nCC1. 1he braln ls Lhe body's mosL complex organ. /("*'-'0 &12(,%-3(+ AfLer sLudy of Lhe asslgned learnlng maLerlals, Lhe sLudenL wlll: 1. uemonsLraLe Lhe four palred lobes of Lhe cerebral corLex and descrlbe Lhe boundarles of each. 2. SkeLch Lhe ma[or feaLures of each cerebral lobe, as seen from Lhe laLeral vlew, ldenLlfylng ma[or gyrl and sulcl LhaL characLerlze each lobe. '"**"%-3( by Leonard L. WPl1L and nell 8. CAn1 uuke lnsLlLuLe for 8raln Sclences ueparLmenL of neuroblology uuke unlverslLy School of Medlclne !"#$"%#& When you vlew Lhe laLeral aspecL of a human braln speclmen (see '%()$#* ,-, and ,./ 0 ), Lhree sLrucLures are usually vlslble: Lhe !"#"$#%& (")*+,("#"+, Lhe !"#"$"&&-), and parL of Lhe $#%*.+/") (alLhough Lhe bralnsLem ls noL vlslble ln Lhe speclmen phoLographed ln laLeral vlew for '%(1 . below). 1he splnal cord has usually been severed (buL we'll conslder Lhe splnal cord laLer), and Lhe resL of Lhe subdlvlslons are hldden from laLeral vlew by Lhe hemlspheres. 1he dlencephalon and Lhe resL of Lhe bralnsLem are vlslble on Lhe medlal surface of a braln LhaL has been cuL ln Lhe mldsaglLLal plane. arLs of all of Lhe subdlvlslons are also vlslble from Lhe venLral surface of Lhe whole braln. Cver Lhe nexL several LuLorlals, you wlll flnd vldeo demonsLraLlons (from Lhe braln anaLomy lab) and phoLographs (ln Lhe LuLorlal noLes) of Lhese braln surfaces, and sufflclenL deLall ln Lhe narraLlve Lo appreclaLe Lhe overall organlzaLlon of Lhe parLs of Lhe braln LhaL are vlslble from each perspecLlve. As you work Lhrough Lhls LexL and lf you have access Lo an lnLeracLlve dlglLal aLlas of Lhe human braln, such as !"#$%&'(, flnd Lhe sLrucLures and reglons LhaL are descrlbed here 3 . 1he !"#"$#%& (")*+,("#"+ are especlally large ln humans. 1hey are enLlrely covered by a 2-3-mm Lhlck layer of cells and cellular processes called Lhe !"#"$#%& !0#/"1. 1he surface of each hemlsphere ls hlghly
1 vlslL "#$%&'$()*+,#- for '456789:4;94 ,764 ,7;94<=8 (2012 SocleLy for neurosclence ) LhaL offer fundamenLal prlnclples abouL Lhe braln and nervous sysLem, Lhe mosL complex llvlng sLrucLure known ln Lhe unlverse. 2 llgure references Lo urves eL al., 1$)%,2+/$0+$3 4 #- 567, Slnauer Assoc., lnc., 2012. [cllck here] 3 1o do so ln !"#$%&'(, launch !"#$%&'( and go Lo 2(0/03#%,(*! 4/&%+, Lhen selecL one of Lhe aLlas fllLers, such as 56#*, 70$"+, or 8-&!* %.9 :*++-#"+. !"#$%"& ()%*"+$ ,* #-$ .%"/0 - Sprlng 2013
2 lnfolded, Lhe rldges Lhus formed are known as 36#* (slngular: gyrus) and Lhe valleys are called +-&!* (slngular: sulcus) or ;*++-#"+ (lf Lhey are especlally deep). 1he appearance of Lhe sulcl and gyrl varles somewhaL from braln Lo braln. (As you mlghL guess, each one has lLs own name, buL lL ls necessary Lo become famlllar wlLh only a few of Lhem.) 1he hemlspheres are convenLlonally dlvlded lnLo lobes named for Lhe bones of Lhe skull LhaL overlle Lhem, namely Lhe ;#0./%&, ,%#*"/%&, 0!!*,*/%& and /"),0#%& &0$"+ (see '%()$# ,-2.
345#$46 4*7#85 9: 5;# <$4%= 1he fronLal lobe ls Lhe mosL anLerlor of Lhe four lobes and ls separaLed from Lhe parleLal lobe by Lhe !"./#%& +-&!-+, whlch ls one of Lhe mosL lmporLanL landmarks ln Lhe cerebral corLex (ln '%()$# ,-, boundary beLween blue and red colored reglons, see LuLorlal on )%*+%*, ./0 10*.23# !&'). An lmporLanL gyrus ln Lhe fronLal lobe ls Lhe ,#"!"./#%& 36#-+. (1he preflx 'pre,' when used Lo refer Lo anaLomlcal poslLlon, refers Lo someLhlng LhaL ls ln fronL of someLhlng else or LhaL ls anLerlor.) 1he corLex of Lhe precenLral gyrus ls Lhe somaLlc 'moLor corLex,' whlch conLalns neurons whose axons pro[ecL Lo Lhe moLor nuclel ln Lhe bralnsLem and splnal cord LhaL lnnervaLe Lhe sLrlaLed muscles of Lhe body. llne, skllled movemenLs are dependenL on Lhe lnLegrlLy of Lhe moLor corLex (and, of course, Lhe axons exLendlng from lL). 1he axons LhaL arlse from neurons ln Lhe moLor corLex and exLend Lo Lhe splnal cord are known as Lhe !0#/*!0+,*.%& /#%!/. 1hose axons LhaL exLend from Lhe moLor corLex Lo nuclel ln Lhe bralnsLem are known as Lhe !0#/*!0$-&$%# /#%!/. (1he bralnsLem ls someLlmes referred Lo as 'bulbar' because lL has a shape resembllng a bulb.) Whlle you can'L see Lhe lndlvldual flbers LhaL make up Lhe LracLs, you can see Lhe sLrucLures Lhrough whlch Lhey pass as Lhey course beLween Lhe corLex and Lhelr LargeLs ln Lhe bralnsLem and splnal cord. Cn Lhe lnferlor-laLeral aspecL of Lhe hemlsphere, you should readlly appreclaLe a deep, falrly sLralghL flssure LhaL separaLes Lhe fronLal and parleLal lobes from Lhe Lemporal lobe, Lhls space ls called Lhe &%/"#%& ;*++-#" or Sylvlan flssure, named afLer Lhe lmporLanL 8enalssance neuroanaLomlsL, 8%"0+/2+)2 (9&:/)2 ; . lf vlewed from above, you would see LhaL Lhe hemlspheres are separaLed by an even deeper flssure called Lhe &0.3*/-9*.%& ;*++-#" (or superlor saglLLal flssure) (see '%()$# ,..2. 1he gyral formaLlons anLerlor Lo Lhe precenLral gyrus on Lhe dorsal-laLeral aspecL of Lhe fronLal lobe beLween Lhe laLeral flssure and Lhe longlLudlnal flssure can be recognlzed as Lhree parallel, longlLudlnal gyrl. Ad[acenL Lo Lhe longlLudlnal flssure ls Lhe +-,"#*0# ;#0./%& 36#-+, whlch ls Lyplcally Lhe mosL obvlous of Lhe Lhree. 1hls gyrus conLlnues on Lhe medlal surface of Lhe hemlsphere ln Lhe depLhs of Lhe longlLudlnal flssure, we wlll see lL agaln when we examlne Lhe mldsaglLLal secLlon Lhrough Lhe braln. !usL lnferlor Lo Lhe superlor fronLal gyrus ls Lhe )*99&" ;#0./%& 36#-+, wlLh Lhe +-,"#*0# ;#0./%& +-&!-+ separaLlng Lhe Lwo. noLe LhaL Lhese Lwo gyral formaLlons are falrly sLralghL and parallel Lo Lhe longlLudlnal flssure ln Lhe parasaglLLal plane. AL Lhelr posLerlor end, Lhey someLlmes merge wlLh Lhe precenLral gyrus, whlch forms, of course, Lhe anLerlor bank of Lhe cenLral sulcus. Powever, Lhere ls ofLen an lnLerrupLed (l.e., a superflclally dlsconLlnuous) sulcus LhaL separaLes Lhem from Lhe precenLral gyrus, named Lhe ,#"!"./#%& +-&!-+.
4 8%"0+/2+)2 (9&:/)2 was Lhe lnsplraLlon for Lhe LlLle of our dlglLal braln aLlas, !"#$%&'(, cllck here for more on Lhls lmporLanL flgure ln Lhe hlsLory of human neuroanaLomy. lf lt were osslble to unfold the cerebrul cortex from one hemlshere (whlch cun be done ln dlgltul reresentutlons of the cerebrul hemlshere), the surfuce ureu of the resultlng, fluttened cerebrul cortex would be roughly uroxlmuted by the crust of u lnch lzzu (thln crust, '4> ?76@ style, of course, glven the thlnness of the cortex). !"#$%"& ()%*"+$ ,* #-$ .%"/0 - Sprlng 2013
3
'%(1 .1 Luterul surfuce of the humun bruln. 1hls flgure ls not lubeled so thut you muy refer to lt for revlew, see '%()$#* ,- > ,./ for un lllustruted und lubeled vlew of the sume hemlshere. (lmuge from !"#$%&'()
!usL lnferlor Lo Lhe mlddle fronLal gyrus, across Lhe *.;"#*0# ;#0./%& +-&!-+, ls a much more complex gyral formaLlon. lor our purposes, we won'L be concerned wlLh Lhe names of Lhese subcomponenLs or Lhelr dlfferenLlal funcLlonal conLrlbuLlons Lo cognlLlon and behavlor. Sufflce lL Lo say LhaL Lhe *.;"#*0# ;#0./%& 36#-+ conLalns a crlLlcal funcLlonal dlvlslon of Lhe moLor corLex LhaL parLlclpaLes ln Lhe producLlon of speech. 1hls dlvlslon has a speclal name, <#0!%=+ 4#"%, glven ln honor of Lhe famous lrench neurologlsL, lerre aul 8roca 3 , who flrsL recognlzed Lhe slgnlflcance of Lhls gyral formaLlon for human speech ln Lhe mld-19Lh cenLury. lnLeresLlngly, Lhe lefL hemlsphere ls domlnanL ln mosL lndlvlduals (especlally males and rlghL-handers) for Lhls funcLlon, such LhaL damage Lo Lhe lefL lnferlor fronLal gyrus ls more llkely Lo produce an lmpalrmenL of language expresslon, called .%,+"<2 "=-"2/" (aphasla means wlLhouL speech"), Lhan a comparable leslon lnvolvlng Lhe rlghL lnferlor fronLal gyrus. lnLeresLlngly, Lhls ls also Lhe dlvlslon of Lhe premoLor corLex where ln non-human prlmaLes neurons wlLh mlrror" properLles have been characLerlzed. 1haL ls, neurons ln Lhe posLerlor parL of Lhe lnferlor fronLal gyrus flre when cerLaln
3 cllck here for more on Lhls lmporLanL flgure ln Lhe hlsLory of human neuroanaLomy. !"#$%"& ()%*"+$ ,* #-$ .%"/0 - Sprlng 2013
4 acLlons are execuLed "06 when Lhose same acLlons are observed (mlrrored) ln Lhe behavlor of anoLher lndlvldual. WheLher Lhese neurons parLlclpaLe ln lmlLaLlon learnlng-or perhaps even decodlng Lhe lnLenLlons of oLhers-remalns an area of lnLense lnvesLlgaLlon. nexL, leL's cross Lhe laLeral flssure. 1he gyral sLrucLure LhaL forms Lhe lnferlor bank of Lhe laLeral flssure ls Lhe +-,"#*0# /"),0#%& 36#-+. 1hls gyrus ls arranged parallel Lo Lhe laLeral flssure and exLends from Lhe anLerlor pole of Lhe Lemporal lobe Lo Lhe parleLal lobe (speclflcally, Lhe angular gyrus of Lhe lnferlor parleLal lobule). 1he elongaLlon of Lhls sLrucLure ln nearly Lhe horlzonLal plane esLabllshes a framework for conslderlng Lhe remalnlng Lemporal gyral formaLlons of Lhe laLeral surface. lncludlng Lhe superlor Lemporal gyrus, lL should be posslble Lo recognlze Lhree elongaLed gyral sLrucLures LhaL lle parallel Lo Lhe laLeral flssure: Lhe +-,"#*0#, )*99&" and *.;"#*0# /"),0#%& 36#*. 1he superlor and mlddle Lemporal gyrl are separaLed by Lhe +-,"#*0# /"),0#%& +-&!-+ and Lhe mlddle and lnferlor gyrl are separaLed by Lhe *.;"#*0# /"),0#%& +-&!-+, alLhough Lhe laLLer sulcus ls ofLen dlsconLlnuous and someLlmes dlfflculL Lo recognlze. noLlce how Lhe laLeral aspecLs of Lhe fronLal and Lemporal lobes bear some resemblance (aL leasL concepLually): Lhey boLh comprlse Lhree, parallel longlLudlnal gyrl LhaL exLend from Lhe anLerlor pole of each lobe back Loward Lhe parleLal lobe. Cnce you are famlllar wlLh Lhese gyral feaLures ln one speclmen, spend some Llme examlnlng as many speclmens as are avallable ln dlglLal formaL (a number of free dlglLal resources are avallable LhaL show laLeral vlews of Lhe human braln). As you do so, look for dlfferences and common sLrucLural Lhemes among bralns and beLween hemlspheres. lf you can, carefully examlne Lhe paLLern and lengLh of Lhe laLeral flssure and conslder Lhe followlng quesLlons. Are Lhe Lwo laLeral flssures of Lhe same braln roughly Lhe same? lf noL, how are Lhey dlfferenL? uoes Lhe lefL laLeral flssure Lend Lo run malnly ln Lhe horlzonLal plane and exLend furLher posLerlorly, compared Lo Lhe rlghL laLeral flssure? ls Lhere an ascendlng branch of Lhe posLerlor laLeral flssure ln Lhe rlghL hemlsphere, buL noL Lhe lefL? 1hese quesLlons have been asked and addressed ln sLudles of hemlspherlc asymmeLrles and gender dlfferences relaLed Lo language laLerallzaLlon ln Lhe human braln. 1he superlor aspecL of Lhe Lemporal lobe conLalns Lhe corLlcal dlvlslons whose funcLlons perLaln Lo audlLlon and Lhe recepLlon of language. 1he posLerlor aspecL of Lhe superlor Lemporal gyrus has a speclal funcLlonal name, >"#.*!?"=+ 4#"%, named ln recognlLlon of Lhe semlnal conLrlbuLlons of Lhe 19Lh cenLury Cerman physlclan, Carl Wernlcke 6 , who descrlbed a dlsLurbance of undersLandlng speech, now called >$%0/+?$<2 "=-"2/". ?ou may flnd dlfferences ln Lhe sLrucLure of Lhe superlor Lemporal gyrus ln Lhe Lwo hemlspheres reflecLed ln Lhe paLLern and lengLh of Lhe laLeral flssure, such dlfferences may, ln Lurn, be relaLed Lo Lhe language domlnance of Lhe lefL hemlsphere. Pemlspherlc dlfferences ln Lhls corLlcal reglon may even relaLe Lo muslcal ablllLles and oLher speclal LalenLs LhaL perLaln Lo audlLlon and semanLlc encodlng. CLher reglons of Lhe Lemporal lobe serve vlsual processlng, emoLlonal processlng, memory, and Lhe lnLegraLlon of sensory experlence, we wlll conslder Lhese funcLlons laLer ln Lhe course. Slnce you already worked Lhrough recognlLlon of Lhe cenLral sulcus, you should now be able Lo vlew Lhe laLeral surface of any hemlsphere and know Lo look for Lhe cenLral sulcus lazlly courslng from Lhe longlLudlnal flssure over Lhe dorsal-laLeral surface of Lhe cerebrum ln a laLeral-venLral and sllghLly anLerlor dlrecLlon (hopefully, Lhe dlrecLlonal Lerms applled Lo Lhe forebraln are also becomlng second- naLure Lo you). Much of Lhe corLex LhaL you observe posLerlor Lo Lhe cenLral sulcus ls parL of Lhe parleLal lobe. 1he promlnenL gyral sLrucLure LhaL forms Lhe posLerlor bank of Lhe cenLral sulcus ls Lhe ,0+/!"./#%&
6 cllck here for more on Lhls lmporLanL flgure ln Lhe hlsLory of human neuroanaLomy. !"#$%"& ()%*"+$ ,* #-$ .%"/0 - Sprlng 2013
5 36#-+, whlch ls parallel Lo Lhe precenLral gyrus of course. 1hls gyrus ls concerned wlLh somaLlc sensaLlon and wlll be a ma[or focus for cllnlclans concerned wlLh Lhe cerebral locallzaLlon of Louch and body poslLlon sensaLlons. lmmedlaLely posLerlor Lo Lhe posLcenLral gyrus ls Lhe ,0+/!"./#%& +-&!-+, whlch separaLes Lhe posLcenLral gyrus from Lwo ma[or gyral formaLlons of Lhe parleLal lobe: Lhe +-,"#*0# and *.;"#*0# ,%#*"/%& &0$-&"+. 1he boundary beLween Lhese Lwo lobules ls ofLen dlfflculL Lo appreclaLe, lL ls recognlzed as a meanderlng and ofLen dlsconLlnuous sulcus called Lhe *./#%,%#*"/%& +-&!-+ LhaL Lends Lo run ln a parasaglLLal plane. We wlll reLurn Lo Lhe superlor parleLal lobule when we explore Lhe medlal surface of Lhe hemlsphere. 1he gyral formaLlons [usL lnferlor Lo Lhe lnLraparleLal sulcus lnclude Lhe +-,#%)%#3*.%& 36#-+ and Lhe %.3-&%# 36#-+, Lwo prlnclpal componenLs of Lhe lnferlor parleLal lobule. 1he lnferlor margln of Lhe parleLal lobe ls formed mosLly by Lhe promlnenL laLeral flssure. Powever, Lhe posLerlor llmlL of Lhe laLeral flssure does noL deflne Lhe enLlre lnferlor boundary of Lhe parleLal lobe. 1he supramarglnal gyrus usually forms a horseshoe" shape around Lhe posLerlor llmlL of Lhe laLeral flssure, wlLh Lhe angular gyrus [usL posLerlor Lo Lhe supramarglnal gyrus. Llkewlse, lL ls ofLen dlfflculL Lo dlscern Lhe boundarles where Lhe posLerlor parleLal and Lemporal lobes meeL Lhe anLerlor occlplLal lobe on Lhe laLeral surface of Lhe cerebrum, and lL may make no funcLlonal sense Lo aLLempL Lo do so. neverLheless, we can deflne an lmaglnary laLeral boundary beLween Lhe parleLal and occlplLal lobes ln Lhls complex and varlable reglon. Lmerglng from Lhe depLhs of Lhe longlLudlnal flssure along Lhe medlal bank of Lhe cerebral hemlsphere (abouL one-fourLh of Lhe lengLh of Lhe flssure from lLs posLerlor llmlL) ls Lhe ,%#*"/0@0!!*,*/%& +-&!-+ (see '%()$# ,./). We wlll see Lhls sulcus more plalnly when we explore Lhe medlal face of Lhe hemlsphere ln anoLher LuLorlal. lor now, appreclaLe LhaL Lhe parleLo-occlplLal sulcus ls Lhe medlal boundary beLween Lhe parleLal and occlplLal lobes. vlew Lhe laLeral surface of Lhe hemlsphere agaln and carefully lnspecL lLs lnferlor margln. !usL above Lhe cerebellum, Lhere ls ofLen a small groove or noLch ln Lhe gyral sLrucLure LhaL can be appreclaLed 3-4 cm anLerlor from Lhe caudal pole of Lhe hemlsphere. 1hls groove ls called Lhe pre- occlplLal noLch" (see '%()$# ,./). now, draw an lmaglnary llne beLween Lhe parleLo-occlplLal sulcus dorsally and Lhe pre-occlplLal noLch venLrally, Lhls llne wlll serve as Lhe laLeral boundary beLween Lhe parleLal and occlplLal lobes. Slmply puL, all gyral sLrucLures posLerlor Lo Lhls llne are occlplLal and have some role Lo play ln elaboraLlng vlsual percepLlon. lorLunaLely for you, Lhe complexlLy and lnLer- lndlvldual varlaLlon of Lhese gyrl defles easy appllcaLlon of sLandard nomenclaLure, for Lhls reason, we wlll refer Lo Lhese gyral sLrucLures as &%/"#%& 0!!*,*/%& 36#* and leave Lhe buslness of namlng Lhese gyrl for Lhe corLlcal carLographers. LasLly, Lhere ls a reglon of corLex called Lhe *.+-&%, whlch ls noL vlslble from Lhe laLeral surface of Lhe hemlsphere because lL ls hldden beneaLh Lhe fronLal, Lemporal, and parleLal lobes. 1he componenLs of Lhese lobes LhaL cover Lhe lnsular corLex are ofLen called 'opercular' componenLs ('opercular' means a lld or cover). lL can be seen lf porLlons of Lhese Lwo lobes are reLracLed (as ls lllusLraLed ln '%()$# ,./). ln splLe of lLs name, Lhe lnsular corLex does noL form an lsland, lL ls a parL of Lhe conLlnuous sheeL of corLex and ls deeply burled only because of Lhe relaLlvely greaLer growLh of Lhe corLex around lL. neurons ln Lhe lnsular corLex are concerned wlLh vlsceral, auLonomlc, and LasLe funcLlons and are LhoughL Lo conLrlbuLe ln complex ways Lo lnLegraLlve braln funcLlons LhaL lmpacL emoLlon and soclal cognlLlon.
!"#$%"& ()%*"+$ ,* #-$ .%"/0 - Sprlng 2013
6 +%)A? B)(+%-&'+ C1. Whlch of Lhe followlng sLaLemenLs concernlng Lhe !"./#%& +-&!-+ ls mosL correcL? A. 1he cenLral sulcus LermlnaLes laLerally ln or very near Lhe longlLudlnal flssure. 8. 1he cenLral sulcus ls Lhe prlnclpal landmark LhaL dlvldes Lhe Lwo cerebral hemlspheres from one anoLher. C. 1he cenLral sulcus LermlnaLes medlally ln or very near Lhe laLeral (Sylvlan) flssure. u. 1he cenLral sulcus ls formed by gyral formaLlons LhaL harbor Lhe somaLlc sensory and moLor dlvlslons of Lhe cerebral corLex ln Lhe human braln. L. 1he cenLral sulcus ls malnly ln Lhe axlal plane of Lhe cranlum.
C2. Whlch Lwo lobes of Lhe cerebral hemlsphere feaLure Lhree ,%#%&&"&A &0.3*/-9*.%& 36#* on Lhelr laLeral aspecL? A. fronLal and Lemporal lobes 8. fronLal and parleLal lobes C. parleLal and occlplLal lobes u. Lemporal and occlplLal lobes L. Lemporal and parleLal lobes