFPJ, 877 MELQLPVPQBJ, 233, 242 MELQLPVPQBJ ), 233, 242 MELQLPVPQBJ )), 233, 241 MELQLQOLME, 604 MELQLQOLMEP, 575 0ELQLQOLMEP, 589 MELQLQOLMFK, 877 MELQLQOLMFKP, 870 0ELQLQOLMFPJ, 868 MELQLQOLMFPJ, 877 MEVIILQ>UV, 855, 877 MEVILDBKBQF@ QOBB, 30, 35, 522, 541 0EVILDBKV, 522 MEVILDBKV, 541 MEVIRJ, 524, 541 MEVPF@>I J>M, 458, 471 MEVPF@>I P@FBK@B, 35 MEVPF@>I P@FBK@BP, 15 MEVPFLILDF@>I AB>A PM>@B, 1152, 1158 MEVQL@EOLJB, 877 MEVQL@EOLJBP, 869 MF> J>QBO, 1008, 1026 MFDJBKQ, 226, 242 MFIF, 581 MFIFAFRJ, 771, 793 MFIRP, 604 MFK>@L@VQB, 793 0FK>@L@VQBP, 752 MFKK>, 1045, 1058 MFKK>QBIV @LJMLRKA IB>C, 877 0FKK>QBIV @LJMLRKA IB>SBP, 855 MFKL@VQLPFP, 162, 166 MFLKBBO PMB@FBP, 1349, 1362 MFPQFI, 710, 724 MFQE, 847, 877 MFQRFQ>OV AT>OCFPJ, 1077, 1089 MFQRFQ>OV DI>KA, 1080, 1089 MFQRFQ>OV PQ>IH, 1080, 1089 MFSLQ GLFKQ, 1129 0FSLQ GLFKQP, 1115 MI>@BKQ>, 1266, 1278 MI>DF>OFPJ, 22, 35 MI>K>O GLFKQ, 1129 0I>K>O GLFKQP, 1114 MI>KHQFSLOB, 1314
MI>KHQFSLOBP, 1304 MI>KHQLK, 621, 638 MI>KLPMFO>I, 773, 793 0I>KQ>O CIBUFLK, 1113 MI>KQ>O CIBUFLK, 1129 MI>KRIFCLOJ, 771, 793 MI>PJ>, 1167, 1184 MI>PJ> @BII, 1225, 1243 MI>PJ> JBJ?O>KB, 116, 137 MI>PJ> JBJ?O>KB ELOJLKB OB@BMQLO, 1089 MI>PJ> JBJ?O>KB ELOJLKB OB@BMQLOP, 1067 MI>PJFA, 419 MI>PJFAP, 402 MI>PJLABPJ>, 137 MI>PJLABPJ>Q>, 133 MI>PJLD>JV, 647, 669 MI>PJLIVPFP, 156, 166 MI>PQFA, 612, 638 MI>QBIBQ, 1184 MI>QBIBQP, 1167 0I>QVOOEFKF, 825, 834 P3,:0(+(70:, 825, 834 MIBRO>, 1150, 1158 0IBROFPV, 1150 MIBROFPV, 1158 MIRJRIB, 919, 929 MKBRJ>QF@ ?LKB, 834 0KBRJ>QF@ ?LKBP, 818 MLFKQ JRQ>QFLK, 392 0LFKQ JRQ>QFLKP, 390 MLI>O @LS>IBKQ ?LKA, 50, 68 MLI>O KR@IBF, 907, 929 MLIIBK DO>FK, 724 MLIIBK DO>FKP, 700 MLIIBK QR?B, 702, 724 MLIIFK>QFLK, 717, 724, 911, 929 MLIV-! Q>FI, 410, 419, 437, 443 MLIV>KAOLRP, 1355 MLIV>KAOV, 1362 0LIV@>OMF@, 927 MLIV@>OMF@, 929 MLIVDBKF@, 465, 471 0LIVDVKLRP, 1355 MLIVDVKV, 1362 MLIVJBO, 104 MLIVJBO>PB @E>FK OB>@QFLK (0#2), 471 MLIVJBOP, 72 MLIVJLOMEF@, 760, 793 MLIVKR@IBLQFAB, 98, 104 MLIVM, 756, 793 MLIVMBMQFAB, 104 MLIVMILFA, 362, 367 MLIVP>@@E>OFAB, 78, 104 MLIVPLJB, 413, 419
MLIVPMBOJV, 1271, 1278 MLIVQLJV, 522, 541 P65.6, 825, 834 MLMRI>QFLK, 28, 35 MLMRI>QFLK ABKPFQV, 1318, 1362 MLMRI>QFLK DBKBQF@P, 503, 517 MLMRI>QFLK DOLTQE O>QB, 1326, 1362 MLMRI>QFLK PFWB (N), 1318, 1362 MLMRI>QFLK S>OF>QFLK, 505, 518 0LOFCBO>, 752, 793 MLPFQFSB CBBA?>@H ILLM, 951, 955 MLPFQFSB DO>SFQOLMFPJ, 871, 877 0LPFQFSB MLI>OFQV, 551 MLPFQFSB MLI>OFQV, 569 MLPFQFSB OBDRI>QLO, 427, 443 MLPQ->K>I Q>FI, 799, 834 MLPQ-QO>KP@OFMQFLK>I, 424, 443 MLPQ-QO>KPI>QFLK>I, 424, 443 MLPQBOFLO MFQRFQ>OV, 1081, 1089 MLPQWVDLQF@ ?>OOFBO, 492, 497 MLQBKQF>I BKBODV, 173, 190 MLQL@VQLPFP, 162, 166 MOB@>MFII>OV PMEFK@QBO, 1184 MOBA>QLO, 1314 MOBA>QLOP, 1304 MOBFKFQF>QFLK @LJMIBU, 407, 419 MOBP?VLMF>, 1051, 1058 MOBWVDLQF@ ?>OOFBO, 492, 497 MOFJ>OV (J>FK) ?OLK@EF, 1140 0OFJ>OV >@QFSB QO>KPMLOQ, 158 MOFJ>OV >@QFSB QO>KPMLOQ, 166 MOFJ>OV ?OLK@ERP, 1158 MOFJ>OV @LKPRJBO, 1393 MOFJ>OV @LKPRJBOP, 1369 MOFJ>OV BIB@QOLK >@@BMQLO, 242 MOFJ>OV BIB@QOLK >@@BMQLOP, 234 MOFJ>OV CB>QEBO, 834 0OFJ>OV CB>QEBOP, 818 MOFJ>OV DOLTQE, 847, 877 MOFJ>OV MOLAR@BO, 1393 MOFJ>OV MOLAR@BOP, 1369 MOFJ>OV PQOR@QROB, 93, 104 MOFJ>OV PR@@BPPFLK, 1349, 1362 MOFJ>PB, 384, 392
1474
).$%8
0OFJ>QBP, 824, 834 MOFJBO, 384, 392 MOFLK, 569 0OFLKP, 564 MOL?B, 471 MOL?BP, 451 MOLAR@Q, 68 MOLAR@Q ORIB, 324, 348 MOLAR@QFSB, 555, 569 MOLAR@QP, 48 0OLDBPQBOLKB, 1260 MOLDBPQBOLKB, 1278 MOLDK>QEF@ G>T, 834 MOLDK>QEF@ G>TP, 828 MOLDVJKLPMBOJ, 724 MOLDVJKLPMBOJP, 700 MOLH>OVLQB, 35, 113, 137 0OLH>OVLQBP, 27 MOLI>@QFK (02,), 1072, 1089 MOLI>@QFK-FKEF?FQFKD ELOJLKB, 1089 MOLI>@QFK-FKEF?FQFKD ELOJLKB (0)(), 1072 MOLI>@QFK-OBIB>PFKD ELOJLKB, 1089 MOLI>@QFK-OBIB>PFKD ELOJLKB (02(), 1072 MOLJBQ>ME>PB, 280, 295 MOLJLQBO, 402, 419 0OLK>QFLK, 1113 MOLK>QFLK, 1129 MOLLCOB>AFKD, 388, 392 MOLME>DB, 555, 569 MOLME>PB, 279, 295 MOLMOFL@BMQFLK, 1011, 1027, 1032, 1058 MOLPFJF>K, 834 0OLPFJF>KP, 824 MOLPQ>QB DI>KA, 1255, 1278 MOLPQEBQF@ DOLRM, 206, 218 MOLQB>PB, 471 MOLQB>PBP, 448 MOLQB>PLJB, 439, 443 MOLQBFK, 104 MOLQBFK PFDK>QROB, 469, 471 0OLQBFKP, 89 MOLQBLJB, 468, 471 MOLQBLJF@P, 468, 471 MOLQL-LK@LDBKB, 295 MOLQL-LK@LDBKBP, 289 MOLQLK, 41, 68 MOLQLKBJ>, 685, 696 MOLQLKBMEOFAF>, 1200 MOLQLPQLJB, 747 0OLQLPQLJBP, 736 0OLQO>@QFLK, 1113 MOLQO>@QFLK, 1129 MOLSBKQOF@RIRP, 963, 985 MOLUFJ>I @LKSLIRQBA QR?RIB (0#4), 1195, 1208
0O0@, 565, 569 0O0P@, 565, 569 MPBRAL@LBILJ>QB, 747 MPBRAL@LBILJ>QBP, 735 MPBRALMBMQFALDIV@>K, 586, 604 MPBRALPQO>QFCFBA, 941, 955 MPV@EOLMEFIB, 604 MRIJL@RQ>KBLRP @FO@RI>QFLK, 1167, 1184 MRIJLK>OV @FO@RI>QFLK, 1166, 1184 MRJM, 166 MRJMP, 158 MRK@QR>QBA BNRFIF?OFRJ, 495, 497 0RKKBQQ PNR>OB, 327, 348 MRMFI, 1058 MROB @RIQROB, 466, 471 MROFKB, 104 MROFKBP, 99 MVOFJFAFKB, 104 MVOFJFAFKBP, 99 MVORS>QB, 200, 218
,
NR>AO>Q, 1319, 1362 NR>QBOK>OV PQOR@QROB, 96, 104 NRFBP@BKQ, 281, 295 NRLORJ PBKPFKD, 263, 268
9-PBIB@QBA PMB@FBP, 1333, 1360 O>AF>I, 736 O>AF>I @IB>S>DB, 747 2>AF>I DIF>, 995 O>AF>I DIF>, 1027 2>AF>I PVJJBQOV, 733 O>AF>I PVJJBQOV, 747 O>AF>QB >OQBOFBP, 1194 2>AF>QFLK EV?OFA J>MMFKD, 460 O>AF>QFLK EV?OFA J>MMFKD, 471 O>AF@IB, 919, 929 O>AFLFPLQLMB, 68 O>AFLFPLQLMBP, 42 O>AFLOBPFPQ>KQ, 576, 604 O>AFRP, 1101, 1129 O>ARI>, 772, 793 O>MEB, 625, 638 OB>@Q>KQ, 68 OB>@Q>KQP, 48 OB>@QFLK @BKQBO, 233, 242 OB>AFKD CO>JB, 399, 419
OB@BMQFLK, 1032, 1058 OB@BMQFSB, 1032 OB@BMQFSB CFBIA, 1058 OB@BMQLO, 268 OB@BMQLO MLQBKQF>I, 1032, 1058 OB@BMQLO-JBAF>QBA BKAL@VQLPFP, 163, 166 OB@BMQLOP, 246, 1066 OB@BPPFSB, 348 OB@BPPFSB IBQE>I, 336, 348 2B@BPPFSB QO>FQP, 323 OB@FMOL@>I @OLPP, 323, 348 OB@LJ?FK>KQ $.!, 453, 471 OB@LJ?FK>KQ MOLQBFK, 471 OB@LJ?FK>KQ MOLQBFKP, 453 OB@LJ?FK>QFLK COBNRBK@V, 357, 367 OB@LJ?FK>QFLK KLARIBP, 303, 315 OB@ORFQJBKQ, 1152, 1158 OB@QRJ, 969, 985 OBA ?ILLA @BII, 1184 2BA ?ILLA @BIIP, 1168 OBALU OB>@QFLK, 218 OBALU OB>@QFLKP, 196 OBAR@QFLK, 237, 242 OBAR@QFLK AFSFPFLK, 309, 315 OBCIBU >@QFLK, 1350, 1362 OBCO>@QLOV MBOFLA, 1000, 1027 OBDRI>QLOV 4 (4OBD) @BII, 1243 OBDRI>QLOV 4 (4OBD) @BIIP, 1228 OBFKCLO@BJBKQ, 495, 497 OBI>QFSB CFQKBPP, 511, 518 2BI>QFSB PMB@FBP >?RKA>K@B, 1346 OBI>QFSB PMB@FBP >?RKA>K@B, 1362 OBK>I >OQBOFBP, 1194 OBK>I >OQBOV, 1208 OBK>I @>MPRIB, 1193, 1208 OBK>I @LIRJK, 1208 OBK>I @LIRJKP, 1194 OBK>I @LOMRP@IB, 1194, 1208 OBK>I C>P@F>, 1193, 1208 OBK>I MBISFP, 1193, 1208 OBK>I MVO>JFA, 1208 OBK>I MVO>JFAP, 1194 OBK>I QR?RIB, 1194, 1209 OBK>I SBFK, 1209 OBK>I SBFKP, 1194 OBKFK, 1069, 1089 OBKFK->KDFLQBKPFK>IALPQBOLKB, 1205, 1209 OBMIF@>QFLK CLOH, 392 OBMIF@>QFLK CLOHP, 383
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1475
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ISBN-10 ISBN-13 Revision

1938168097 978-1-938168-09-3 BM-1-001-DW

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