Apostila Completa Bioquimica Rita Luz

Fccu|ccce ce Mecicinc Mcc|c ll.
l - 8iccu|micc
1

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Metobo||smo: ccnjunIc ce Iccc: c: (ecccce: cu|micc: inIec(ccc: nc meic ce|u|c(. cue
pc(Iicipc nc I(ccc ce mcI(ic e ene(cic ccm c meic exI(cce|u|c(.

Funes pr|nc|po|s do metobo||smo:
CLIenccc ce ene(cic cu|micc {mc|cu|c: nuI(ienIe: cu pc( cL:c(ccc ce |uz :c|c()
Ccnve(:cc cc: nuI(ienIe: exccenc: em pe(cu(:c(e: cc: ccmpcnenIe:
{mcc(cmc|cu|c: ce|u|c(e:): ccnve(:cc mc|ecu|c(. Lic::|nIe:e:
S|nIe:e ce mcc(cmc|cu|c: ce|u|c(e: {p(cIe|nc:. |ipicc:. ccicc: nuc|eicc:. c||cicc:.
eIc.)
Fc(mcccc e cec(cccccc ce Licmc|cu|c: ccm funcce: e:pecic|izccc:.
Exc(eccc ce meIcLc|iIc:
Fe:e(vc: meIcLc|icc:
LimiIccc: c|icccnic {f|cccc)
Ncc |imiIccc: I(icci|c|ice(ci: {Iecicc ccipc:c)
{exce::c ce c|icc:e I(cn:fc(mccc em I(icci|c|ice(ci:)

C|oss|||coo dos orgon|smos quonto s |ontes de motr|o e energ|o:

Or|gens do corbono e energ|o I|po de orgon|smo
Cc(Lcnc
CC2 /uIcI(cficc
Mc|cu|c: ccmp|exc: ccm cc(Lcnc HeIe(cI(cficc
Ene(cic
Luz :c|c( FcIcI(cficc
Fecccce: ce Cxiccccc-(ecuccc
c) mc|cu|c: c(ccnicc:
L) SuL:Icncic: inc(ccnicc:
CuimicI(cficc
c) cuimic-c(ccncI(cficc:
L) cuimic-|iIcI(cficc:

C|oss|||coo dos orgon|smos heterotro||cos quonto oce|too de e|ectres
/e(cLic: - uIi|izcm cxicnic {(e:pi(cccc)
/nce(cLic: - uIi|izcm cuI(c: mc|cu|c: cxicccc: {fe(menIcccc)
/nce(cLic: (e:I(iIc: - nuncc uIi|izcm cxicnic
Fccu|IcIivc: - cxicnic cu mc|cu|c: cxicccc:.

C|c|os de motr|o-|nterdependnc|o (s|ntro||o) dos orgon|smos v|vos
C: cuIcI(cficc: e c: heIe(cI(cficc: ccexi:Iem nc Lic:fe(c. inIe(cepencencc em
Ie(mc: ce mcI(ic e ene(cic
C: cuIcI(cficc: uIi|izcm CC2 pc(c fc(mc( Licmc|cu|c:. ccn:umicc: pe|c:
heIe(cI(cficc:. cue |iLe(Icm H2C e CC2 pc(c c meic
G(cnce pc(Ie cc: cuIcI(cficc: Im fcIc::|nIe:e. cLIencc c ene(cic cue p(eci:cm
cc |uz :c|c(. encucnIc cue c: heIe(cI(cficc: c cLIm pc( cxiccccc cc: mc|cu|c:
c(ccnicc:.
Ne:Ic: I(cn:fe(ncic:. hc ene(cic ncc cp(cveiIccc {cumenIc cc ene(cic cc meic).
encucnIc umc pecuenc pc(Ie uIi|izccc pc(c p(ccuzi( I(cLc|hc {ene(cic |iv(e).
C f|uxc ce ene(cic unici(eccicnc| e :em pc::iLi|iccce ce (ecupe(cccc. cevicc c
ccn:IcnIe ci::ipcccc {cumenIc c enI(cpic cc unive(:c)

Or|gem e ut|||zoo do energ|o qum|co ce|u|or - c|c|o do AIF

FcIc::|nIe:e cu
cxiccccc ce
nuI(ienIe:
Ene(cic
/DF + Fi
/IF
Ene(cic I(cLc|hc ce|u|c(
Fccu|ccce ce Mecicinc Mcc|c ll.l - 8iccu|micc
2

C /IF. ccnIinucmenIe (ecene(ccc cc /DF e Fi. funcicnc ccmc I(cn:pc(Iccc( e:pecic|
ce umc f(ccccc ce ene(cic |iv(e. enI(e (ecccce: ence(ccnicc: e exe(ccnicc:.

V|os metobo||cos
F(incipic: inIe(venienIe:
SuL:I(cIc:/meIcLc|iIc:/p(ccuIc:
Enzimc: {em :ecuencic: mu|IienzimcIicc:)
Ccenzimc:/ccfccIc(e:
F(incipci: cLjecIivc:
Fc(mcccc ce /IF
CLIenccc ce pcce( (ecuIc( {N/DFH + H
+
)
CLIenccc ce p(ecu(:c(e: mcc(cmc|ecu|c(e:

Si:Iemc: mu|IienzimcIicc:
Em cccc eIcpc ccIc|i:ccc pc( umc enzimc e:pec|ficc. :cc I(cn:fe(icc: c(upc
funcicnci:. cIcmc: cu mc|cu|c:
Em ce(c| c 1 eIcpc ccIc|i:ccc pc( umc enzimc (ecu|ccc(c

Ccmp|exiccce cc: :i:Iemc: mu|IienzimcIicc:
Si:Iemc :c|ve|: c: :uL:I(cIc: ce cimen:ce: mci: (ecuzicc: e muiIc mcvei:. ce:|cccm-
:e ce umc enzimc pc(c cuI(c. em :ecuncic. cI cLIenccc cc p(ccuIc finc|.
Ccmp|exc: mu|IienzimcIicc:: c: enzimc: ci::cciccc: :cc inccIivc:: em ce(c| c:
:uL:I(cIc: ccn:e(vcm-:e |icccc: cc :i:Iemc pc( |icccce: ccvc|enIe: cI c fc(mcccc
ce p(ccuIc.
Si:Iemc: c::cciccc: c e:I(uIu(c: :up(c-mc|ecu|c(e: {pc( exemp|c: memL(cnc:)

I(cn:pc(Iccc(e: ce e|ecI(ce:
Nc cxiccccc ce(cLic cc: mc|cu|c: ce nuI(ienIe: :cc (emcvicc: e|ecI(ce: ccm c:
:ecuinIe: ce:Iinc: pc::|vei::
1. CLIenccc ce /IF
2. CLIenccc ce pcce( (ecuIc( pc(c Lic::|nIe:e
Em cuc|cue( cc: cc:c:. c: e|ecI(ce: :cc veicu|ccc: {cc: mc|cu|c: inicic|menIe
(ecuzicc: pc(c c cceiIccc( finc|) pc( I(cn:pc(Iccc(e: e:pecici: {nuc|ecIicc: pi(ic|nicc:
e f|cvinc:).
A t|cn:|e|nc|c ce e|ect|ce: |ec|e:entc c t|cn:cc|te ce ene|g|c cun|cc.

F(incipci: I(cn:pc(Iccc(e: ce e|ecI(ce::
Em ce(cLic:e. c C2 ccIuc ccmc cceiIccc( finc| ce e|ecI(ce: I(cn:pc(Iccc: pe|c F/DH2
cu N/DH + H
+
e ce:Ie ve|cu|c: pc(c ccceic miIcccnc(ic| ce I(cn:pc(Ie ce e|ecI(ce:.
Fe|c f|uxc ce e|ecI(ce: cI(cv: ce:Ic ccceic ce(ccc c(cnce pc(Ie cc /IF ce|u|c(
{fc:fc(i|cccc cxiccIivc)

Mc|cu|c (ecuzicc e
-
F/DH2. N/DH + H
+
ccceic miIcccnc(ic| C2

C /IF ce(ccc (cpiccmenIe uIi|izccc em (ecccce: meIcLc|icc: ence(ccnicc:.
p(cp(ic: cc I(cLc|hc ce|u|c(: c ene(cic cu|micc I(cn:pc(Iccc pe|c /IF I(c:nfe(icc. c
pc( cc fc:fcIc Ie(minc|. pc(c mc|cu|c: cceiIccc(e: cue c::im cccui(em ene(cic
ccmp|emenIc(.

Mc|cu|c (ecuzicc
Mc|cu|c cxicccc
I(cn:pc(Iccc( cxicccc
I(cn:pc(Iccc( (ecuzicc
e|ecI(ce:
/ceiIccc( finc|
ce e|ecI(ce:
3

Fcce( (ecuIc(
/|m cc /IF. c|cumc: eIcpc: cc: Lic::|nIe:e: (ecue(em pcce( (ecuIc(. pc(c c
fc(mcccc ce :uL:Icncic: mci: (ecuzicc: cue c: pe(cu(:c(e:.
C: e|ecI(ce: :cc I(cn:pc(Iccc: cc: eIcpc: ccIcLc|icc: pc(c c: eIcpc: cncLc|icc:
cue ce|e: cepencem cI(cv: cc N/DF. Cccc mc|cu|c ce N/DF cxicccc
{N/DF
+
) I(cn:pc(Ic. pc( meccni:mc :eme|hcnIe cc cc N/D
+
. cci: e|ecI(ce:
exc|u:ivcmenIe uIi|izccc em Lic::|nIe:e: (ecuIc(c:: pe|c ccnI(c(ic. c N/D
+

uIi|izccc cuc:e uniccmenIe nc ce(cccc ce /IF.
Fc:e: Ge(ci: cc meIcLc|i:mc:
C ccIcLc|i:mc e c cncLc|i:mc cecc((em em 3 c(cnce: fc:e:. cue envc|vem
(e:pecIivcmenIe. c uIi|izcccc cu fc(mcccc ce c||cicc:. |ipicc: e p(cIe|nc:.

CcIcLc|i:mc:
Dec(cccccc exe(ccnicc pc( eIcpc:
Ccn:e(vcccc pc(cic| ce ene(cic |iv(e :cL c fc(mc ce /IF
Mc|cu|c: ccmp|exc: muiIc cife(enIe: mc|cu|c: :imp|e: ccmun:. c uIi|izc(
numc vic ccmum
/ncLc|i:mc:
S|nIe:e ence(ccnicc pc( eIcpc:
Ccn:umc ce /IF
Ccmpc:Ic: ce:Ic vic ccmum mcc(cmc|cu|c: {c||cicc:. p(cIe|nc: e |ipicc:)

CcIcLc|i:mc e cncLc|i:mc
Secuncic ce eIcpc: em :i:Iemc: mu|IienzimcIicc:
Fecu|cccc incepencenIe
Ccn:e(vcccc e:I(uIu(c| e I(cn:fe(ncic ene(cIicc :imu|Icnec:.
Ncc :cc inIei(cmenIe c (eve(:c
Lccc|izcm-:e em ccmpc(IimenIc: ce|u|c(e: ci:IinIc:.

Vic cnfiLc|icc: Im funcce: ccIcLc|icc: e cncLc|icc:.

Corocterst|cos d|nm|cos do metobo||smo
Ccc((e em eIcpc:. inIec(ccc: em :ecuncic: mu|IienzimcIicc: incepencenIe:
{mc: :imu|Icnec:) e ccm (ecu|cccc p(cp(ic
/ I(cn:fc(mcccc ce cccc p(ecu(:c( em um cu vc(ic: p(ccuIc: meciccc pc(
m|Iip|c: inIe(mecic(ic: {meIcLc|iIc:)
Cccc eIcpc ce I(cn:fc(mcccc meIcLc|icc ccIc|i:ccc pc( umc enzimc {ce
e:pecificiccce vc(icve|)
Simu|IcnecmenIe ccm c I(cn:fc(mcccc e:I(uIu(c| {ccvc|enIe). cccc eIcpc
meIcLc|icc pc::iLi|iIc I(cn:fe(ncic: ene(cIicc:.
Em ceIe(minccc: eIcpc: meIcLc|icc:. pc(Ie cc ene(cic cu|micc cc: meIcLc|iIc:
ccn:e(vccc :cL c fc(mc ce /IF. pc( :uc vez uIi|izccc em cuI(c: eIcpc:
e:pecici:.
Mc|cu|c: nuI(ienIe:
pcIencic|menIe
ene(cIicc:
F(ccuIc: ce Lic::|nIe:e
e ccmpcnenIe:
mc|ecu|c(e:
Energ|o L|vre
Mc|cu|c: :imp|e:
H2C. CC2
/ncLc|i:mc CcIcLc|i:mc
4

C cecu(:c cc meIcLc|i:mc em m|Iip|c: eIcpc: :ecuencici: ccnfe(e c(cnce
f|exiLi|iccce e ve(:cIi|iccce cc ccnjunIc
Cccc eIcpc :ujeiIc c (ecu|cccc incife(enciccc {cepencenIe cc ccncenI(cccc
cc: inIe(venienIe: ci(ecIc: e ccncicce: cc meic): cpenc: c|cumc: eIcpc: em
cccc :ecuncic :cc ccIc|i:ccc: pc( enzimc: e:pecificcmenIe (ecu|ccc(c: ce::c
:ecuncic.
DeIe(minccc: ccnjunIc: ce (ecccce: ccIcLc|icc: fc(mcm cic|c: meIcLc|icc:.
pc::iLi|iIcncc c (ecene(cccc {ccmc p(ccuIc) ce um :uL:I(cIc inicic|.

kegu|oo gero| do metobo||smo
kegu|oo ccnfe(e c(cnce f|exiLi|iccce c: (ecccce: meIcLc|icc: {c(cnce
ccpcciccce ce cccpIcccc c ncvc: :iIuccce:)
/ ccIiviccce meIcLc|icc ncc ccnI(c|ccc pe|c: ci:pcniLi|iccce: mcmenIcnec: em
nuI(ienIe: cu meIcLc|iIc:. mc: pe|c: nece::iccce: ene(cIicc: :cL c fc(mc ce /IF

Meccni:mc (ecu|ccc(e: cc meIcLc|i:mc
lnIe(ce|u|c(e:: neu(c| e hc(mcnc|
lnI(cce|u|c(e:: exp(e::cc cenIicc e mccu|cccc enzimcIicc {c|c:I(icc e ccvc|enIe)

/cIiviccce ce cccc (eccccc enzimcIicc {pH. ccncenI(cccc ce :uL:I(cIc:.
p(ccuIc: e ccfccIc(e:) - c|Ie(cccc cevicc c meccni:mc: pc:iIivc: cu neccIivc:.
/cIivcccc enzimcIicc: ccIivcccc ce zimccnic: {enzimc: cue nece::iIcm ce :e(
ce:cccificccc: pc(c :e(em funcicnci:)
CcncenI(cccc ce mc|cu|c: enzimcIicc: {cepence cc :|nIe:e - incuccc cu
(ep(e::cc cc exp(e::cc enzimcIicc - e cec(cccccc p(cIeicc)
/cIiviccce cc: enzimc: (ecu|ccc(c: {c|c:I(icc: cu ce mccu|cccc ccvc|enIe)
Ccmpc(Iiccc cc: vic:: c: memL(cnc: ccncicicncm c: I(ccc: ce mcI(ic enI(e
ccmpc(IimenIc: ci:IinIc: e fc(mcm ccmpc(IimenIc: {cue c(icincm c(ccienIe: ce
ccncenI(cccc e ce pH ccecuccc: c: ccIiviccce: cc meic)
E:Iccc ene(cIicc ce|u|c( - pcce :e( cvc|iccc pc( 2 |ncice:: cc(cc ene(cIicc e
pcIencic| ce fc:fc(i|cccc.
l:cenzimc:

Cc(cc ene(cIicc
F(e::upce c :i:Iemc /IF-/DF-/MF. cujc vc|c( ceIe(minccc pe|c ccnIecc ce c(upc:
fc:fcIc ce c|Ic ene(cic.

Cc(cc ene(cIicc = {/IF] + {/DF]
{/MF]+{/DF]+{/IF]
Em ecui||L(ic meIcLc|icc:
F(ccuccc ce /IF = uIi|izcccc ce /IF: cc(cc ene(cIicc = 0.

Cc(cc ene(cIicc < 0. - c: (ecccce:
ce(ccc(c: ce /IF :cc cce|e(ccc:: c:
(ecccce: uIi|izccc(c: ce /IF :cc
ciminu|cc:
Cc(cc ene(cIicc > 0. - c: (ecccce:
ce(ccc(c: ce /IF :cc ciminu|cc:: c:
(ecccce: uIi|izccc(c: ce /IF :cc
cce|e(ccc:

/: (ecccce: ce(ccc(c: ce /IF :cc iniLicc: pe|c cumenIc cc cc(cc ene(cIicc.
encucnIc cue c: (ecccce: cue uIi|izcm /IF :cc e:Iimu|ccc: pc( cc(cc ene(cIicc c|Ic.
FcIencic| cc fc:fc(i|cccc
FcIencic| ce fc:fc(i|cccc = {/IF]
{/IF]{Fi]
/cIiviccce
(e|cIivc
Fecccce:
exe(ccnicc:
Fecccce:
ence(ccnicc:
+ /IF
- /IF
Cc(cc ene(cIicc
5

/ c|u|c Ie(c IcnIc mci: ene(cic cucnIc mcic( fc( c pcIencic| ce fc:fc(i|cccc.

Metobo||smo nos d||erentes c|u|os

Hepofc|fos {mcnuIenccc cc c|icemic)
Ccnve(:cc ce cmincccicc: em c|icc:e - c|iccneccne:e
Ccnve(:cc ce ccicc: cc(cc: em cc(pc: ceIcnicc:
S|nIe:e ce ccicc: cc(cc: c pc(Ii( cc c|icc:e
Cxiccccc ce ccicc: cc(cc:
Cerebro
Cxiccccc cc c|icc:e e cc(pc: ceIcnicc:
Muscu|o esque|ef|co
Cxiccccc ce ccicc: cc(cc:. cc(pc: ceIcnicc:. cmincccicc: e c|icc:e
Ccn:e(vcccc ce c|icc:e {c|icccnic)
Fc(nece cmincccicc: pc(c c f|cccc
Ad|poc|fos
Fe:e(vc ce I(icci|c|ice(ci:

6

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||c
cco
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Metobo||smo dos G|c|dos

G||cicc: {ex: c|icc:e. f(uIc:e. cc|ccIc:e e mcnc:e)
Funccc Fc(nece( ene(cic {pc(cic|menIe ccn:e(vccc :cL c fc(mc ce /IF)
Ccmc /I(cv: ce (ecccce: (eccx. Mcximc cc (encimenIc em ce(cLic:e.

V|os de metobo||zoo dos g|c|dos
1. G||co||se (ou v|o de Embden-Meverho|): Secuncic cc (ecccce: cue cxiccm c
c|icc:e em pi(uvcIc {cu |ccIcIc)
2. C|c|o do c|do ctr|co: :ecuncic cc (ecccce: cm cue cccc (e:icuc cceIcIc
cxicccc em CC2. C p(ccuIc finc| {cxc|ccceIcIc) IcmLm um cc: :uL:I(cIc:
cc p(imei(c (eccccc. em cue ccc((e c :uc ccncen:cccc ccm c cceIi|-Cc/.
3. Fos|or||oo ox|dot|vo {c::cciccc cc I(cn:pc(Ie ce e|ecI(ce: pc(c c C2):
Fc:fc(i|cccc cc /DF em /IF. pc( cp(cveiIcmenIc ce c(cnce pc(Ie cc ene(cic
|iv(e cLIicc cu(cnIe c I(cn:pc(Ie {cI(cv: cc ccceic (e:pi(cIc(ic) ce e|ecI(ce:.
ce mc|cu|c: ccm pcIencic| ene(cIicc e|evccc pc(c c C2.
4. G||coneognese: :|nIe:e cc c|icc:e c pc(Ii( ce p(ecu(:c(e: ncc-c|icicicc:.
5. G||cognese: :|nIe:e cc c|icccnic. pc( pc|ime(izcccc cc: (e:|cuc cc c|icc:e
. G||cogeno||se: ceccmpc:iccc cc c|icccnic em c|icc:e
7. V|o dos |os|opentoses: Vic c|Ie(ncIivc c c|icc|i:e. pe(miIe c cxiccccc cc c|icc:e
CC2 e c ce(cccc ce pcce( (ecuIc( {e c:e: ccm 3 c 7 cc(Lcnc:)
8. V|o do c|do uron|co: IcmLm umc vic c|Ie(ncIivc. pe(miIe c cxiccccc cc
c|icc:e em ccicc c|icu(cnicc e penIc:e:.
. Metobo||smo do |rutose: Cxiccccc cc f(uIc:e em pi(uvcIc {cu |ccIcIc). em
ccnexcc ccm c c|icc|i:e
10. Metobo||smo do go|octose: Ccnve(:cc cc cc|ccIc:e em c|icc:e e :|nIe:e cc
|ccIc:e
11. Metobo||smo dos om|no-oses: Fc(mcccc cc: cminc-n:e: {c pc(Ii( ce c|icc:e e
cuI(c: c:e:). ccm pc:Ie(ic( cp(cveiIcmenIc nc :|nIe:e cc: c|iccp(cIe|nc: e
muccpc|i::ccc(icc:

7

G||co||se

bt|||dode metobo||co do g||cose
Hic(c|i:e enzimcIicc - cice:Icc
1. Am|do e g||cogn|o:
/mi|c:e: {Lccc e :ucc pcnc(ecIicc) (upIu(c cc: |icccce: c|icc:|cicc: u -1.4. Hic(c|i:e
IcIc| em ccceic: |inec(e: {cmi|c:e)
Am||ose Mo|tose + G||cose
/mi|c:e
G|icc:icc:e: {mucc:c inIe:Iinc|) (upIu(c cc: |icccce: c|icc:|cicc: u -1.
-f(uIc:icc:e cu Sccc(c:e {inIe:Iinc cc Hcmem) hic(c|i:e cc Sccc(c:e em G|icc:e e
F(uIc:e.
$ocorose Frutose + G||cose
Sccc(c:e
-cc|ccIc:icc:e cu LccIc:e {inIe:Iinc cc hcmem) hic(c|i:e cc LccIc:e em Gc|ccIc:e
e G|icc:e.
Loctose Go|octose + G||cose
Sccc(c:e

2. Ironsporte oct|vo ce c|icc:e cc n|ve| cc: c|u|c: inIe:Iinci: pc(c c :cncue pc(Ic| e
cepci: pc(c c ci(cu|cccc ce(c|.
3. G|icc:e c|icc:e--fc:fcIc {hexccinc:e: cu G|icccinc:e) c|icc:e |iv(e {G|icc:e--
fc:fcIc:e)
4. C ||godo c 1 c(ccc c Ie( c cpc(Iuniccce ce (emcve( c|icc:e cc :cncue pc(Ic|.
G|icemic c|Ic - (emcccc cc c|icc:e {pc( pc(Ie cc f|cccc). pc(c c: p(cce::c: ce
G|icccne:e e G|icc|i:e. cue ccn:cmem c|icc:e.
G|icemic Lcixc - fc(nece c|icc:e cc :cncue pe|c: p(cce::c: p(ccuIc(e: ce c|icc:e. c
G|icccenc|i:e e c Necc|icccne:e.

G|icc:e: p(incipc| hic(cIc ce cc(Lcnc cue cL:c(vicc nc inIe:Iinc e cp(cveiIccc pe|c:
c|u|c: cc cc(pc cue ce|c (eIi(cm ene(cic.
Exi:Iem Iipc: ce|u|c(e: cue cepencem cc c|icc:e ccmc fcnIe ene(cIicc em exc|u:ivc
{e(iI(cciIc:). :uL:Icncic|menIe {|euccciIc:. c|u|c: ne(vc:c:) cu em :ecuncc p|cnc
{hepcIcciIc)

G||co||se
/ ceccmpc:iccc {ccIcLc|icc) cc c|icc:e pcce ccc((e(:
Nc p(e:encc ce cxicnic {g||co||se oerob|o)
Nc cu:ncic ce cxicnic {g||co||se onoerob|o ou |ermentoo)
/ c|icc|i:e pe(miIe cc m:cu|c-e:cue|Iicc n|vei: Lc:IcnIe e|evccc: ce ccIiviccce.
me:mc ncc ci:pcncc ce cxicnic e pe(miIe cue Iecicc: ccm ccpcciccce c|icc||Iicc
:icnificcIivc :cL(evivcm c epi:ccic: cncxic: {cu:ncic ce C2).
/ fe(menIcccc ccc((e e:pcnIcnecmenIe {ccm ccenIuccc cLcixcmenIc cc
ene(cic |iv(e) e fc(mc /IF

I|pos de |ermentoo:
LccIicc: 1 mc| c|icc:e 2 mc| |ccIcIc
/|ccc|icc {em mic(cc(ccni:mc:) : 1 mc| c|icc:e 2 mc| eIcnc|

G||co||se oerob|o: Ncc hc ccumu|cccc ce |ccIcIc: um cc: p(ccuIc: {pi(uvcIc)
IcIc|menIe cxicccc em CC2 e H2C pe|c (e:pi(cccc

Lccc|izcccc: ciIc:c|

CLjecIivc: ccnve(:cc cc c|icc:e em pi(uvcIc cu |ccIcIc e cLIenccc ce /IF
8

Funcce: ce(ci: cc c|icc|i:e
Deccmpc:iccc cc c|icc:e em c|u|c: :em ccIiviccce meIcLc|icc {ex. c|cLu|c
ve(me|hc)
F(epc(cccc cc :ecuncic ce(cLic cc ccIcLc|i:mc cc c|icc:e
SuL:IiIui-:e c (e:pi(cccc. nc cLIenccc cc ene(cic cu(cnIe cu(Ic: pe(|ccc: em
cnce(cLic:e
F(ccuccc ce pcIencic| (ecuIc(
F(ccuccc ce pi(uvcIc ccmc pe(cu(:c( cc: ccicc: cc(cc: nc f|cccc e nc Iecicc
ccipc:c
F(ccuccc ce pe(cu(:c(e: cc: Lic::inIe:e: ce cmincccicc: e penIc:e:.

Meccni:mc: e cc(ccIe(|:Iicc: ce(ci:
Nc ccnve(:cc cc c|icc:e em |ccIcIc inIe(vm 11 enzimc: cife(enIe:. inIec(ccc: numc
:ecuncic mu|IienzimcIicc em :c|uccc nc ciIcp|c:mc. cue pc(Iicipcm em cive(:c:
Iipc: ce (eccccc:
I(cn:fe(ncic cc c(upc fc:fc(i|c {cinc:e:)
l:cme(izcccc
De:ic(cIcccc
C|ivccem c|ccc|icc
Fecuccc

ExcepIucncc c c|icc:e. pi(uvcIc e |ccIcIc. Iccc: c: (e:IcnIe: meIcLc|iIc: e:Icc
fc:fc(i|ccc: {pc( |icccce: :Ie( cu cnic(icc ccm 1 cu 2 F)

Funes dos grupos |os|oto:
Fc(mcm um ccmpc:Ic pc|c( ccm cc(cc neccIivc {c pH 7) c cue cificu|Ic c :uc
cifu:cc cI(cv: cc memL(cnc p|c:mcIicc
Ccn:IiIuem c(upc: ce unicc cu (eccnhecimenIc nc fc(mcccc cc: ccmp|exc:
enzimc-:uL:I(cIc
Fc::iLi|iIcm c ccn:e(vcccc cc ene(cic. inci:pen:cve| c fc(mcccc cc /IF

GLICLI$E - FA$E$

1 Fose preporotor|o do g||cose
Ccn:umc ce 2mc| /IF/mc| ce c|icc:e
F(ccuccc ce 1mc| ce f(uIc:e 1.Li:F/mc| ce c|icc:e

1. Fos|or||oo do g||cose - Fos|or||oo

/ c|icc:e enI(c nc: c|u|c: pc( cifu:cc fcci|iIccc c cue ncc pe(miIe c
ccumu|cccc nc c|u|c ce ccncenI(ccce: ce c|icc:e :upe(ic(e: c: exi:IenIe: nc
:cncue fc:fc(i|cccc cc c|icc:e pe|c enzimc hexoc|nose.

/ memL(cnc ce|u|c( impe(mecve| c c|icc:e--fc:fcIc. cue pcce pc( i::c :e(
ccumu|ccc nc c|u|c.
Ne:Ic eIcpc nece::c(ic c ut|||zoo de AIF ccmc cccc( ce fc:fcIc: cue
I(cn:fc(mccc em /DF.
/ (eccccc cccmpcnhccc pc( pe(cc :icnificcIivc ce ene(cic |iv(e :cL c fc(mc
ce cc|c(. c cue Ic(nc c (eccccc |rreversve| em ccncicce: fi:ic|ccicc:.
/ fc(mcccc ce c|icc:e c pc(Ii( cc (eccccc G|icc:e -F e ccuc ncc c (eccccc
inve(:c. cife(enIe. :encc ccIc|i:ccc pc( umc enzimc cife(enIe e em
ccmpc(IimenIc: cife(enIe:. C Fi |iLe(Iccc pc(c cuI(c meic. ncc vci fc:fc(i|c( /DF
pc(c fc(mc( c|icc:e e /IF.
G||cose + AIF G||cose -|os|oto + ADF
Hexccinc:e cu G|icccinc:e {Mc
2+
cu Mn
2+
)
9

Dest|nos do g||cose--|os|oto em oerob|ose:
Fc(mcccc ce I(ic:e:
Fc(mcccc ce penIc:e: - vic cc: fc:fcpenIc:e:
Fc(mcccc ce ccicc c|ucu(cnicc - vic cc ccicc c|ucu(cnicc
Hic(c|i:e G|icc:e :cncue {pe|c c|icc:e--fc:fcIc:e)
ExcecenIe c|icc:e 1-F c|icccnic c|icccne:e

Hexccinc:e:
umc cinc:e cue ccIc|izc c I(cn:fe(ncic cc c(upc fc:fc(i|c Ie(minc| cc /IF pc(c
um ccepIc( nuc|ecf||icc. Scc umc :uLc|c::e cc: I(cn:fe(c:e:.
Fc::ui umc e|evccc cfiniccce pc(c c c|icc:e fc:fc(i|cncc Iccc c c|icc:e cue
enI(c nc c|u|c. me:mc cucncc c: :uc: ccncenI(ccce: :cncu|nec: :cc Lcixc:.
umc enzimc regu|odoro. pci: so|re retro-|n|b|o o|ostr|co pe|c p(ccuIc ce:Ic
(eccccc: G|icc:e -fc:fcIc.
G|icccinc:e
e:pec|ficc cpenc: pc(c c D-c|icc:e {Iencc um KM muiIc :upe(ic( cc cc
hexccinc:e)
Exi:Ie cuc:e :c nc: hepcIcciIc:
/mLc: c: enzimc: (ecue(em Hc
2+
cu Mn
2+
{ccfccIc(e:). cue fc(mcm ccmp|exc: ccm
c /IF.

2. Converso do g||cose--|os|oto em |rutose--|os|oto - Isomer|zoo
Fc(c pcce( :e( uIi|izccc nc p(ccuccc ce ene(cic. c c|ucc:e--fc:fcIc p(imei(c
|somer|zodo c f(uIc:e--fc:fcIc pe|c |os|og||co|somerose.
Fc:icicncmenIc cc c(upc cc(Lcni| cc C2 p(epc(cccc pc(c c c|ivccem
Ccnve(:cc ce umc c|cc:e numc ceIc:e

Fc:fcc|icci:cme(c:e: ccIc|izc c i:cme(izcccc (eve(:|ve| cc c|icc:e--fc:fcIc em
f(uIc:e--fc:fcIc.

3. Fos|or||oo do |rutose--|os|oto - Fos|or||oo
/ f(uIc:e--fc:fcIc cepci: |os|or||odo c f(uIc:e-|.-Li:fc:fcIc. ccm cc:Ic ce /IF pe|c
|os|o|rutoc|nose.
E:Ie c pcnIc ce ncc-(eIc(nc ce:Ic vic meIcLc|icc: c pc(Ii( cc mcmenIc em cue c
c|ucc:e I(cn:fc(mccc em f(uIc:e-|.-Li:fc:fcIc jc ncc pcce :e( u:ccc em nenhumc
cuI(c vic - umc reoco |rreversve|.

Fc:fcf(uIccinc:e:
CcIc|i:c c I(cn:fe(ncic cc c(upc fc:fc(i|c cc /IF pc(c c f(uIc:e--fc:fcIc.
fc(mcncc f(uIc:e-1.-Li:fc:fcIc.
Fecue( Mc
2+
cue fc(mc ccmp|exc: ccm c /IF
umc enz|mo regu|odoro ccIc|izcncc c p(incipc| pcnIc ce (ecu|cccc cc c|icc|i:e.
Iem mccu|ccc(e: m|Iip|c:
oct|vodo pe|c exce::c /MF e /DF - p(ccuIc: ce c|ivccem cc /IF
|n|b|do pc( ccncenI(ccce: e|evccc: ce /IF e ciI(cIc.

G||cose -|os|oto Frutose -|os|oto
Fc:fcc|icci:cme(c:e
Frutose -Fos|oto + AIF Frutose 1,-8|s|os|oto + ADF
Fc:fcf(uIccinc:e {Mc
2+
)
10

2 Fose do g||co||se

4. C||vogem do |rutose-1,-b|s|os|oto - A|do||zoo
/ f(uIc:e-1.-Li:fc:fcIc c|ivccc pc(c c(icinc( 2 I(ic:e: fc:fcIccc: cife(enIe:: c
c|i(c|ceicc-3-fc:fcIc {c|cc:e) e c ciic(cxiceIcnc-fc:fcIc {ceIc:e).
/ c|ivccem ence(ccnicc. e:pcnIcnec nc :enIicc inve(:c. Fc(c cue e|c ccc((c nc
:enIicc ci(ecIc. c: p(ccuIc: ce (eccccc vcc :e( ccn:umicc: ncuI(c: (ecccce:.
E:Ic: cuc: mc|cu|c: :cc fcci|menIe inIe(ccnve(I|vei: pc( i:cme(izcccc ccIc|i:ccc
pe|c I(ic:e-fc:fcIc-i:cme(c:e.

/|cc|c:e {f(uIc:e-1.-Li:fc:fcIc c|cc|c:e):
CcIc|i:c umc (eccccc ce ccncen:cccc c|cc| (eve(:|ve|

5. Interconverso dos tr|oses-|os|oto
/penc: c c|ice(c|ceicc-3-fc:fcIc pcce :e( ci(ecIcmenIe cec(ccccc nc pc::c:
:uL:ecuenIe: cc c|icc|i:e
/ ciic(cxiceIcnc-fc:fcIc. (cpicc e (eve(:ive|menIe ccnve(Iicc em c|icec|ceicc-3-
fc:fcIc pe|c I(ic:e-fc:fcIc-i:cme(c:e.
Fc(IcnIc. Lc:Ic umc vic meIcLc|icc pc(c cc( c: cuc:. pc( e:Ic (czcc cue c
c|ucc:e--F fci i:cme(izccc c f(uIc:e--F: c c|ivccem cc c|ucc:e cc(ic c(icem c
cuc: mc|cu|c: Lc:IcnIe cife(enIe:. ce cci: e cucI(c cIcmc: ce cc(Lcnc.
(e:pecIivcmenIe. cue exici(icm cuc: vic: meIcLc|icc: cife(enIe: pc(c c :uc
cec(cccccc.

E:Ic (eccccc ccmp|eIc c fc:e p(epc(cIc(ic cc c|icc:e. / mc|cu|c ce hexc:e fci
fc:fc(i|ccc em C1 e C {ccn:umc ce 2 mc|cu|c: ce /IF) e cepci: c|ivccc pc(c fc(mc(
2 mc|cu|c: ce c|ice(c|ceicc-3-fc:fcIc. CuI(c: hexc:e:. ccmc c D-f(uIc:e. D-mcnc:e e
D-cc|ccIce:e IcmLm pccem :e( ccnve(Iicc: em c|ice(c|cei(c-3-fc:fcIc.

3 Fose do g||co||se
F(ccuccc ce 2mc| ce pi(uvcIc/mc| ce c|icc:e
F(ccuccc ce 4 mc| ce /IF/mc| ce c|icc:e/2mc| ce N/DH

lnc|ui pc::c: cc fc:fc(i|cccc ccn:e(vccc(c ce ene(cic em cue c|cumc cc ene(cic |iv(e
cc c|icc:e ccn:e(vccc :cL c fc(mc ce /IF. / ccnve(:cc ce 2 mc|cu|c: ce
c|ice(|cceicc-3-F em 2 mc|cu|c: ce pi(uvcIc cccmpcnhccc pe|c fc(mcccc ce 4
mc|cu|c: ce /IF c pc(Ii( ce /DF. CcnIucc. c ccnhc ce /IF/mc|cu|c: c|icc:e
cec(ccccc cpenc: 2. pc(cue 2 /IF fc(cm inve:Iicc: nc fc:e p(epc(cIc(ic cc
c|icc|i:e pc(c fc:fc(i|c( c: cuc: exI(emiccce: cc mc|cu|c ce hexc:e.

. Ox|doo do g||cero|de|do-3-F em 1,3-b|s|os|og||ceroto - ox|doo e |os|or||oo

C: c|ce|cc: Im pcIencici: ce cxiccccc-(ecuccc Lc:IcnIe Lcixc: {ce(cc ce -00
c -500 mV) e:pcnIcneiccce cc (eccccc ce cxiccccc cc c|ice(c|ce|cc-3-
fc:fcIc pe|c N/D
+
{E=-320 mV).
Umc vez cue c: c|u|c: ccnIm cucnIiccce: |imiIccc: ce N/D
+
. c c|icc|i:e pc(c(ic
(cpiccmenIe :e c N/DH fc(mccc ne:Ic eIcpc ncc fc::e ccnIinucmenIe cxicccc.
Fc( fc:fc(c|i:e ccicicnccc um fc:fcIc inc(ccnicc {Fi).

G|ice(c|ceicc-3-fc:fcIc-ce:ic(ccenc:e
CcIc|i:c c cxiccccc cc c|ice(c|ce|cc-3-fc:fcIc c 1.3-8i:fc:fcc|ice(cIc
Frutose 1,-b|s|os|oto
D|h|drox|cetono |os|oto
G||cero|de|do 3-|os|oto
/|cc|c:e I(ic:e fc:fcIc
i:cme(c:e
11

N/D
+
cepencenIe. I(cn:fc(mcncc-:e e:Ie em N/DH + H
+
. pc( I(cn:fe(ncic cc:
ecuivc|enIe: (ecuIc(e: (emcvicc: nc cxiccccc.

Ne:Ic (eccccc. c c(upc c|ce|cc em C1 I(cn:fc(mc-:e em cci|-fc:fcIc {cnic(icc mi:Ic
cc ccicc fc:fc(icc e ccicc cc(Lcx||icc) ccnfe(incc c mc|cu|c um e|evccc pcIencic|
ene(cIicc. Fep(e:enIc umc fc:fc(i|cccc c n|ve| cc :uL:I(cIc. em cue c p(ccuIc
(e:u|IcnIe ccnIm cci: c(upc: fc:fcIc ju:Icpc:Ic:. cccc um ccm cc(cc neccIivc c
e|evccc cen:iccce ce e|ecI(ce: junIc cc cxicnic cc |iccccc cnic(icc.

7. Irons|ernc|o do grupo |os|or||o do 1,3-b|s|os|og||ceroto poro o ADF

/ con||guroo 1,3-b|s|os|og||ceroto fcvc(ece e:Ic eIcpc:
/pc: c hic(c|i:e. c (ec::ccicccc cc: p(ccuIc: cificu|Iccc pe|c: fc(cc: ce
(epu|:cc {cc(cc neccIivc) cue ccnIm.
C: e|ecI(ce: cc |iccccc cnic(icc Iencem pc(c pc:icce: mci: e:Icvei: nc: c(upc:
fc:fcIc e cc(Lcx||icc.
Em ccn:ecuncic ce:Ie pcIencic| ene(cIicc c c(upc fc:fc(i|c C1 {cc(Lcxi|c)
I(cn:fe(icc pc(c c /DF ccncc c(icem cc 3-|os|og||ceroto.
/ :c|cc cc c(upc fc:fcIc cc c(icem c e:pcie: muiIc mci: e:IcLi|izccc: pc(
(e::cncncic

Fc:fcc|ice(cIc-cinc:e:
I(cn:fe(e c c(upc fc:fcIc cc 1.3-Li:fc:fcc|ice(cIc pc(c c /DF. fc(mcncc /IF e
cicc 3-fc:fcc|ic(icc.
umc enzimc c|c:I(icc. ccIivccc pe|c N/D
+
{cccpe(cIiviccce neccIivc)
De:fc:fc(i|c c :uL:I(cIc. I(cn:fe(incc um c(upc fc:fcIc pc(c c /DF

C: pc::c: e 1 em ccnjunIc ccn:IiIuem um p(cce::c ccp|ccc( ce ene(cic. em cue
1.3-Li:fc:fcc|ice(cIc c inIe(mecic(ic ccmum: fc(mccc nc 1 (eccccc {cue :e(ic
ence(ccnicc cucncc i:c|ccc). e c :eu c(upc fc:fcIc I(cn:fe(icc pc(c c /DF nc 2
(eccccc {fc(IemenIe exe(ccnicc).

Nc pc::c
AG = AG
0
+ FI |n {1.3-Li:fc:fcc|ice(cIc]{N/DH]
{c|ice(c|ceicc -3F] {Fi] {N/D
+
]

Nc (eccccc 7. ccn:umincc 1.3-Li:fc:fcc|ice(cIc p(ccuzicc em . ciminui c :uc
ccncenI(cccc. fczencc ccm cue AG :ejc muiIc neccIivc.
/ fc(mcccc ce /IF. pc( I(cn:fe(ncic ce um c(upc fc:fc(i|c ce um :uL:I(cIc ccmc c
1.3-Li:fc:fcc|ice(cIc (efe(icc ccmc fc:fc(i|cccc cc n|ve| cc :uL:I(cIc.
G||cero|de|do 3-|os|oto + F| + NAD
+
1,3-8|s|os|og||ceroto + NADH + H
+

G|ice(c|ceicc 3-fc:fcIc ce:ic(ccenc:e
1,3 - 8|s|os|og||ceroto + ADF 3-|os|og||ceroto + AIF
Fc:fcc|ice(cIc cinc:e
G3F + ADF + F| + NAD
+
2FG + AIF + NADH + H
+
{AG
0
= -12.5 Kj/mc|)
12

8. Converso do 3-|os|og||ceroto em 2-|os|og||ceroto

Fc:fcc|ice(cIcmuIc:e:
CcIc|i:c c muccncc (eve(:|ve| cc c(upc fc:fc(i|c enI(e C2 e C3 cc c|ice(cIc.
MuIc:e - enzimc: cue ccIc|izcm c I(c:nfe(ncic ce um c(upc funcicnc| ce umc
pc:iccc pc(c c cuI(c nc me:mc mc|cu|c {:uLc|c::e cc: i:cme(c:e:).
Fc(c e:Ic( ccIivc nece::iIc ce (eceLe( um c(upc fc:fcIc cc 2.3-Li:fc:fcc|ice(cIc
{:inIeIizccc c pc(Ie) cue funcicnc ccmc inIe(mecic(ic:

. Des|drotoo do 2-|os|og||ceroto em |os|oeno|p|ruvoto

/pe:c( cc 2-fc:fcc|ice(cIc e cc fc:fcenc|pi(uvcIc ccnIe(em cp(cximcccmenIe c
me:mc cucnIiccce IcIc| ce ene(cic. c pe(cc cc mc|cu|c ce ccuc pe|c 2-
fc:fcc|ice(cIc p(cvccc c (eci:I(iLuiccc ce ene(cic cenI(c cc mc|cu|c.
cumenIcncc c ene(cic |iv(e pcc(cc cc hic(c|i:e cc c(upc fc:fcIc.
/ ce:ic(cIcccc vci cumenIc( c pcIencic| ce I(cn:fe(ncic cc c(upc F

10. Irons|ernc|o do grupo |os|or||o de |os|oeno|p|ruvoto poro o ADF
/ ce:ic(cIcccc cue c(icinc c fc:fcenc|pi(uvcIc e|evc ccenIucccmenIe c
pcIencic| ce I(cn:fe(ncic cc c(upc fc:fc(i|c. cevicc c cxiccccc-(ecuccc
inI(cmc|ecu|c(e: {c: e|ecI(ce: ccncenI(cm-:e mci: junIc cc C2. cevicc c
fc(mcccc cc |iccccc cup|c e ciminuiccc cc ci:Icncic ce |iccccc ce C2 ccm
cIcmc: envc|venIe:).
Fc( cp(cximcccc (e|cIivc cc c(upc fc:fcIc e cc(Lcxi|c {ccm cc(cc neccIivc).
cumenIc c Ien:cc inI(cmc|ecu|c(. fcvc(cve| c hic(c|i:e. Dccui (e:u|Ic c cecncic
cc c(upc fc:fc(i|c cc /DF. nc (eccccc em cue :e fc(mc c pi(uvcIc.

Fi(uvcIc-cinc:e:
c 3 enzimc (ecu|ccc(c cc c|icc|i:e
:ujeiIc c mccu|cccc ccvc|enIe pc( vc(ic: efecIc(e: {ex. iniLicc pe|c /IF e
ciI(cIc)
/ (eccccc cue e:Ic enzimc ccIc|i:c e::encic|menIe i((eve(:|ve|. em ccncicce:
inI(cce|u|c(e:.

3-|os|og||ceroto 2-|os|og||ceroto
Fc:fcc|ice(cIcmuIc:e {Mc
2+
)
2.3-8FG 3-FG 2-FG 2.3-8FG
+ +
2-Fos|og||ceroto Fos|oeno| p|ruvoto + H2O
Enc|c:e
Fos|oeno|p|ruvoto + ADF F|ruvoto + AIF
Fi(uvcIc cinc:e {Mc
2+
. K
+
)
13

keoco g|obo|:

G|icc:e + 2Fi + 2/DF + 2N/D
+
2 pi(uvcIc + 2/IF + 2N/DH + H
+
+ 2H2C

8o|ono energt|co no converso do g||cose em p|ruvoto

keoces Vor|oo AIF/g||cose
G|icc:e c|icc:e -fc:fcIc -1
F(uIc:e -F f(uIc:e 1. -8i:fc:fcIc -1
2x{1.3-cifc:fcc|ice(cIc 3-fc:fcc|ice(cIc] +2
2x{Fc:fcenc|pi(uvcIc pi(uvcIc] +2
IcIc| +2

Conc|uso do g||co||se
Em ccncicce: ce(cLic:. c pi(uvcIc cxicccc c cceIcIc. cue enI(c nc cic|c cc
ccicc c|I(icc e cxicccc c CC2 e H2C. :encc c N/DH fc(mccc pe|c ce:ic(ccencccc
cc c|ice(c|ceicc 3-F (ecxicccc em N/D
+
pe|c pc::ccem cc: :eu: e|ecI(ce: cc C2 nc
(e:pi(cccc miIcccnc(ic|.
CcnIucc. em ccncicce: ce hipcxic. ccmc em m:cu|c: e:cue|Iicc: muiIc
ccIivc:. c N/DH ce(ccc nc c|icc|i:e ncc pcce :e( (ecxicccc pe|c C2. / pc(ccem cc
(ecene(cccc cc N/D
+
ceixc(ic c c|u|c :em cceiIccc(e: ce e|ecI(ce: pc(c c
cxiccccc ce G 3-F. c: c(ccni:mc: mci: mcce(nc: cccui(i(cm c ccpcciccce ce
(ecene(c( c N/D
+
cu(cnIe c c|icc|i:e cnce(cLic. I(cn:fe(incc e|ecI(ce: cc N/DH pc(c
fc(mc( um p(ccuIc finc| (ecuzicc. ccmc c |ccIcIc cu c eIcnc|.

Fermentoo |ct|co
Cucncc c: Iecicc: cnimci: ncc :cc cLc:Iecicc: ccm C2 :uficienIe pc(c :upc(Ic( c
cxiccccc ce(cLic cc pi(uvcIc e N/DH p(ccuzicc: nc c|icc|i:e. c N/D
+

(ecene(ccc cc N/DH pe|c (ecuccc cc pi(uvcIc c |ccIcIc
Nc c|icc|i:e. c ce:ic(ccencccc ce 2 mc|cu|c: ce G 3-F ccnve(Ie 2 mc|cu|c: ce
N/D
+
em N/DH. Ccmc c (ecuccc ce 2 mc|cu|c: ce pi(uvcIc c 2 mc|cu|c: ce
|ccIcIc (ecene(c 2 mc|e: ce N/D
+
. c p(cce::c c|cLc| Lc|cnceccc e pcce
ccnIinuc( incefiniccmenIe.
/ fe(menIcccc ccc((e ccm f(ecuncic nc: c|u|c: mu:cu|c(e: pc(cue c
ve|cciccce cc c|icc|i:e excece c Icxc meIcLc|icc cc cic|c ce K(eL: e cc
(e:pi(cccc.

Fermentoo o|coo||co
/: |evecu(c: e cuI(c: mic(cc(ccni:mc: fe(menIcm c c|icc:e c eIcnc| e CC2. em vez ce
|ccIcIc. / c|icc:e ccnve(Iicc c pi(uvcIc pe|c c|icc|i:e. e c pi(uvcIc ccnve(Iicc c
eIcnc| e CC2 num p(cce::c ccm 2 pc::c:.
1 Fc::c: c pi(uvcIc ce:cc(Lcxi|ccc numc (eccccc i((eve(:|ve| ccIc|izccc pe|c
pi(uvcIc ce:cc(Lcxi|c:e. cue nece::iIc ce Mc
2+
e ce umc ccenzimc fc(IemenIe
|icccc. c IFF {Iicminc pi(cfc:fcIc).
2 Fc::c: c cceIi|c|ce|cc (ecuzicc c eIcnc| pe|c ccccc cc ce:ic(ccenc:e c|ccc|icc.
:encc c pcce( (ecuIc( fc(necicc pe|c N/DH ce(ivccc cc ce:ic(ccencccc cc G 3-F.

Em cmLc: c: Iipc: ce fe(menIcccc ncc cxi((ecuccc |icuicc. pc(cue c N/D
+
. vci :e(
uIi|izccc nc c|icc|i:e.
2 F|ruvoto + 2NADH 2 Loctoto + 2 NAD
+

De:ic(ccenc:e LccIicc
F|ruvoto + H
+
Aceto|de|do + CO2
Fi(uvcIc ce:cc(Lcxi|c:e {IFF)
Aceto|de|do + NADH Etono| + NAD
+

De:ic(ccenc:e c|ccc|icc
14

A|ternot|vos metobo||cos
Vic: c|Ie(ncIivc pc(c c cihic(cxi-cceIcnc fc:fcIc
Fcce:
Scf(e( c ccccc cc enzimc I(ic:e fc:fcIc i:cme(c:e :encc ccnve(Iicc em
c|ice(c|ceicc 3-F
Fc(Iicipc( nc vic cc: fc:fcpenIc:e: {em cc:c ce exce::c ce c|icc:e p(cvccc c
:|nIe:e ce ||picc: - I(icci|c|ice(ci:)
Fcce :e( ccnve(Iicc em G|ice(c| 3 fc:fcIc {pe(cu(:c( |ipicicc) numc
(eccccc(eve(:|ve| ccIc|izccc pe|c c|ice(c| 3-fc:fcIc ce:ic(ccenc:e em cue c
N/DH cxicccc c N/D
+
:

Vic c|Ie(ncIivc pc(c c 1.3 - Li:fc:fcc|ice(cIc
Fcce:
Fc(mc( ci(ecIcmenIe 3-fc:fcc|ice(cIc {e :|nIe:e ce /IF)
I(cn:fc(mc(-:e em 2.3-Li:fc:fcc|ice(cIc.
Nc: e(iI(cciIc: exi:Iem ccncenI(ccce: e|evccc: ce 2.3-Li:fc:fcc|ice(cIc. muiIc
:upe(ic(e: c: cc: (e:IcnIe: Iipc: ce|u|c(e:.
De:Ic pc(Iicu|c(iccce (e:u|Ic cue c 2.3-Li:fc:fcc|ice(cIc exe(ce um efeiIc
mccu|ccc( impc(IcnIe nc cfiniccce cc hemcc|cLinc pc(c c cxicnic e
ccn:ecuenIemenIe nc I(cn:pc(Ie e I(ccc: ce cxicnic enI(e c :cncue e c: Iecicc:.
C 2.3-Li:fc:fcc|ice(cIc :inIeIizccc c pc(Ii( cc 1.3-Li:fc:fcc|ice(cIc e
pc:Ie(ic(menIe I(cn:fc(mccc em 3-fc:fcc|ice(cIc ce:Ie mccc u|Icpc::cncc-:e c
eIcpc ccIc|izccc. nc c|icc|i:e. pe|c fc:fcc|ice(ccinc:e.
/ I(cn:fc(mcccc cc 2.3-Li:fc:fcc|ice(cIc ccIc|i:ccc pc( umc p(cIe|nc ccIc||Iicc
mu|IifccIc(ic| {2.3-Li:fc:fcc|ice(cIc-:inIc:e) cue ccIuc ccmc muIc:e e fc:fcIc:e.
/: cc(cc: neccIivc: cc 2.3-Li:fc:fcc|ice(cIc unem-:e c: cc(cc: pc:iIivc: cc: 2
ccceic: ce hemcc|cLinc. cI(cincc-c: e cumenIcm. c::im. c expu|:cc ce cxicnic
pc(c c: Iecicc: {ce:|cccmenIc cc cu(vc ce ci::ccicccc cc hemcc|cLinc pc(c c
ci(eiIc). c cue um Leneficic em :iIuccce: ce fc|Ic ce cxicnic

C ccn:umc ce c|icc:e e c c|icc|:e :cc ce(cc ce 10x mci: (cpicc: em Iumc(e:
:c|icc: cc em cue Iecicc: ncc ccnce(|cenc:. Ccmc c: c|u|c: Iumc(ci: :cf(em hipcxic.
cepencem cc f(emeIcccc pc(c c p(ccucc cc mcic(ic cc /IF. UIi|izcm mci: c|icc:e
cue c: c|u|c: nc(mci:. ccnve(Iencc-c em pi(uvcIc e cepci: em |ccIcIc pc(c (ecic|c(
c N/DH. cue :e vci ccumu|cncc ne:Ic: c|u|c:.
D|h|drox|cetono |os|oto + NADH G||cero| 3-|os|oto + NAD
+

G|ice(c| 3-fc:fcIc ce:ic(ccenc:e
3-8i:fc:fcc|ice(cIc
cinc:e
G|ice(c|ceicc 3-F
1.3-8i:fc:fcc|ice(cIc
3-Fc:fcc|ice(cIc
2-Fc:fcc|ice(cIc
NAD
+
N/DH + H
+

Fi
AIF
/DF
G|ice(c|ceicc 3-F
De:icc(cenc:e
Fc:fcc|ice(cIccinc:e
Fc:fcc|ice(cIcmuIc:e
1.3-8i:fc:fcc|ice(cIc muIc:e
fc:fcIc:e
Fi
Fi
Fi(uvcIc
H2C
15

G|icc:e
G|icc:e -fc:fcIc
G|icc:e -fc:fcIc:e G|icccinc:e
H2C
Fi /IF
/DF
kEGbLAO DA GLICLI$E
Funcce: cc vic c|icc||Iicc:
Ge(c( /IF
Fc(nece( L|ccc: ce ccn:I(uccc pc(c (ecccce: ce :|nIe:e
Fc( i::c c ccnve(:cc ce c|icc:e em pi(uvcIc cc-:e ccn:ccnIe e::c: 2 nece::iccce: -
nece::iccce ce ccnI(c|c cI(cv: ce 3 enzimc: {hexccinc:e. fc:fcf(uIccinc:e e
pi(uvcIc-cinc:e) (ecu|ccc(c:
Enzimc: (ecu|ccc(c::
Scc ccnI(c|ccc: pc( efecIc(e: c|c:I(icc: cu pc( mccificcccc ccvc|enIe.
CcIc|izcm eIcpc: i((eve(:|vei:

EmLc(c c enzimc mci: impc(IcnIe :ejc :emp(e c 1 nc vic meIcLc|icc. nc c|icc|i:e c
|os|o|rutoc|nose mci: impc(IcnIe cue c hexoc|nose. pc(cue:
Sc ccn:ecuimc: iniLi( c hexccinc:e :e c p(imei(c iniLi(mc: c fc:fcf(uIccinc:e.
/ (ecu|cccc mci: eficcz :e fc( (ec|izccc cI(cv: ce umc enzimc cue ccIc|i:e
umc (eccccc :em (eIc(nc. cu :ejc. cujc p(ccuIc Ienhc :c um ce:Iinc. c cue ncc
cccnIece ccm c hexccinc:e. pci: c c|icc:e -p ncc um inIe(mecic(ic exc|u:ivc
cc c|icc|i:e. :cf(e vc(ic: ce:Iinc: {vic cc: fc:fcpenIc:e:. c|icccne:e) ncc :e
:ucecencc c me:mc ccm c f(uIc:e-1.-Li:fc:fcIc.

/ 1 (eccccc |imiIcIivc exc|u:ivc cpenc: ce c|icc|i:e ccIc|izccc pe|c
fc:fcf(uIccinc:e {eIcpc ccmp(cmeIicc cu (ecu|ccc(c") eIcpc mci: impc(IcnIe
pc(c c ccnI(c|e cc vic meIcLc|icc.
/ ve|cciccce cc ccnve(:cc cc :uL:I(cIc inicic| nc p(ccuIc finc| ccncicicnccc
pe|c eIcpc mci: |enIc cc vic {eIcpc mc(cc pc::c)
Gene(iccmenIe pcce cize(-:e cue c (ecu|cccc cc c|icc|i:e mccu|ccc pe|c
e:Iccc ene(cIicc ce|u|c( {/IF//DF) e ci:pcniLi|iccce em pe(cu(:c(e: pc(c c:
Lic::|nIe:e:. Cucncc cmLc: c: ccncicce: e:Icc ceficienIe:. Iccc: c: enzimc:
(ecu|ccc(c: :cc ccIivccc:. :ucecencc c inve(:c cucncc exi:Ie exce::c ce cc(Lcnc
e/cu ene(cic.

Hexoc|nose:
CcIc|i:c c fc:fc(i|cccc cc c|icc:e c c|icc:e--fc:fcIc
/ hexccinc:e iniLicc pe|c p(ccuIc cc (eccccc c|icc:e--fc:fcIc. Se c
meIcLc|izcccc cc c|icc:e--fc:fcIc menc( cue c :uc :|nIe:e. e:Ic ccumu|c-:e
iniLincc c hexccinc:e. nc enIcnIc i:Ic :c cccnIece cucncc c fc:fcf(uIccinc:e e:Ic
iniLicc.
Cucncc c hexccinc:e iniLicc enI(c em ccccc. nc f|cccc. c c|icccinc:e
{i:cenzimc cc hexccinc:e).
/ iniLiccc IcmLm pcce ccc((e( cucncc :e ve(ificcm e|evccc: vc|c(e: ce /IF.
Fcce :e( ccIivccc cucncc hc um e|evccc n|ve| ce c|icc:e.
lniLicc(e:: c|icc:e -fc:fcIc: /IF
/cIivccc(: c|icc:e

G|icccinc:e:
Iem um Km Lc:IcnIe mci: e|evccc pc(c c c|icc:e
Ncc iniLicc pe|c p(ccuIc finc| {c|icc:e -fc:fcIc)
iniLicc pe|c f(uIc:e--F
ccIivccc pe|c f(uIc:e-1-F
CcnI(iLui pc(c mcnIe( ccn:IcnIe: c:
n|vei: ce c|icemic: ccIuc cucncc c
ccncenI(cccc ce c|icc:e :cncu|nec
cIince e|evccc: vc|c(e:.
Nc f|cccc c ccIiviccce cc
c|icccinc:e cpc:Ic c cc c|icc:e--
fc:fcIc:e. cue ccmLinccc: fc(mcm
um cic|c fIi|.
16

Fos|o|rutoc|nose
CcIc|i:c c I(cn:fe(ncic cc c(upc fc:fc(i|c cc /IF pc(c c f(uIc:e--fc:fcIc.
fc(mcncc f(uIc:e-1.-Li:fc:fcIc.
un||unc|ono| pci: inccpcz ce ccIc|i:c( c (eccccc inve(:c cue :e efecIuc nc
necc|icccne:e pe|c ccccc cc f(uIc:e-l.-Li:fc:fcIc:e.
o|ostr|co: pc::ui vc(ic: ccIivccc(e: e iniLicc(e::

lniLicc(e::
/IF
Sencc c /IF um p(ccuIc finc| cc c|icc|i:e.
cucncc n|vei: e|evccc: ce /IF nc c|u|c
:inc|izcm cue c /IF e:Ic c :e( p(ccuzicc mci:
(cpicc cc cue c :e( ccn:umicc. c /IF |icc-:e
cc cenI(c c|c:I(icc cc fc:fcf(uIccinc:e.
ciminuincc c cfiniccce ce:Ic pc(c c f(uIc:e -
fc:fcIc.
C /DF e /MF. cujc ccncenI(cccc cumenIc c
mecicc cue c /IF ccn:umicc. ccIucm
c|c:Ie(iccmenIe pc(c |iLe(Ic( c iniLiccc pe|c
/IF.
H
+
{pH Lcixc):
/ pc(Ii( ce ce(Ic: vc|c(e: ce pH cc-:e c
iniLiccc cc fc:fcf(uIccinc:e.
/ccnIece muiIc cucncc :e (ecc((e c
fe(menIcccc |ccIicc pc(c umc cLIenccc mci:
(cpicc ce /IF.
C LccIcIc pc( :e( ccicc. Lcixc c pH.
Exi:Ie um :i:Iemc p(cIecIc( - :c|cc ccnjunIc ce
|ccIcIc e p(cIcc.
/ fe(menIcccc ce::c cucncc ncc :c c
ccncenI(cccc ce /IF :uficienIe ccmc
IcmLm c umc e|evccc ccncenI(cccc ce H
+

{Lcixc cemc:iccc c pH)
CiI(cIc
um inIe(mecic(ic nc cxiccccc ce(cLic cc
pi(uvcIc e IcmLm funcicnc ccmc (ecu|ccc(
c|c:I(icc
/ um cumenIc cc ccncenI(cccc ce ciI(cIc
cc((e:pcnce um cumenIc cc efeiIc iniLiIc(ic
cc /IF
Iecicc: cue uIi|izem p(efe(encic|menIe cc(pc:
ceIcnicc: e ccicc: cc(cc: cc(e:cem c
ccncenI(cccc ce ciI(cIc ciIcp|c:mcIicc
Um c|Ic n|ve| ce ciI(cIc imp|icc IcmLm um
c|Ic n|ve| ce /IF.
F(uIc:e 1. Li:fc:fcIc - iniLiccc pe|c p(ccuIc finc|
{feec-Lcck neccIivc)

/cIivccc(e:
F(uIc:e 2. Li:fc:fcIc
/umenIc c cfiniccce cc enzimc pc(c c f(uIc:e -F
Diminui c efeiIc iniLicc( cc /IF
/ ccIivcccc pe|c F(uIc:e 2. Li:fc:fcIc fcz pc(Ie cc (ecu|cccc ccvc|enIe
c::cciccc c fc:fcf(uIccinc:e.

17

/MF e Fi
/IF + /DF + /MF ccn:IcnIe em ccncicce: fi:ic|ccicc:
{/IF]>>{/DF]>>{/MF] umc pecuenc vc(icccc nc {/IF] ccu:c umc c(cnce
vc(icccc cc {/MF] l::c fcz ccm cue c /MF :ejc um cpIimc incicccc( cc e:Iccc
ce ene(cic cc c|u|c e um efecIc cc ccIiviccce cc fc:fcf(uIccinc:e
/cIivc c fc:fcf(uIccinc:e e iniLe c f(uIc:e-1.-Li:fc:fcIc:e pc(cue c ccccc ce:Ic.
junIcmenIe ccm c fc:fcf(uIccinc:e p(cvccc um cic|c fIi|.

EfeiIc Fc:Ieu(
Nc p(e:encc ce cxicnic p(ccuz-:e mci: /IF cue em ccncicce: cne(cLic:. |ccc c
f|uxc c|icc||Iicc em cnce(cLic:e e menc(.
Fc(IcnIc cucncc c cxicnic inI(ccuzicc nc :i:Iemc c /IF cumenIc. ciminuincc
/MF e Fi. iniLincc-:e c::im c fc:fcf(uIccinc:e e ccn:ecuenIemenIe c vic
c|icc||Iicc.
Fc(c cLIe( c me:mc (encimeInc ene(cIicc nece::c(ic muiIc mci: c|icc:e em
cnce(cLic:e cue em ce(cLic:e. E cnce(cLic:e c ccn:umc ce c|icc:e Lc:IcnIe
e|evccc. |cncc cucncc enI(e cxicnic nc :i:Iemc c /IF :cL(e exc(LiIcnIemenIe.

F|ruvoto-c|nose
CcIc|i:c c |Iimc (eccccc cc vic. c ccnve(:cc cc ccicc FEF em ccicc pi(vicc.

lniLicc(e:
/IF: |icc-:e cc cenI(c c|c:I(icc cc pi(uvcIc-cinc:e
CiI(cIc//|cninc: pe(cu(:c(e: pc(c Lic::|nIe:e: e|evccc:
/cIivccc(e::
F(uIc:e-1.-Li:fc:fcIc: cucncc em exce::c ccIivc inci(ecIcmenIe c pi(uvcIc-cinc:e
pc(c cue c c|icc|i:e :e c ccIivcmenIe.

Scf(e IcmLm (ecu|cccc ccvc|enIe
Fc:fc(i|ccc: - ccIivc
De:fc:fc(i|ccc: + ccIivc

Necess|dode regu|oo o|ostr|co e tombm covo|ente
Nc (ecu|cccc c|c:I(icc vc(ic: enzimc: e:Icc c::cciccc:. c cue p(cmcve umc
mcic( cmp|ificcccc cc :inc|
8c:Ic um e:I|mu|c pc(c ce:encccec( Iccc um p(cce::c em cc:ccIc
Hc c::im um mcic( ccnI(c|c cc (ecu|cccc: ccnI(c|cncc c 1 mc|cu|c (ecu|ccc(c
ccnI(c|c-:e c p(cce::c
Umc enzimc cpenc: ccm efeiIc c|c:I(icc p(ccuz um efeiIc (ecu|ccc( muiIc
menc(.

18

/DF
AIF
Glicose
Glicose 6-P
Frutose 6-P
Frutose 1,6- bisP
Gliceraldeido 3-P Dihidroxicetona P
1,3-Bisfosfoglicerato
3-Fosfoglicerato
2-Fosfoglicerato
Fosfoenolpiruva
to
Piruvat
o
Lactato
AIF
/DF
NAD
+
N/DH + H
+

Fi
AIF
/DF
/DF
AIF
N/DH + H
+

NAD
+
Hexoc|nose
Fos|og||comutose
A|do|ose
Fos|o|rutoc|nose
Ir|ose Fos|oto
Isomerose
G||cero|de|do 3-F
Des|drogenose
Fos|og||cerotoc|nose
Fos|og||cerotomutose
Eno|ose
F|ruvoto c|nose
Loctoto
Des|drogenose

19

3
33.
.. D
DDo
oo F
FF|
||r
rru
uuv
vvo
oot
tto
oo o
ooo
oo C
CC|
||c
cc|
||o
oo d
dde
ee K
KKr
rre
eeb
bbs
ss

3.1. Descorbox||oo ox|dot|vo do p|ruvoto

/ ce:cc(Lcxi|cccc cxiccnIe cc pi(uvcIc c /ceIi|-Cc/ c e|c ce |iccccc enI(e c
c|icc|i:e e c cic|c ce K(eL:
C pi(uvcIc I(c:npc(Iccc pc(c
cenI(c cc miIcccnc(ic cI(cv: ce um
I(cn:pc(Iccc e:pecic| ce ccicc
pi(uvicc cue cjucc c :uc pc::ccem
cI(cv: cc memL(cnc miIcccnc(ic|
inIe(nc. c cue envc|ve um
meccni:mc ce :impc(Ie nc cuc| um
p(cIcc cc-I(c:npc(Iccc.
/ ce:ic(ccencccc e
ce:cc(Lcxi|cccc ccmLinccc: cc
pi(uvcIc c cceIi|-Cc/ (ecue(em c
ccccc :ecuencic| ce 3 enzimc:
cife(enIe: {inIec(ccc: num ccmp|exc
mu|IienzimcIicc - ccmp|exc pi(uvcIc
ce:ic(ccenc:e) e 5 ccenzimc: cu
c(upc: p(c:IIicc: cife(enIe:.

Comp|exo F|ruvoto Des|drogenose
Enzimc:
Des|drogenose p|rv|co {c(upc p(c:IIicc IFF)
D|h|dro||po||tronscet||ose {c(upc p(c:IIicc Lipccmicc)
D|h|dro||po||des|drogenose {c(upc p(c:IIicc F/D)

Ccenzimc:
I|om|no p|ro|os|oto {IFF) - (ecce ccm c pi(uvcIc
L|poom|do - cceiIc c(upc cceIi| e I(cn:fe(e-c pc(c c Cc/
CoA - cceiIc c(upc cceIi|
FAD - cceiIc ecuivc|enIe: (ecuIc(e:
NAD
+
- cceiIc ecuivc|enIe: (ecuIc(e:

ViIcminc: cife(enIe: (ecue(icc: ne:Ie :i:Iemc ccmc ccmpcnenIe: viIci::
Iicminc IFF
FiLcf|cvinc F/D
NiccIinc N/D
FcnIcIencIc Cc/

Etopos
1. C C1 cc pi(uvcIc |iLe(Iccc ccmc CO2 pe|c ccccc cc ce:ic(ccenc:e pi(uvicc
e c C2 unicc cc IFF ccmc grupo h|drox|et||
2. E:Ie c(upc cxicccc c cceIcIc. cue I(cn:fe(icc ccm mci: 2 e
-
pc(c c fc(mc
cxicccc cc grupo ||poom|do cc cihic(c|ipci|I(cn:ceIi|c:e.
3. Dc-:e umc I(cn:e:Ie(ificcccc nc cuc| c c(upc -SH cc CoA :uL:IiIui c c(upc -SH
cc cihic(c|ipci|I(cn:ceIi|c:e pc(c c(icinc( Acet|| CoA e c fc(mc ccmp|eIcmenIe
(ecuzicc cc |ipccmicc.
4. Fc(c c (ecene(cccc cc ||poom|do ox|dodo. hc I(cn:fe(ncic: ce e|ecI(ce:
pe|c ccccc cc enzimc cihic(c|ipci|ce:ic(ccenc:e. p(epc(cncc c ccmp|exc
pc(c cuI(c ce:ic(ccencccc. C: e|ecI(ce: (emcvicc: pc::cm pc(c c NAD
+

cI(cv: cc FAD.
20

keoco g|obo|
F|ruvoto + CoA + NAD
+
Acet||-CoA + CO2 + NADH + H
+
AG0'= -8 kco|/mo|

/ pc::ccem cc pi(uvcIc c /ceIi|-Cc/ p(cmcvicc pe|c ccmp|exc cc
ce:ic(ccenc:e pi(uvicc nc :cmcIc(ic cc: (ecccce: umc ce:cc(Lc|ixcccc
cxiccIivc
Lc:IcnIe ence(ccnicc. |cncc i((eve(:|ve|
umc cc: p(incipci: (czce: pe|c fccIc cc: ccicc: cc(cc: ncc :e ccnve(Ie(em c
hic(cIc: ce cc(Lcnc.

kegu|oo
/ (ecu|cccc c|c:I(icc e pc( mccificcccc ccvc|enIe
/ pi(uvcIc ce:ic(ccenc:e iniLicc pe|c: :eu: p(ccuIc: {(eI(c-iniLiccc): /ceIi|-Cc/
e N/DH.
/ pi(uvcIc ce:ic(ccenc:e iniLicc:
Fc( um c|Ic pcIencic| ene(cIicc
Nc: ccncicce: ce cxiccccc ce ccicc: cc(cc:
cue ccncuzem c um cumenIc cc: (czce:
[Acet||-CoA]/[CoA]: [AIF]/[ADF]: [NADH]/[NAD
+
]
/ mccificcccc ccvc|enIe cI(cv: cc fc:fc(i|cccc
{cinc:e) e ce:fc:fc(i|cccc {fc:fcIc:e) cue :cc
(ecu|ccc: pe|c: c|Ie(ccce: cccue|c: (czce:.
Cucncc cumenIcm
ccIivccc c cinc:e.
inccIivcncc c ccmp|exc
Cucncc ciminuem c
ccmp|exc ce:fc:fc(i|ccc
pe|c fc:fcIc:e. ccIivcncc-c.
C cumenIc ce in:u|inc {Iecicc
ccipc:c) e ce epinef(inc {Iecicc
mu:cu|c( cc(c|ccc) ccIivc c pi(uvcIc
ce:ic(ccenc:e.
C /MFc ncc Iem cuc|cue( (e|cccc
ccm e:Ic (ecu|cccc.

Hic(cxieIi|-IFF
IFF
CC2
Fi(uvcIc
Fi(uvcIc
ce:ic(ccenc:e
Lipccmicc
cxicccc
/ceIi|
Lipccmicc
Lipccmicc
(ecuzicc
Dihic(c|ipci|
I(cn:ceIi|c:e
Dihic(c|ipci|
De:ic(ccenc:e
/ceIi|-Cc/
Cc/
F/D
N/D
+

N/DH +H
+

F/DH2
21

3.2. C|c|o de Krebs {Cic|c cc: cicc: I(icc(Lcxi|icc: cu Cic|c cc cicc C|I(icc)

Loco|: miIcccnc(ic {emLc(c exi:Icm i:cenzimc: nc ciIc:c|)

C cic|c ce K(eL: c vic finc| ccmum pc(c c cxiccccc ce mc|cu|c: c|imenIc(e:. /
mcic(ic ce:Ic: mc|cu|c: enI(c nc cic|c ccmc /ceIi|-Cc/
E:Ie cic|c fc(nece inIe(mecic(ic: pc(c Lic::|nIe:e. cc| cue :ejc cnfiLc|icc.
Fc(c ccmecc( umc vc|Ic nc cic|c c /ceIi|-Cc/ {2C) ccmLinc-:e ccm
cxc|ccceIcIc {4C) (e:u|Icncc nc fc(mcccc ce ciI(cIc {C).
Secue-:e um ccnjunIc ce (ecccce: cI(cv: cc: cuci: 2 mc|cu|c: ce cicxicc ce
cc(Lcnc :e pe(cem. e (ecene(ccc c ccicc cxc|cccIicc.
Vi:Ic cue ncc :cc p(eci:c: mci: cc cue pecuenc: cucnIiccce: ce ccicc
cxc|cccIicc pc(c I(cn:fc(mc( c(cnce nme(c ce uniccce: cceI||icc: c CC2.
ccn:ice(c-:e cue c cxc|ccceIcIc ce:empenhc c pcpe| ccIc||Iicc.
4 cc: 8 (ecccce: {pc::c:) :cc cxicccce:. :encc c ene(cic cc cxiccccc
ccn:e(vccc :cL c fc(mc ce ccenzimc: (ecuzicc:: F/DH2 e N/DH
E:Ie p(cce::c nece::iIc ce C2 ccmc (ecepIc( finc| cc: ecuivc|enIe: (ecuIc(e:.

1. Formoo de C|troto
/ 1 (eccccc cc cic|c ccn:i:Ie nc ccncen:cccc cc Acet||-CoA ccm c c|do
Oxo|ooct|co pc(c fc(mc( c cicc C|I(icc. ccIc|i:ccc pe|c CiI(cIc :inIc:e. cue
efecIuc umc |iccccc cc(Lcnc-cc(Lcnc enI(e c cc(Lcnc-meI||icc cc /ceIi|-Cc/ e
c cc(Lcnc cc(Lcni| cc ccicc cxc|cccIicc.
/ (eccccc ce ccncen:cccc c CiI(ci|-SCc/. cccmpcnhccc pe|c hic(c|i:e cc
|iccccc Iic:Ie( cc Cc/. pe(cencc-:e c(cnce pc(Ie cc ene(cic |iv(e ccmc cc|c(. c
cue cc(cnIe cue c (eccccc ccc((c cI cc fim.
/pe:c( cc Lcixc ccncenI(cccc ce cxc|ccceIcIc c (eccccc ccc((e cevicc cc
Lcixc vc|c( ce AG0.
C ciI(cIc pcce Ie( cuI(c: ce:Iinc: meIcLc|icc: ccmc c: Lic::|nIe:e: (ecuIc(c: ce
|ipicc:.

2. Formoo de |soc|troto
C c|troto i:cme(izccc c |soc|troto pe|c ccccc cc enzimc /ccniIc:e cue ccnIm
fe((c nc e:Iccc Fe
2+
nc fc(mc ce umc p(cIe|nc fe((c-enxcf(e {Fe:S).
E:Ic ccnve(:cc ccc((e em 2 eIcpc:: ce:ic(cIcccc c cicc Ci:-cccn|Iicc. e c
(ehic(cIcccc c l:cciI(cIc.
/ (eccccc iniLicc pe|c cicc f|c(-ccIicc. c cuc|. nc fc(mc ce f|uc(cceIi|-Cc/
ccncen:c-:e ccm c ccicc cxc|cccIicc pc(c fc(mc( c ccicc f|uc(c|I(icc. E:Ie
|Iimc iniLe c cccniIc:e. ccc:icncncc c ccumu|cccc ce ccicc c|I(icc.
pc::|ve| cue c ci:-cccniIcIc ncc :ejc um inIe(mecic(ic cL(iccIc(ic enI(e c ciI(cIc
e c i:cciI(cIc. :encc nc (ec|iccce umc (cmificcccc |cIe(c| cc vic p(incipc| cc cic|c.
/ i:cme(izcccc p(ccuz um ccmpc:Ic ce cc(Lcnc (cmificccc mci: fcci| ce
meIcLc|izc(.

3. Formoo de u-cetog|utoroto
C ccicc i:cc|I(icc :cf(e ce:ic(ccencccc em p(e:encc cc l:cciI(cIc-ce:ic(ccenc:e
pc(c fc(mc( ccicc Cxc|c:ucc|nicc.
/ cxiccccc. |icccc c ccceic (e:pi(cIc(ic cc i:cciI(cIc ccc((e cuc:e exc|u:ivcmenIe
cI(cv: cc enzimc N/D
+
-cepencenIe.
Acet||-CoA + Oxo|oocetoto + H2O C|troto + CoA$H
CiI(cIc SinIc:e
C|troto C|s-ocon|toto + H2O Isoc|troto
/ccniIc:e
22

Secue-:e umc ce:cc(Lcxi|cccc c u-ceIcc|uIc(cIc. IcmLm ccIc|i:ccc pe|c
l:cciI(cIc-ce:ic(ccenc:e.
pc::|ve| cue c ccicc Cxc|c:ucc|nicc pe(mcnecc |icccc c enzimc ccmc
|nIe(mecic(ic nc (eccccc c|cLc|.

4. Formoo de $ucc|n|| CoA
Ic| ccmc c p|ruvoto. c u-cetog|utoroto um u -cetoc|do {i:Ic . pc::ui um c(upc
cc(Lcni|c ccjccenIe cc c(upc ccicc cc(Lcx||icc).
Lccc. (ecce exccIcmenIe ccmc c pi(uvcIc. i:Ic . c :uc ce:cc(Lcxi|cccc fc(nece
ene(cic :uficienIe pc(c cue :e fc(me umc |iccccc Iic:Ie( ccm c ccenzimc /.
/ enzimc (e:pcn:cve| pc( e:Ic (eccccc. c u-ceIcc|uIc(cIc ce:ic(ccenc:e.
:eme|hcnIe c pi(uvcIc ce:ic(ccenc:e nc :uc ccmpc:iccc e ccfccIc(e::
C ecui||L(ic ce:Ic (eccccc ce Ic| mccc c fcvc( cc fc(mcccc ce Succini|-Cc/
cue. fi:ic|cc|ccmenIe. e|c ceve :e( ccn:ice(ccc unici(eccicnc|.
C /(:eniIc iniLe c (eccccc. ccu:cncc c ccumu|cccc ce u-ceIcc|uIc(cIc.

5. Formoo de $ucc|noto
C $ucc|n||-CoA ccnve(Iicc c $ucc|noto pe|c Succini|-Cc/ SinIeIc:e.
E:Ic (eccccc (ecue( GDF. cue ccnve(Iicc em GIF. em p(e:encc ce fc:fcIc
inc(ccnicc.

E:Ie c nicc exemp|c. nc cic|c cc ccicc c|I(icc. cc fc(mcccc ce um fc:fcIc ce
c|Ic ene(cic cc n|ve| cc :uL:I(cIc. e :u(ce em vi(Iuce cc |iLe(Icccc cxiccIivc cc u-
ceIcc|uIc(cIc.
C AIF pcce :e( fc(mccc c pc(Ii( cc GIF pc( meic ce umc nuc|ec:icc-cifc:fcIc-
cinc:e:

ADF + GIF AIF + GDF

. kegeneroo do Oxo|oocetoto

C :uccincIc :cf(e meIcLc|izcccc ccicicnc| pc::cncc. p(imei(c pc( umc
ce:ic(ccencccc. :ecuicc pe|c cciccc ce H2C. e :uL:ecuenIemenIe. pc( umc
ncvc ce:ic(ccencccc. cue (ecene(c c cxc|ccceIcIc.

/ SuccincIc-ce:ic(ccenc:e e:Ic |icccc c :upe(f|cie inIe(nc cc memL(cnc
miIcccnc(ic| inIe(nc. cc ccnI(c(ic cc: cuI(c:. :encc c nicc ce:ic(ccencccc cc
cic|c ce K(eL: cue envc|ve c I(cn:fe(ncic ci(ecIc ce hic(ccnic cc :uL:I(cIc :em
c pc(Iicipcccc cc N/D
+
. umc vez cue c cceiIccc( c F/D.
C: e|ecI(ce: :cc I(cn:fe(icc: ci(ecIcmenIe cc: cc|cme(ccc: Fe-S cc enzimc e
ce:Ie: pc(c c ULicuincnc {ccceic ce I(cn:pc(Ie ce e|ecI(ce:).
/ cciccc ce Mc|cIc cu Cxc|ccceIcIc iniLe ccmpeIiIivcmenIe c SuccincIc-
ce:ic(ccenc:e p(cvcccncc c ccumu|cccc ce SuccincIc.
Isoc|troto + NAD
+
Oxo|osucc|noto + NADH + H
+

Oxo|osucc|noto u-cetog|utoroto + CO2
l:cciI(cIc ce:ic(ccenc:e
u-cetog|utoroto + CoA + NAD
+
$ucc|n||-CoA + NADH + CO2
u-ceIcc|uIc(cIc ce:ic(ccenc:e
$ucc|n||-CoA + GDF + F| $ucc|noto + CoA$H + GIF
Succini|-Cc/ :inIeIc:e
1. $ucc|noto + FAD Fumoroto + FADH2
SuccincIc ce:ic(ccenc:e
23

EmLc(c c ecui||L(ic ce:Ic (eccccc :ejc muiIc fcvc(cve| cc mc|cIc. c f|uxc efecIivc
nc ci(ecccc cc cxc|ccceIcIc pc(cue e::e ccmpc:Ic. junIcmenIe ccm cuI(c p(ccuIc
cc (eccccc (emcvicc nc: (ecccce: :uL:ecuenIe:.

keoco g|obo|:
Acet||-CoA + 3NAD
+
+FAD + 2H2O + GDF+F| 2CO2 + 3NADH + 3H
+
+ FADH2 + GIF+CoA

8o|ono Energt|co
2. Fumoroto + H2O Mo|oto
Fumc(c:e
3. Mo|oto + NAD
+
Oxo|oocetoto + NADH + H
+

Mc|cIc ce:ic(ccenc:e
24

C: ecuivc|enIe: (ecuIc(e: fc(mccc: ccmc (e:u|Iccc cc ccccc cc: ce:ic(ccenc:e:
:cc I(cn:fe(icc: pc(c c ccceic (e:pi(cIc(ic nc memL(cnc inIe(nc cc miIcccnc(ic.
Du(cnIe c pc::ccem. pe|c ccceic I(cn:pc(Iccc(c ce e|ecI(ce:. c: ecuivc|enIe: ce
(ecuccc cc N/DH + H
+
c(icinc 3 /IF:. CcnIucc. c F/DH2 p(ccuz :cmenIe 2 /IF:.
vi:Ic cue I(cn:fe(e c :eu ecuivc|enIe (ecuIc( ci(ecIcmenIe c ULicuincnc. c cue
:icnificc cue enI(c nc ccceic (e:pi(cIc(ic cepci: cc N/DH.
Um cuI(c fc:fcIc ce c|Ic ene(cic fc(mccc nc p(cp(ic cic|c
/::im 12 /IF: :cc fc(mccc: pc( cccc cic|c ce ccicc c|I(icc. e ccmc cccc
mc|cu|c ce c|icc:e c(icinc 2 cic|c. ccn:ice(cm-:e 24 /IF:.

kegu|oo
C cic|c ce K(eL:
E:Ic :ujeiIc c: nece::iccce: cc c|u|c em /IF
ccnI(c|ccc funccmenIc|menIe pe|c ci:pcniLi|iccce ce :uL:I(cIc:. iniLiccc pe|c:
p(ccuIc: e pc( cuI(c: inIe(mecic(ic: cc cic|c.
cepencenIe cc fc(necimenIc cc: cc-fccIc(e: cxicccc: cc: ce:ic(ccenc:e:. c:
cuci:. pc( :uc vez. cevicc c fc(Ie |iccccc enI(e c cxiccccc e c fc:fc(i|cccc nc
CIE. :cc cepencenIe: cc ci:pcniLi|izcccc ce /DF e. pc(IcnIc. cc ve|cciccce ce
uIi|izcccc ce /IF.
IcmLm (ecu|ccc cc n|ve| ce c|cumc: enzimc:. cue :cc (e:pcn:cvei: pe|c
e:Iccc ce ene(cic exp(e::c pe|c: (e|ccce: /IF//DF e N/DH/N/D
+

kegu|oo no entrodo de Acet|| -CoA (2 enz|mos)
Ccmp|exc cc Fi(uvcIc-De:ic(ccenc:e:
CiI(cIc SinIc:e
ln|b|dores:
/IF
N/DH
Succini|-Cc/
De(ivccc: ce /ci|-Cc/ ce
ccceic: Lcncc:
CiI(cIc

Acf|vodores:
/DF e /MF

/ ccncenI(cccc ce cxc|ccceIcIc Icmnm ce:empenhc um pcpe| impc(IcnIe nc
(ecu|cccc ce:Ic enzimc.

kegu|oo do v|o propr|omente d|to
l:cciI(cIc- De:ic(ccenc:e
CcIc|izc c 3 eIcpc cc cic|c
Scf(e (ecu|cccc c|c:I(icc:
ln|b|dores:
/IF
N/DH

Acf|vodores:
/DF e /MF
lcc Cc
2+

u-CeIcc|uIc(cIc-De:ic(ccenc:e
CcIc|izc c eIcpc 4 cc cic|c:
IcmLm c|c:I(icc
ln|b|dores:
/IF
N/DH
GIF
Succini|-Cc/
Acf|vodores:
/DF e /MF
lcc Cc
2+

SuccincIc De:ic(ccenc:e
Fccu|ccce ce Mecicinc ce Li:Lcc Mcc|c ll.l - 8iccu|micc
25

ln|b|dore : cxc|ccceIcIc

keoces onop|erot|cos
Scc (ecccce: cue ccmp|eIcm c: ccncenI(ccce: ce inIe(mecic(ic: cc Cic|c ce K(eL:.
cucncc c ccncenI(cccc ce um ce|e: ciminui nc c|u|c. cc(cnIincc c::im c
ccnIinuiccce cc vic.

C|c|o de Krebs - c|c|o on||bo||co

c fc:e Ie(minc| cc cxiccccc ce(cLic: cc: p(incipci: mc|cu|c: nuI(ienIe: e
ccn:IiIui c pcnIc ce pc(Iicc pc(c c Lic::|nIe:e.
E:Ic: vic: ccmp(eencem p(cce::c: ccmc c Necc|icccne:e. I(cn:cmincccc.
De:cmincccc e S|nIe:e ce ccicc: cc(cc:.
Nc :uc ve(IenIe ccIcLc|icc. c cic|c ce K(eL: pc::iLi|iIc c fc(mcccc ce /IF.
Iccc: c: meIcLc|iIc: inIe(mecic(ic: cc cic|c ce:ce c ccicc c|I(icc cI cc ccicc
cxc|cccIicc :cc pcIencic|menIe c|icccnicc:. vi:Ic cue pccem c(icinc( umc
p(ccuccc efecIivc ce c|icc:e nc f|cccc cu nc (im. umc vez cue e:Ie: c(ccc: :cc
c: nicc: cue pc::uem c: enzimc: nece::c(ic: c necc|icccne:e.
/ enzimc chcve cue pe(miIe c I(cn:fe(ncic efecIivc pc(c fc(c cc cic|c. ce um
cc: :eu: ccmpcnenIe:. pc(c c vic cc necc|icccne:e. c fc:fcenc|pi(uvcIc-
cc(Lcxicinc:e. cue ccIc|i:c c ce:cc(Lcxi|cccc cc ccicc cxc|cccIicc c ccicc
fc:fcenc|pi(vicc. funcicncncc c GIF ccmc fcnIe ce ene(cic.
/: (ecccce: ce I(cn:cmincccc p(ccuzem pi(uvcIc c pc(Ii( cc c|cninc.
cxc|ccceIcIc c pc(Ii( cc c:pc(IcIc e ccicc c-ceIcc|uIc(icc c pc(Ii( cc ccicc
c|uIclnlcc.
Fe|c fccIc ce e:Ic: (ecccce: :e(em (eve(:|vei:. c cic|c fc(nece IcmLm c:
e:cue|eIc: cc(Lcnccc: pc(c c :|nIe:e ce c|cun: cc: cmincccicc: ncc e::encici:.
C cceIi|-Cc/. fc(mccc c pc(Ii( cc pi(uvcIc. c p(incipc| ccmpc:Ic cue inicic c
:|nIe:e cc: ccicc: cc(cc:. Nc enIcnIc. c cceIi|-Cc/ ncc ccn:ecue cI(cve::c( c
memL(cnc miIcccnc(ic| inIe(nc. e pc(IcnIc Iem cue :e( I(cn:fc(mccc em ciI(cIc
pc(c :e( I(cn:pc(Iccc pc(c fc(c cc miIcccnc(ic. cnce :e (efcz c cceIi|-Cc/. umc
vez cue c: enzimc: (e:pcn:cvei: pe|c :|nIe:e cc: ccicc: cc(cc: :cc
exI(cmiIcccnc(ici: e c pi(uvcIc-ce:ic(ccenc:e exc|u:ivcmenIe miIcccnc(ic|.
Em ccncicce: excecenIc(ic: ce cc(cc ene(cIicc. c|cun: cc: inIe(mecic(ic: cc
cic|c :cc cp(cveiIccc: nc :|nIe:e cc: pc(fi(inc:. c|cun: cmincccicc:. pi(imicinc: cu
c|icc:e {e cuI(c: pc|i::ccc(icc:).
C cic|c ce K(eL: ceixc ce funcicnc( cucncc c: :eu: inIe(mecic(ic: meIcLc|icc: :cc
ce:viccc: pc(c Lic::inIe:e:. c menc: cue c cxc|ccceIcIc e/cu mc|cIc. :ejcm
fc(mccc: c pc(Ii( cc pi(uvcIc. pc::iLi|iIcncc c cxiccccc ce ncvc: mc|cu|c: ce
/ceIi|-Cc/.

F|ruvoto + CO2 + AIF + H2O Oxo|oocetoto + ADF + F| + H
+

Fi(uvcIc cc(Lcxi|c:e
26

Fi(uvcIc
Cxc|ccceIcIc
CiI(cIc
u-ceIcc|uIc(cIc
Succini|-Cc/
Fumc(cIc
Mc|cIc
/ceIi|-Cc/
/ceIccceIcIc /|cninc
/mincccicc:
cicc:
cc(cc:
E:Ie(cice:
G|uIcmcIc
/mincccicc:
F(cpicni|-Cc/ /mincccicc:
cicc: cc(cc:
Fc:fcenc| pi(uvcIc
G|icc:e
/:pc(IcIc
Cic|c ce K(eL:
Lipicc:
Fc(fi(inc:
Fi(imicinc:
27

G|icc:e -F G|icc:e 1-F
G|icccnic
G|icc:e
CcIcLc|i:mc cu ccnI(c|c
cc: n|vei: ce c|icmic
5 5. . G G| || |c co og ge en no o| || |s se e e e I In nt te er rc co on nv ve er rs s o o d de e
M Mo on no os ss so oc c r r| |d do os s

5.1 - G||cogeno||se

G||cogeno||se: p(cce::c ccIcLc|icc ce fc(mcccc ce mc|cu|c: ce c|icc:e--fc:fcIc c
pc(Ii( cc pc|i::ccc(icc ce c|icc:e. c|icccnic.
Nc: c|u|c: em cue exi:Ie c enzimc c|icc:e -fc:fcIc:e {ex.: hepcIcciIc. c|u|c
epiIe|ic| inIe:Iinc|) c p(ccuIc finc| cc c|icccenc|i:e c c|icc:e.
Nc: c|u|c: mu:cu|c(e: cnce c c|icc:e -fc:fcIc:e ncc exi:Ie c c|icc:e -fc:fcIc
:uL:I(cIc ce(ccc( cc: p(ccuIc: /IF. CC2 e H2C {m:cu|c ve(me|hc) cu ce |ccIcIc
{m:cu|c L(cncc).

G|icccnic: pc||me(c ce (e:|cuc: ce c|icc:e unicc: enI(e :i pc( |icccce: c|icc:|cicc:
1.4 e ccm |icccce: c|icc:icicc: 1.. nc: (cmificccce:.
Fr|nc|po| |uno: (e:e(vc ce c|icc:e (cpiccmenIe ci:pcn|ve| pc(c c: nece::iccce:
meIcLc|icc:. em cc:c ce cc(ncic ene(cIicc.

Etopos do g||cogeno||se

G|icccnic G|icc:e 1-fc:fcIc G|icc:e -fc:fcIc

I - Decompos|o do g||cogn|o em resduos de g||cose 1-|os|oto

1. /: |icccce: c|icc:icicc: 1.4 cc:
exI(emiccce: ncc (ecuIc(c: cc
c|icccnic :cc :uce::ivcmenIe c|ivccc:
pc( fc:fc(c|i:e. |iLe(Icncc c: (e:|cuc: ce
c|icc:e :cL fc(mc ce c|icc:e 1-fc:fcIc.
2. / ccccc cc fc:fc(i|c:e cc c|icccnic
cecc((e c G Lcixc. pci: cue cccc
|iccccc c|icc:|cicc :uL:IiIu|cc pc(
umc |iccccc :Ie(-fc:fcIc.
3. / ccccc cc fc:fc(i|c:e inIe((cmpe-:e c
ce(cc ce 4 (e:|cuc: ce cccc |iccccc
c|icc:|cicc 1..
4. Secue-:e c ccccc ce cuI(c enzimc {
1.4 1.4 c|iccn I(cn:fe(c:e) cue
I(cn:fe(e um I(i::ccc(icc ce um
:ecmenIc pc(c cuI(c. cpc: c|ivccem cc
|iccccc 1.4.
5. /cc(c c (e:|cuc ce c|icc:e cue
c::ecu(c c |iccccc 1. ficc expc:Ic c
ccccc hic(c||Iicc cc 1. c|icc:icc:e.
:encc |iLe(Iccc ccmc c|icc:e |iv(e.
28

4/IF
Fc:fc(i|c:e cinc:e
Fc:fc(i|c:e fc:fcIc:e
G|icccnic
Fc:fc(i|c:e L
G|icccnic
Fc:fc(i|c:e c
4/DF
G|icccnic
G|icccnic 1-fc:fcIc
Fi
4Fi 4H2C
. Fe:Ic ccc(c umc :ecuncic exIen:c cpenc: ccm |icccce: c|icc:|cicc: 1.4
ncvcmenIe expc:Ic: c ccccc cc fc:fc(i|c:e.
7. ExcepIucncc c: (e:|cuc: cc c|icc:e |iv(e cue fc(mcm c: |icccce: 1.. Iccc: c:
(e:IcnIe: :cc (emcvicc: cc c|icccnic :cL c fc(mc ce c|icc:e 1-fc:fcIc

Fos|oro||se: c|ivccem ce umc |iccccc pc( inI(ccuccc cc: ccmpcnenIe: cc ccicc:
fc:fc(icc.
Vontogens do |os|or||oo de resduos de g||cose:
Ncc hc ci:pncic cc /IF pc(c fc:fc(i|c( c c|icc:e. c cue Ic(nc imecicIcmenIe
cce::|ve| c: eIcpc: meIcLc|icc:.
/ c|icc:e 1-fc:fcIc ncc pcce cifunci( cc c|u|c {pe|c (czcc jc mencicnccc c
p(cpc:iIc cc: meIcLc|iIc: cc c|icc|i:e).

G|icccnic fc:fc(i|c:e {cinc:e):
Enzimc cue c|ivc umc |iccccc 1.4 ccm fc:fcIc inc(ccnicc {Fi)
Nc cenI(c ccIivc cp(e:enIc c |os|oto de p|r|doxo| {fc(mc ccIivc cc viIcminc 8)
ccmc c(upc p(c:IIicc.
ccn:IiIu|cc pc( 4 :uLuniccce:
/p(e:enIc ccmpc(IcmenIc cinIicc c|c:I(icc ccm e:Iccc: ccnfc(mccicnci:
ci:IinIc: cmLc: (ecu|ccc: pc( mccu|cccc ccvc|enIe {fc:fc(i|cccc/ce:fc:fc(i|cccc
cc: (e:|cuc: ce :e(inc).
2 Fc(mc: inIe(ccnve(Iivei::
Fos|or||ose o {fc:fc(i|ccc. ccIivc)
Fos|or||ose b {ce:fc:fc(i|ccc. inccIivc)
/ pc::ccem cc fc:fc(i|c:e Lc fcz-:e nc p(e:encc cc c|icccnic fc:fc(i|c:e cinc:e
e /IF. encucnIc cue c (eccccc inve(:c ccc((e nc p(e:encc ce c|icccnic
fc:fc(i|c:e fc:fcIc:e.

/ c|icccnic fc:fc(i|c:e Lc:IcnIe mci: (cpicc cc cue c: cuI(c: enzimc:
inIe(venienIe:. pe|c cue c: (cmc: exIe(ic(e: cc c|icccnic :cc cec(ccccc: muiIc
(cpiccmenIe nc m:cu|c em pcucc: :ecuncc: cucncc nece::c(ic muiIc ene(cic. /
cec(cccccc cc c|icccnic pc(c |c ce:Ie pcnIc exice c I(cn:fe(c:e e c 1.
c|icc:icc:e e pc(IcnIc mci: |enIc. c cue exp|icc em pc(Ie c fccIc cc m:cu|c :c
pcce( exe(ce( c :uc mcximc fc(cc cu(cnIe pcucc: :ecuncc:.

/ceni|cic|c:e l
/ceni|cic|c:e /
/MF
/MFc
F(cIe|nc cinc:e l
F(cIe|nc cinc:e /
G|icccnic
Fc:fc(i|c:e L
G|icccnic
Fc:fc(i|c:e c
Cc|cic
/c(enc|inc
G|ucccinc
29

II - Irons|ormoo do g||cose 1-|os|oto em g||cose -|os|oto - eIcpc ccmum c
c|icccne:e.

/ c|icc:e 1-fc:fcIc. cLIicc ccmc p(ccuIc finc| cc fc:fc(i|cccc cc c|icccnic.
I(cn:fc(mccc em c|icc:e -fc:fcIc pc( ccccc cc fc:fcc|iccmuIc:e.
Nc inIe(ccnve(:cc ce cmLc: c: meIcLc|iIc: fc(mc-:e um inIe(mecic(ic {c|icc:e
1.-Li:fc:fcIc).

Fc:fcc|uccmuIc:e:
Nc cenI(c ccIivc cp(e:enIc um (e:|cuc ce :e(inc cujc c(upc CH :e enccnI(c
fc:fc(i|ccc pe|c ccicc fc:fc(icc.
E:Ie c(upc fc(fc(i|c I(cn:fe(icc pc(c c c(upc CH ce C cc c|icc:e 1-fc:fcIc.ccncc
c(icem c c|icc:e 1.-Li:fc:fcIc.
Nc :ecuncic e:Ie inIe(mecic(ic I(cn:fe(e c c(upc fc:fc(i|c ce C1 pc(c c (e:|cuc ce
:e(inc cc enzimc. (ecene(cncc c fc:fcenzimc e :imu|IcnecmenIe. c(icincncc c
c|icc:e -fc:fcIc.
Ccm c Iempc. c enzimc pe(ce c :eu c(upc fc:fc(i|c. pc( hic(c|i:e: c (efc:fc(i|cccc
cc enzimc c::ecu(ccc pc( I(cn:fe(ncic c pc(Ii( cc c|icc:e 1.-Li:fc:fcIc. pc( :uc
vez ce(ivccc cc (eccccc ccIc|i:ccc pe|c fc:fcc|icccinc:e. c pc(Ii( cc c|icc:e 1-
fc:fcIc e /IF.

Dest|no metobo||co do g||cose -|os|oto
/ c|icc:e -fc:fcIc ce(ccc c pc(Ii( cc cec(cccccc cc c|icccnic Iem 2 ce:Iinc:
pc::|vei::
ccn:umicc pe|c c|u|c
Difunce-:e pc(c c ci(cu|cccc e e:pccc: exI(c-ce|u|c(e:.

Nc f|cccc {(im e inIe:Iinc:). c c|icc:e -fc:fcIc hic(c|i:ccc pe|c c|icc:e -
fc:fcIc:e. c cue pe(miIe c c|icc:e cI(cve::c( c: memL(cnc: ciIcp|c:mcIicc:.
De:Ic fc(mc. c ccn:ecuncic finc| cc c|icccenc|i:e hepcIicc ccn:i:Ie nc
cumenIc cc c|icemic. :cL(eIucc Ii| nc mcnuIenccc cc: n|vei: ci(cu|cnIe: ce c|icc:e
nc: inIe(vc|c: enI(e c: (efeicce: cu nc cecu(:c cc exe(c|cic mu:cu|c( inIen:c.
Nc enIcnIc. ccue|c enzimc ncc exi:Ie nc m:cu|c e nc c(eL(c. Dccui (e:u|Ic c
(eIenccc e ccn:ecuenIe cp(cveiIcmenIc meIcLc|icc cc c|icc:e -fc:fcIc pc( ccue|e:
Iecicc:. cpc: c|icccenc|i:e |ccc|.
/ cife(encc nc cp(cveiIcmenIc meIcLc|icc cc c|icc:e -fc:fcIc exp|icccc
pe|c uIi|iccce cc c|icc:e ccmc nuI(ienIe cccue|e: Iecicc:: funccmenIc| pc(c c:
ccIiviccce: meIcLc|icc: ce(eL(ci: e cc m:cu|c {:cL ccnI(ccccc inIen:c) e
:ecuncc(ic pc(c c f|cccc {em cue :up|cnIccc pe|c: ccicc: cc(cc:). /::im. c f|cccc
c(mczenc c c|icc:e pc(c fc(nece( c cuI(c: Iecicc: cue ce|c cepencem mci:
e:I(eiIcmenIe. ccmc c m:cu|c e c c(eL(c.

G||cogenoses {cu ccencc: ce cepc:iIc: ce c|icccnic"): I(ccuz ceficincic:
he(eciIc(ic: {(ece::ivc:) ce c|cumc: enzimc: cc c|icccenc|i:e e c: :uc: vc(icnIe:.
IcmLm :e enc|cLcm c: ceficincic: cc c|icc:e -fc:fcIc:e e cc 1.4 c|icc:icc:e
p(e:enIe nc: |i:c::cmc:. E:Ic: ccencc: pccem :e( |eIci: nc infcncic. encucnIc cuI(c:
ccc((em ccm :inIcmcIc|ccic mcce(ccc.

G||cose 1-Fos|oto G||cose 1, b|s|os|oto G||cose -Fos|oto
Fc:fcc|uccmuIc:e
30

5.2 - V|os de |nterconverso dos monossocr|dos

G(cnce pc(Ie cc: mcnc::ccc(icc: p(e:enIe: nc meic ce|u|c( ce(ivcm cc c|icc:e. /:
inIe(ccnve(:ce: imp||ciIc: Im cLjecIivc: ene(cIicc: e/cu vi:cm c cLIenccc ce
ccmpc:Ic: ccm nme(c vc(icve| ce cc(Lcnc: pc(c Lic::|nIe:e: e:pec|ficc:.

I - Interconverso dos pr|nc|po|s hexoses

/|m cc mc|Ic:e. cuI(c: ci::ccc(icc: ccmun: nc c|imenIcccc :cc c |ccIc:e e c
:ccc(c:e: pc( hic(c|i:e enzimcIicc {mc|Ic:e. :ccc(c:e. |ccIc:e). e:Ie: ci::ccc(icc: ccc
c(icem c 3 Iipc: ce mcnc::ccc(icc:: c c|icc:e. c f(uIc:e e c cc|ccIc:e.

Cofobo||smo do frufose
Frutose:
Ccn:IiIui umc f(ccccc impc(IcnIe cc: c||cicc: c|imenIc(e:. cue( nc fc(mc |iv(e.
cue( ccmc ccmpcnenIe cc :ccc(c:e.
Fcce :e( ccIc|izccc ce 2 fc(mc::

1. Frutose Frutose 1-Fos|oto {f|cccc. inIe:Iinc e (in:)
f(uIccinc:e: enzimc ccm e|evccc cfiniccce pc(c c f(uIc:e.

/ f(uIc:e pcce :e( fc:fc(i|ccc nc f|cccc c f(uIc:e 1-fc:fcIc pc( umc f(uIccinc:e
e:pecic|. CcnIucc. ncc exi:Ie nenhumc muIc:e ccpcz ce c ccnve(Ie( c f(uIc:e -
fc:fcIc. nem pcce c fc:fcf(uIccinc:e :inIeIizc( c f(uIc:e 1.-Li:fc:fcIc c pc(Ii( cc f(uIc:e
1-fc:fcIc. Em vez ci::c. c f(uIc:e 1-fc:fcIc c|cc|c:e c|ivc c f(uIc:e 1-fc:fcIc em DH/F.
cue enI(c ci(ecIcmenIe nc vic c|icc||Iicc. e em c|ice(c|ceicc. cue p(imei(c
fc:fc(i|ccc em c|ice(c|ceicc 3-fc:fcIc cnIe: ce enI(c( nc vic c|icc||Iicc. / cu:ncic
ce:Ic enzimc {c|cc|c:e) p(cvccc inIc|e(cncic c f(uIc:e.

2. Frutose Frutose -Fos|oto* {Iecicc ccipc:c e m:cu|c)
Hexccinc:e: cpenc: umc f(ccccc m|nimc ce f(uIc:e fc:fc(i|ccc em f(uIc:e -
fc:fcIc pe|c hexccinc:e. pc(cue:
c ccncenI(cccc cc c|icc:e muiIc :upe(ic( c cc f(uIc:e
c cfiniccce cc hexccinc:e pc(c c c|icc:e ce(cc ce 20 veze: :upe(ic( cue
pc(c c f(uIc:e
Nc Iecicc ccipc:c. cnce c ccncenI(cccc ce f(uIc:e excece :uL:Icncic|menIe c ce
c|icc:e. cecc((e c fc(mcccc p(efe(encic|menIe cc f(uIc:e -fc:fcIc. pe|c hexccinc:e.

* EnI(c |ccc nc vic c|icc||Iicc {e pc( ccccc cc fc:fcf(uIccinc:e ccnve(Iicc em 1.-
f(uIc:e Li:fc:fcIc)

|rutose 1-|os|oto d||drox|cetono 3-|os|oto + g||cero|de|do
f(uIc:e 1-fc:fcIc c|cc|c:e
g||cero|de|do + AIF g||cero|de|do 3-|os|oto + ADF
c|ice(c|ceicc cinc:e {I(ic:ecinc:e)
d||drox|cetono 3-|os|oto g||cero|de|do 3-|os|oto
I(ic:e-fc:fcIc i:cme(c:e:
31

Cofobo||smo do go|ocfose
Go|octose:
Fe:u|Ic cc cec(cccccc cc |ccIc:e cc |eiIe
I(cn:fc(mccc nc f|cccc em c|icc:e 1-fc:fcIc cc |cncc ce 4 eIcpc:. pe|c: cuci:
cp(cveiIccc nc vic c|icc||Iicc cu c(mczenccc ccmc c|icccnic.

De:Iinc: cc UDF c|icc:e
S|nIe:e ce c|icccnic
S|nIe:e cc ccicc c|icu(cnicc {ce:cc(ccenc:e N/D
+
cepencenIe)
EnI(c nc vic c|icc||Iicc cI(cv: cc |iLe(Icccc ce c|icc:e 1-fc:fcIc {ccm
I(cn:fc(mcccc em c|icc:e -fc:fcIc e :uL:ecuenIe cp(cveiIcmenIc meIcLc|icc)

De:Iinc: cc UDF cc|ccIc:e
lncc(pc(cccc em e:fincc||picc:
Ccnve(:cc em UDF-c|icc:e meIcLc|i:mc cc: c||cicc:

/ (eccccc ce ccnve(:cc (eve(:|ve| cc cc|ccIc:e em c|icc:e. ccm c c:e :cL c
fc(mc ccnjucccc ccm um nuc|ecIicc. exemp|ificc c: meccni:mc: ce(ci: ce
ccnve(:cc. I(cn:pc(Ie cu incc(pc(cccc cc: c:e: em mc|cu|c: mcic(e:. cue fczem
inIe(vi( c: fc(mc: ccnjucccc: {ccIivccc:) cc: mcnc::ccc(icc: - c: nuc|ecIicc-c:e:

Go|octose + AIF Go|octose 1-|os|oto + ADF
Gc|ccIccinc:e
{ce:envc|vimenIc e c(icncc)
Go|octose 1-|os|oto + bDF-g||cose bDF-go|octose + g||cose 1-|os|oto
Gc|ccIc:e 1-fc:fcIc u(ici| I(cn:fe(c:e
Ob
{/cc|e:cenIe)
Go|octose 1-|os|oto + bIF bDF-go|octose + FF|
UDF - cc|ccIc:e-pi(cfc:fc(i|c:e
bDF-go|octose bDF g||cose
UDF cc|ccIc:e 4-epime(c:e
bDF-g||cose + FF| g||cose 1-|os|oto + bIF
UDF c|icc:e pi(cfc:fcIc:e
G||cose 1-|os|oto g||cose -|os|oto
Fc:fcc|iccmuIc:e
32

Cofobo||smo de oufros oses

1. Mcnc:e f(uIc:e -fc:fcIc

2. G|ice(c| ciic(cxiceIcnc-fc:fcIc
C c|ice(c| {p(ccuIc cc ccIcLc|i:mc |ipicicc) pcce :e( uIi|izccc cpenc: pe|c: Iecicc:
cue pc::uem c|ice(c|cinc:e {p(incipc|menIe f|cccc e (in:).

C c|ice(c|ceicc fc(mccc Ie(c ccmc ce:Iinc: pc::|vei: c c|icc|i:e e c c|iccneccne:e.

Monose monose -|os|oto
Hexccinc:e
Monose -|os|oto |rutose -|os|oto
Fc:fcmcnc:e i:cme(c:e
G||cero| + AIF G||cero| 3-|os|oto + ADF + H
+

c|ice(c|cinc:e
G||cero| 3-|os|oto + NAD
+
D||hrox|cetono-|os|oto
c|ice(c| 3-fc:fcIc ce:ic(ccenc:e
D||hrox|cetono-|os|oto G||cero|de|do
I(ic:e fc:fcIc i:cme(c:e
33

5.3 - V|o dos |os|opentoses
{cu vic cc: hexc:e-mcncfc:fcIc: vic cxiccIivc cc fc:fcc|iccncIc: vic cc: penIc:e:)

V|os dos |os|opentoses: :ecuncic mu|Iifuncicnc| ci:pcn|ve| pc(c c cec(cccccc cc
c|icc:e. Decc((e. Ic| ccmc c c|icc|i:e. nc ciIc:c|. e c :uc ccIiviccce cepence cc
c(ccni:mc e e:Iccc meIcLc|icc. /c ccnI(c(ic cc c|icc|i:e. ncc ce(c ci(ecIcmenIe /IF
mc: um Iipc cife(enIe ce ene(cic meIcLc|icc: c pcce( (ecuIc(.

Object|vos:
CLIenccc ce pcce( (ecuIc( {:cL c fc(mc ce N/DF (ecuzicc) pc(c Lic::|nIe:e
(ecuIc(c: {ex.: :|nIe:e ce ccicc: cc(cc:: e:Ie(cice: e cmincccicc:). :encc
pc(IcnIc muiIc ccIivc cc f|cccc. Iecicc ccipc:c. cc(Iex :up(c-(enc|. c|cncu|c
mcmc(ic. Ii(ciceic. Ie:I|cu|c: e e(iI(cciIc:.
Ccnve(:cc cc: hexc:e: em penIc:e:. ce cue :e ce:Iccc c (iLc:e 5-fc:fcIc.
inci:pen:cve| c :|nIe:e cc: ccicc: nuc|eicc: e nuc|ecIicc: impc(IcnIe: {F/D.
N/D
+
.N/DF
+
). cu :ejc. cLIenccc ce ccucc(e: ccm 3 c 7 cc(Lcnc:.
Cxiccccc cc: penIc:e: em hexc:e: pc(c uIi|izcccc c|icc||Iicc e ccn:ecuenIe
fc(mcccc ce /IF cu (ecene(cccc cc c|icc:e -fc:fcIc pc(c pc:Ie(ic(
:up|emenIcccc cc vic cc: fc:fcpenIc:e:.

/c ccnI(c(ic cc cue :ucece nc G|icc|i:e. ncc ccn:cme /IF. e um p(cce::c
e::encic|menIe ce(cLic. pci: c (ecxiccccc cc: ccenzimc: (ecuzicc: :c pc::|ve|
cI(cv: cc CIE cu ce (ecccce: ce Lic::|nIe:e. cue uIi|izem c N/DFH e ce(em.
pc(IcnIc. N/DF
+
.
/ ccIiviccce ce(c| ce:Ic vic ccnI(c|ccc pe|c ccncenI(cccc ce N/DF
+
{fc:e
cxiccIivc) e pe|c ccncenI(cccc ce :uL:I(cIc: {fc:e ncc cxiccIivc)

Foses
1. Fose ox|dot|vo
/ c|icc:e--fc:fcIc ce:cc(Lcxi|ccc c FiLu|c:e-5-fc:fcIc. p(ececicc pc( 2
(ecccce: ce cxiccccc. ccm c fc(mcccc ce N/DFH

De:ic(ccenc:e: {cc c|icc:e -fc:fcIc e -fc:fcc|iccncIc):
/mLc: ccIucm e:pecificcmenIe ccm c N/DF
+
.
C KM cc ce:ic(ccenc:e cc c|icc:e -fc:fcIc pc(c c N/D
+
ce(cc ce 1000 veze:
:upe(ic( cc cc N/DF
+
.
/ (eccccc ccIc|i:ccc pc( e:Ic enzimc fc(IemenIe iniLicc pc( :uL:Icncic:
cxiccnIe: {cnIi-mc|c(icc:. :u|fcncmicc:. c:pi(inc).
DeIe(minccc: inciv|cuc: cp(e:enIcm ceficincic e(iI(cciIc(ic em ce:ic(ccenc:e
cc c|icc:e -fc:fcIc. ce I(cn:mi::cc he(eciIc(ic.
Fc( expc:iccc c :uL:Icncic: cxiccnIe:. pc(Ie cc: c|cLu|c: ve(me|hc: {ccm
ceficincic enzimcIicc) (cpiccmenIe ce:I(u|cc c(icincncc cnemic.
Nc c(icem ce:Ic: c(i:e: hemc||Iicc: pc(ece e:Ic( um cefeiIc cc (ecuccc cc:
c(upc: SH cc memL(cnc e(iI(cciIc(ic. inci:pen:cve| c mcnuIenccc cc :uc
e:I(uIu(c e funcce:.

o) G||cose -|os|oto NADF
+
-|os|og||cono|octono + NADFH
c|icc:e -fc:fcIc ce:ic(ccenc:e
b) -|os|og||cono|octono + H2O -|os|og|uconoto + H
+

-fc:fc-c|icc-|ccIcnc:e
c) -|os|og|uconoto + NADF
+
k|bu|ose 5-|os|oto + NADFH + H
+
+ CO2
-fc:fcc|uccncIc-ce:ic(ccenc:e
34

Ccc((em c::im. ne:Ic I(cn:fc(mcccc. cuc: cxicccce: ccm fc(mcccc ce N/DFH.
cujc: e|ecI(ce: pcce(cc :e( I(cn:fe(icc: pc(c c :i:Iemc I(cn:pc(Iccc( ce e|ecI(ce: ccm
p(ccuccc ce 3 c 5 /IF.

2. Fose no ox|dot|vo
2.1 l:cme(izcccc em (iLc:e-5-fc:fcIc e xi|u|c:e-5-fc:fcIc
/ (iLu|c:e 5-fc:fcIc pcce :e(vi( ce :uL:I(cIc c 2 enzimc: cife(enIe:. cue c I(cn:fc(mcm
em (iLc:e 5-fc:fcIc cu xi|u|c:e 5-fc:fcIc. /mLc: c: (ecccce: :cc (eve(:|vei:.

umc I(cn:fc(mcccc ceIc:e-c|cc:e. pc( i::c inc|ui um inIe(mecic(ic enecic|.
Fc( veze: c vic inIe((cmpe-:e ccm c fc(mcccc ce (iLc:e 5-fc:fcIc {ex.: cucncc
e:Ie meIcLc|iIc ccIuc ccmc pe(cu(:c( nc :|nIe:e cc: nuc|ecIicc: e ccicc:
nuc|eicc:): ne:Ie cc:c c vic Iem ccmc finc|iccce e::encic| c p(ccuccc ce (iLc:e
5-fc:fcIc.

Ccm c fc(mcccc ce xi|u|c:e 5-fc:fcIc c vic p(c::ecue c(icincncc cccc(e: ccm 3
c 7 cc(Lcnc:.
Ne:Ic ci(cun:Icncic. c: c|u|c: pccem cLIe( muiIc mci: N/DFH cue (iLc:e 5-
fc:fcIc.
E:Ie. em exce::c. cucnIiIcIivcmenIe ccnve(Iicc em meIcLc|iIc: cc c|icc|i:e
{c|ice(c|ceicc 3-fc:fcIc e f(uIc:e -fc:fcIc)

2.2 lnIe(ccnve(:cc cc: penIc:e:-fc:fcIc e cc: hexc:e:-fc:fcIc pc( I(cn:c|cc|izcccc e
I(cn:ceIc|i:cccc

Ne:Ic fc:e ccc((em I(cn:fe(ncic: ce c(upc: ccm 3 cIcmc: ce cc(Lcnc {c(upc
ciic(cxiceIcnc) - I(cn:c|cc|i:cccc {I(cn:c|cc|c:e) - e ccm 2 cIcmc: ce cc(Lcnc
{c(upc c|ice(c|ceicc) - I(cn:ceIc|i:cccc {I(cn:ceIc|c:e).
Ne:Ic eIcpc. c: 3 (ecccce: :cc (eve(:|vei:.
E:Ic eIcpc cepence cc: nece::iccce: cc c|u|c: :e c c|u|c :c p(eci:c( ce
N/DFH e ncc p(eci:c( ce (iLc:e-5-F. e:Ic pcce(c :e( (ecp(cveiIccc.

/ (iLc:e-5-F (eceLe cci: cc(Lcnc: cc xi|u|c:e-5-F
/ I(cn:ceIc|c:e (ecue( c pc(Iicipcccc cc ccenzimc Iicminc pi(cfc:fcIc {IFF).

ccmc :eu c(upc p(c:IIicc. cue ccIuc ccmc I(cn:pc(Iccc inIe(mecic(ic cc c(upc
c|ice(c|ceicc e Mc
2+

Scc I(cn:fe(icc: I(: cc(Lcnc: cc :eccepIu|c:e 7 -F pc(c c c|ice(c|ce|cc-3-F
o) k|bu|ose 5-|os|oto k|bose 5-|os|oto
Fc:fcIcpenIc:e i:cme(c:e
b) k|bu|ose 5-|os|oto X||u|ose 5-|os|oto
Fc:fcpenIc:e epime(c:e
o) X||u|ose 5-|os|oto + r|bose 5-|os|oto g||cero|dedo 3-|os|oto + sedoeptu|ose 7-
|os|oto
I(cn:ceIc|c:e
b) G||cero|dedo 3-|os|oto + sedoeptu|ose 7-|os|oto er|trose 4-|os|oto + |rutose -
|os|oto
I(cn:c|cc|c:e
35

/ I(cn:c|cc|c:e umc enzimc cnc|ccc c c|cc|c:e cc c|icc|i:e. IcnIc num cc:c
ccmc nc cuI(c. c (eccccc ce:enccceccc cpc: |iccccc cc :uL:I(cIc cc c(upc
-cminc ce um (e:|cuc ce |i:inc cc enzimc. Ccc((e enIcc umc (upIu(c :ecuicc ce
umc ccncen:cccc. ce:icnccc: c|cc|icc: {nc c|icc|i:e ccc((ic cpenc: c (upIu(c).

Em conjunto:

2 xi|u|c:e 5-fc:fcIc + (iLc:e 5-fc:fcIc 2 f(uIc:e -fc:fcIc + c|ice(c|ceicc 3-fc:fcIc

V|os dos |os|opentoses e g||co||se
/ f(uIc:e--F e c c|ice(c|ce|cc-3-F pccem :e( uIi|izccc: nc c|icc|i:e pc(c p(ccuccc
ce ene(cic. cu (ecic|ccc: pe|c Necc|icccne:e pc(c fc(mc( ncvcmenIe c|icc:e -
fc:fcIc.
/ ci(ecccc cc f|uxc e ce:Iinc cc G|icc:e -fc:fcIc :cc ceIe(minccc:
e::encic|menIe pe|c nece::iccce ce|u|c( em N/DFH cu (iLc:e 5-fc:fcIc.

SiIucccc 1 - nece::c(ic mci: (iLc:e 5-fc:fcIc cc cue N/DFH
/ c|icc:e -fc:fcIc p(efe(encic|menIe cxicccc pe|c vic c|icc||Iicc. funcicncncc
cpenc: c fc:e ncc cxiccIivc cc vic.
Fc( ccccc cc I(cn:c|cc|c:e e I(cn:ceIc|c:e. cccc 2 mc|cu|c: ce f(uIc:e -fc:fcIc
e umc cc c|ice(c|ceicc 3-fc:fcIc p(cvenienIe: cc c|icc|i:e ccc c(icem c 3
mc|cu|c: ce (iLc:e 5-fc:fcIc. :em fc(mcccc ce N/DFH.
5 G||cose -F + AIF k|bose 5-F + ADF + H
+

SiIucccc 2 - c: cucnIiccce: (ecue(icc: {N/DFH e (iLc:e) :cc ecuivc|enIe:
/ G|icc:e -fc:fcIc cxicccc pe|c vic cc: fc:fcpenIc:e: em cucnIiccce: mcce(ccc:.

G||cose -|os|oto + 2 NADF
+
+ H2O k|bose 5-|os|oto + 2 NADFH + 2H
+
+ CO2

c) X||u|ose 3-|os|oto + er|trose 4-|os|oto g||cero|de|do 3-|os|oto + |rutose -|os|oto
I(cn:ceIc|c:e
36

SiIucccc 3 - (ecue(icc mci: N/DFH cc cue (iLc:e 5-fc:fcIc. ccm cxiccccc ccmp|eIc
ce c|icc:e -fc:fcIc em CC2
F(uIc:e -fc:fcIc e c|ice(c|ceicc 3-fc:fcIc. pc( inve(:cc cc: eIcpc: c|icc||Iicc:.
(ecene(cm c|icc:e fc:fcIc. fczencc-c (eci(cu|c( pe|c vic cc: fc:fcpenIc:e: cI
ccmp|eIc cxiccccc em CC2.
Simu|IcnecmenIe hc fc(mcccc ce N/DFH

g||cose -|os|oto + 12NADF
+
+ H2O r|bose 5-|os|oto + 12 NADFH + 12H
+
+ CO2
r|bose 5-|os|oto 4 |rutose -|os|oto + 2 g||cero|de|do 3-|os|oto
4 |rutose -|os|oto + 2 g||cero|de|do 3-|os|oto + H2O 5 g||cose -|os|oto + F|

Cu :ejc:
G||cose -|os|oto + 12NADF
+
+ 7H2O CO2 + 12 NADFH + 12H
+
+ F|

SiIucccc 4 - (ecue(icc mci: N/DFH e /IF
Ccm c fc(mcccc cc (iLc:e 5-fc:fcIc ce(ccc N/DFH.
/ f(uIc:e -fc:fcIc e c|ice(c|ceicc 3-fc:fcIc fc(mccc: c pc(Ii( cc (iLc:e 5-fc:fcIc
:cc I(cn:fc(mccc: {nc c|icc|i:e) cI pi(uvcIc. ccm p(ccuccc :imu|Icnec ce /IF.

3 g||cose -|os|oto + NADF
+
+ 5 NAD
+
+ 5F| + ADF 5 p|ruvoto + 3CO2 + NADFH + 5
NADH + 8 AIF + 2H2O + 8H
+

8o|ono energt|co

Ccc((em 2 cxicccce:. ccm c fc(mcccc ce N/DFH. cujc: e|ecI(ce: pcce(cc :e(
I(cn:fe(icc: pc(c c CIE ccm c fc(mcccc ce /IF. nc enIcnIc. c ce:Iinc hcLiIuc| cc
N/DFH p(ccuzicc ncc c p(ccuccc ce /IF. mc: :im ccIuc( ccmc cccc( ce
hic(ccnic e e|ecI(ce: nc: Lic::|nIe:e: (ecuIc(c:.
Numc vc|Ic ce cic|c c ecuivc|enIe c umc mc|cu|c ce c|icc:e em cccc :ei:
ccmp|eIcmenIe cxicccc.
Fc( cccc mc|cu|c ce c|ucc:e--F ccmp|eIcmenIe cxicccc c :ei: mc|cu|c: ce
CC2 :cc (ecuzicc: 12 mc|cu|c: ce N/DF
+
ccm c fc(mcccc ce 12 N/DFH.

G||cose--F + 12 NADF
+
5 G||cose--F + CO2 + 12 NADFH + 12H
+
+ F|

37

keoces Enz|mo
Fose ox|dot|vo do v|o
G|icc:e--F + N/DF
+
-Fc:fcc|iccnc|ccIcnc + N/DFH + H
+
G|icc:e -F-
ce:ic(ccenc:e
-Fc:fcc|iccnc|ccIcnc + H2C -fc:fcc|iccncIc + H
+
LccIcnc:e
-fc:fcc|iccncIc + N/DF
+
(iLu|c:e 5-F + CC2 + N/DFH -fc:fcc|iccncIc -
-ce:ic(ccenc:e
Fose no ox|dot|vo do v|o
FiLu|c:e-5-F FiLc:e-5-F

fc:fcpenIc:e-
i:cme(c:e
FiLu|c:e-5-F Xi|u|c:e-5-F fc:fcpenIc:e-
epime(c:e
Xi|u|c:e 5-F+ FiLc:e 5-F SeccepIu|c:e 7-F + G|ice(c|ce|cc 3-F I(cn:ceIc|c:e
SeccepIu|c:e 7-F + G|ice(c|ce|cc 3-F F(uIc:e -F + E(iI(c:e 4-F I(cn:c|cc|c:e
Xi|u|c:e 5-F + E(iI(c:e 4-F F(uIc:e -F + G|ice(c|ce|cc 3-F I(cn:ceIc|c:e

kegu|oo
C fccIc( ce (ecu|cccc mci: impc(IcnIe cc vic cc: penIc:e: cc n|ve| cc N/DF
+
.
cue c cceiIccc( ce e|ecI(ce: nc cxiccccc cc c|icc:e--fc:fcIc c ccicc--
fc:fcc|iccnicc.

C pe(cxicc ce hic(ccnic (emcvicc pe|c c|uIcIicc pe(cxicc:e. C c|uIcIicc cxicccc
(ecuzicc pe|c c|uIcIicc (ecuIc:e. nc p(e:encc ce N/DFH. cujc ccncenI(cccc
ciminui. ccIivcncc c vic. / vic cc: penIc:e: fc:fcIc muiIc impc(IcnIe nc e(iI(cciIc.
pc(c mcnIe( c c|uIcIicc (ecuzicc. pc(c cue e:Ie (emcvc c pe(cxicc ce hic(ccnic.
|e:ivc pc(c c e(iI(cciIc. em e:pecic| pc(c c memL(cnc ce|u|c(.
38

.
.. C
CCo
oot
tto
oob
bbo
oo|
|||
||s
ssm
mmo
oo L
LL|
||p
pp|
||d
dd|
||c
cco
oo

Lipicc::
8icmc|cu|c: ce(ivccc: ce ccicc: cc(cc: {ccicc: mcnccc(Lcxi|icc:) cu ccceic
c|ifcIicc vc(icve| em ccm cimen:ce: vc(icvei:
/pc|c(e: cu hic(cfcLicc:. in:c|vei: em ccuc. mc: :c|vei: em :c|venIe: c(ccnicc:.
Exi:Iem ||picc: {ccicc: cc(cc:. fc:fc||picc:. e:fincc||picc:. ccicc: Li|ic(e: e
cc|e:Ie(c|) cue ccnIm c(upc: pc|c(e:. :encc umc pc(Ie cc mc|cu|c hic(cf||icc
encucnIc c cuI(c pc(Ie hic(cfcLicc - :cc :uL:Icncic: c|ifcIicc:. /::im :e exp|icc
c c(ienIcccc ce:Ic: :uL:Icncic: em ccnI(ccIc ccm c ccuc. fc(mcncc-:e
memL(cnc: |ipicicc:. emu|:ce:. mice|c: e |ipc::cmc:.

Funccc Lic|ccicc:
CcmpcnenIe: cc: memL(cnc: ce|u|c(e:. junIcmenIe ccm c: p(cIe|nc: {fc:fc||picc:
e cc|e:Ie(c|) impc(Icncic cc pc(Ie hic(cfcLicc
Ccmpc:Ic Liccu|micc mci: cc|c(icc. :encc c p(incipc| fc(mc ce c(mczencmenIc
{I(cci|c|ice(cie:). e ce(cccc ce ene(cic meIcLc|icc cI(cv: cc -cxiccccc ce
ccicc: cc(cc:.
McnIm c inIec(iccce cc :upe(f|cie c|vec|c( nc: pu|mce:
Sc|uLi|izcm c: :uL:Icncic: ncc pc|c(e: cc: f|uicc: cc(pc(ci:.
Scc ccmpcnenIe: cc :i:Iemc ce I(cn:pc(Ie ce e|ecI(ce: nc inIe(ic( cc memL(cnc
miIcccnc(ic| {uLicuincnc)
Fc(mcm umc pe||cu|c pc(IecIc(c {i:c|cnIe I(micc) :cL(e c epice(me ce muiIc:
cnimci: {Iecicc: ccipc:c)
Im funcce: e:pecic|izccc: ccmc hc(mcnc: e viIcminc: |ipc::c|vei:: ViIcminc /.
D. E e K: cccce: meIcLc|icc: e:pecic|izccc:: hc(mcnc: {e:Ie(cice:. eicc:cncice:)
cu men:ccei(c: ce ccccc hc(mcnc| {fc:fc||picc: e ce(ivccc:).

I. G||cr|dos
Mcnc-. ci-. cu I(i:Ie(e: cc c|ice(c| ccm ccicc: cc(cc: ce:icncm-:e
(e:pecIivcmenIe mcnccci|c|ice(c|. cicci|c|ice(c| e I(icci|c|ice(c|.
C: mcnccci|c|ice(ci: e cicci|c|ice(ci: e:Icc p(e:enIe: :cmenIe em cucnIiccce:
(e|cIivcmenIe pecuenc: e ccc((em em c(cnce pc(Ie. ccmc inIe(mecic(ic:
meIcLc|icc: nc Lic::|nIe:e e nc cec(cccccc cc: ||picc: cue ccnIm c|ice(c|.
C: I(icci|c|ic(icc: :cc c: mci: f(ecuenIe:.
C f|cccc e c Iecicc ccipc:c {nc :uc mcic( pc(Ie) (ec|izcm c p(cce::c ce
ccnve(:cc ce ccicc: cc(cc: em I(ic|ic(icc:.
C Iecicc ccipc:c {ccn:IiIu|cc pc( ccipcciIc:) um Iecicc ccnjunIivc
e:pecic|izccc. ce:Iinccc c :|nIe:e. cc c(mczencmenIc e c hic(c|i:e cc:
I(ic|ic(icc:.
C: I(ic|ic(icc: :cc c(mczenccc: ccmc ccI|cu|c: ||cuicc: nc ciIcp|c:mc. mc: i:Ic
ncc um Iipc ce c(mczencmenIc e:IcIicc". umc vez cue :cc (ecic|ccc:. em
pcucc: cic:. /::im numc :iIucccc hcmec:IcIicc. ccc((e :|nIe:e e cueL(c ccnI|nuc
cc: I(ic|ic(icc:. nc Iecicc ccipc:c.

VcnIccen: ene(cIicc: cc: cc(ccIe(|:Iicc: ncc-pc|c(e: cc: I(ic|ic(icc::
1. C meIcLc|i:mc ce I(icci|c|ice(ci: pe(miIe c fc(mcccc ce mcic( cucnIiccce ce
/IF cue c meIcLc|i:mc cc c|icccnic.
2. C: I(icci|c|ice(ci: pccem :e( c(mczenccc: :em e:Ic(em c::cciccc: c ccuc.
encucnIc cue c c|icccnic muiIc hic(cf||icc e nece::iIc cc ccL(c cc :eu
pe:c em ccuc. cucncc e:Ic c(mczenccc em Iecicc:. /::im. c ecuivc|enIe ce
cucnIiccce ce ene(cic c(mczenccc ccmc c|icccnic hic(cIccc pe:c(c 4
veze: mci: cc cue :e e:Iive::e c(mczenccc ccmc I(icci|c|ice(c|.
39

/ceni|cic|c:e l
/ceni|cic|c:e /
/MF
/MFc
Fc:fc(i|c:e
Cinc:e
Lipc:e l
Lipc:e /
/c(enc|inc
G|ucccinc
3. Numc pe::cc ccm 70Kc. cpenc: 350c ce cc(Lchic(cIc: e:Icc c(mczenccc:. c
cue (ep(e:enIc cpenc: 1400Kcc| ce ene(cic. encucnIc cue c: (e:e(vc: ce
||picc: cc(cnIem c :cL(evivncic pc(c vc(ic: :emcnc: :em c|imenIcccc.
4. C: I(ic|ic(icc: pc::uem muiIc mci: cIcmc: ce cc(Lcnc pc( uniccce ce pe:c
cue c: pc|i::ccc(icc:. pe|c cue pe(miIem ccn:IiIui( (e:e(vc: ene(cIicc: ccm
umc mc::c muiIc menc(.

H|dro||se dos tr|g||cr|dos (L|po||se)
Ccc((ncic:
8cixc (e:e(vc ene(cIicc
Jejum
Exe(c|cic vicc(c:c
Fe:pc:Ic cc :I(e::
F(cce::c hc(mcnc-cepencenIe ccncucenIe c ccicc: cc(cc:. c|ice(c|.
C: I(ic|ic(icc: exccenc: I(cn:pc(Iccc: pe|c: cui|cmic(c e c: encccenc:
I(cn:pc(Iccc: pe|c: VLDL :cc ce:I(u|cc: nc enccI|ic pe|c |ipcp(cIeinc |ipc:e.

E:Ic |ccc|izccc nc :upe(f|cie cc: c|u|c: cc enccI|ic cc: ccpi|c(e:.
/ |ipc:e ccIivccc pe|c c/MF.
C: c|icccc(Iiccice:. c: ccIecc|cminc:. e c c|ucccinc pc( ccIivc(em c
cceni|cic|c:e. cumenIcm c ccccc cc |ipc:e. e ccn:ecuenIemenIe c |iLe(Icccc
ce ccicc: cc(cc: pc(c c p|c:mc. pe|c Iecicc ccipc:c.
/ in:u|inc Iem um efeiIc ccnI(c(ic.

C: ccicc: cc(cc: |iLe(Iccc::
Ci(cu|cm nc :cncue c::cciccc: c c|Luminc
/I(cve::cm c: memL(cnc: ce|u|c(e: pc( ccI(cn:pc(Ie ce :ccic
Ncc uIi|izcccc nc c(eL(c nem nc: c|cLu|c: ve(me|hc:.
C: mcnccci|c|ice(ci: pccem pc::c( pc(c cenI(c cc: c|u|c: cu :e(em hic(c|izccc:
pe|c mcnccci|c|ice(c| hic(c|c:e.
Ccmc c Iecicc ccipc:c ncc Iem c|ice(c|cinc:e. c c|ice(c| (e:u|IcnIe cc
ccIcLc|i:mc. ncc pcce :e( uIi|izccc e cifunce-:e pc(c c p|c:mc. pc(c :e( uIi|izccc
pe|c: Iecicc: cue ccnIm c|ice(c|cinc:e.

Ir|oc||g||cero|s + 3H2O 3 c|dos gordos + G||cero|
Lipc:e
40

/p(cveiIcmenIc meIcLc|icc cc c|ice(c|

UIi|izcccc cc c|ice(c| nc hepcIcciIc
/cccc cc c|ice(c| cinc:e {cu:enIe nc ccipcciIc) - fc:fc(i|cccc pc( c|ice(c|-cinc:e
em c|ice(c| 3-fc:fcIc - ccm I(cn:fc(mcccc ci(ecIc cu inci(ecIc {cpc: I(cn:fe(ncic ce
cIcmc: ce hic(ccnic pc(c c ccceic (e:pi(cIc(ic) em ciic(cxiceIcnc-fc:fcIc {e
:uL:ecuenIemenIe cp(cveiIcmenIc pe|c c|icc|i:e/c|iccneccne:e)

De:Iinc: cc c|ice(c| 3-fc:fcIc
S|nIe:e ce I(icci|c|ice(ci:
S|nIe:e ce fc:fc||picc:
SuL:I(cIc cc c|iccneccne:e nc f|cccc
S|nIe:e ce c|icc:e

II. c|dos gordos
cicc: mcnccc(Lcxi|icc c|ifcIicc
Fcce |iLe(Ic(-:e pc( hic(c|i:e c pc(Ii( ce c|ec: cu cc(cu(c: ncIu(ci:.
C: mci: impc(IcnIe: e f(ecuenIe: Im ccceic |inec(. ncc (cmificccc ccm n pc(
ce cIcmc: ce cc(Lcnc {enI(e 4 e 30)
Fccem :e( :cIu(ccc: {cpenc: ccm |icccce: :imp|e:) cu in:cIu(ccc: {ccm |icccce:
cup|c: e I(ip|c:)
Fccem :e( hic(cxi|ccc:.

Ncmenc|cIu(c
/ nume(cccc cc: cc(Lcnc: cc: ccicc: cc(cc: fcz-:e c pc(Ii( cc cc(Lcxi|c Ie(minc|
{C1) pc(c c c(upc CH3 {Cn).
C: cc(Lcnc: 2 e 3 :cc chcmccc: ce e . (e:pecIivcmenIe.
Em cuc:e Iccc: c: ccicc: cc(cc: in:cIu(ccc: c cup|c |iccccc Iem i:cme(ic ci:.
Fc(c (ep(e:enIc( c: |icccce: cup|c:: n IcIc| ce cIcmc: ce cc(Lcnc : n ce
|icccce: cup|c: {c|cc(i:mc{:) cc((e:pcncenIe{:) cc 1 cc(Lcnc cc |iccccc cup|c).
Ex.: 18:2{.12)
Cucncc ccnvenienIe cc(ccIe(izc( c ccicc cc(cc c pc(Ii( cc exI(emiccce mci:
p(cximc cc c(upc meIi| Ie(minc|. u:c-:e 18:2 n {cu c) .. c :icnificc( ccicc cc(cc
ccm 18 cIcmc: ce cc(Lcnc 2 cuc: cup|c: |icccce:. enccnI(cncc-:e c mci:
p(cximc cc c(upc meIi|c c ci:Icncic ce cIcmc: ce cc(Lcnc.

Cotobo||smo dos c|dos gordos
Lccc|izcccc: mcI(iz miIcccnc(ic|

Foses
1. Act|voo dos c|dos gordos
/ ccIivcccc cc: ccicc: cc(cc: (ecue( ene(cic fc(necicc pe|c /IF
Ccc((e nc memL(cnc miIcccnc(ic| pc( ccccc cc cci|ccenzimc /-:inIeIc:e
E:Ic enzimc ccIc|i:c c fc(mcccc ce umc |iccccc {:inIeIc:e) Iic:Ie( muiIc
ene(cIicc. |ccc c /IF hic(c|i:ccc c /MF {ecuivc|enIe c hic(c|i:e ce 2 /IF em 2
/DF)

k-COO
-
+ CoA + AIF + H2O Ac||-coA + AMF + 2 F| + 2H
+

cci|ccenzimc /-:inIeIc:e
41

2. Ironsporte dos mo|cu|os de Ac||-CoA
/ memL(cnc cc miIcccnc(ic impe(mecve| cc: cci|-Cc/ ce ccceic |cncc. mc:
ncc cc: cci|-Cc/ ce ccceic cu(Ic.
/::im. c: cci|-Cc/ ce ccceic |cncc pc(c enI(c(em nc miIcccnc(ic Im cue (ecci(
ccm um cmincccicc e:pecic|". c corn|t|no. |iLe(Icncc c ccenzimc /.
Umc enzimc. c Cc(niIinc Fc|miIci| I(cn:fe(c:e l {CFI l). e:Ic c::cciccc c
memL(cnc exIe(nc cc miIcccnc(ic e ccnve(Ie c(upc cci|-Cc-/ ce ccceic |cncc
c /ci|-Cc(niIinc. :encc ce:Ic fc(mc ccpcze: ce peneI(c( nc miIcccnc(ic.
/ cc(niIinc e:Ie(ificccc I(cn:pc(Iccc pc(c cenI(c cc miIcccnc(ic pc( um
I(cn:pc(Iccc( e:pec|ficc - Cc(niIinc-cci|cc(niIinc I(cn:|ccc:e.
DenI(c cc miIcccnc(ic. c cc(niIinc I(cn:fe(e c c(upc cci|c pc(c umc cuI(c
mc|cu|c ce Cc/. (eccccc ccIc|i:ccc pe|c Cc(niIinc Fc|miIci| I(cn:fe(c:e ll {CFI
ll).
/ cc(niIinc |iv(e vc|Ic enIcc pc(c c ciIcp|c:mc cI(cv: cc I(cn:pc(Iccc(.
Ne:Ie p(cce::c ncc exi:Ie I(cn:pc(Ie ce Cc/ pc(c cenI(c cc miIcccnc(ic. c::im
c: (e:e(vc: ciIcp|c:mcIicc e miIcccnc(ic| ce Cc/ ncc :e mi:Iu(cm.
/ enI(ccc ce ccicc: cc(cc: ce ccceic cu(Ic incepencenIe cc cc(niIinc.
pc(cue e|e: c(uzcm c memL(cnc miIcccnc(ic| inIe(nc ci(ecIcmenIe e :cc
ccIivccc: cc: :eu: cci|-Cc/. nc ccmpc(IimenIc cc mcI(iz miIcccnc(ic|.

3. -ox|doo dos oc||-CoA

C cci|-cc/ cue fci fc(mccc cxicccc pe|c enzimc
cci| Cc/ ce:ic(ccenc:e. umc f|cvcp(cIe|nc cue u:c
F/D ccmc cceiIccc( ce e|ecI(ce:.
C F/DH2 fc(mccc I(cn:pc(Ic c: e|ecI(ce: pc(c c
ccceic ce I(cn:pc(Ie ce e|ecI(ce:. (ecene(cncc F/D.
Fc(mcccc ce um ccmpc:Ic ccm |iccccc cup|c.

.-I(cn:-enci|-Cc/ fc(mccc nc (eccccc cnIe(ic(
hic(cIccc.
ScIu(cccc cc |iccccc c|cenc
E:Ic (eccccc e:Ie(ece:pec|ficc. IcnIc cue c i:cme(c
L c p(ccuIc cucncc c cup|c |iccccc hic(cIccc.

E:Ic enzimc (ecu|c c e:Ie(ece:pecificiccce ce:Ie
p(cce::c. pci: e:pec|ficc pc(c c i:cme(c L.
Ac||-CoA + FAD , -trons-eno||-CoA + FADH2
cci| Cc/ ce:ic(ccenc:e
, -trons-eno||-CoA + H2O L- -h|drox|oc||-CoA
enci|-Cc/ hic(c|c:e
L- -h|drox|oc||-CoA + NAD
+
-cetooc||-CoA + NADH + H
+

L--hic(cxicci|-Cc/ ce:ic(ccenc:e
4 reoco: -cetooc||-CoA + CoA$H Ac|| CoA (com menos 2C) + Acet|| CoA
Iic|c:e
42

Conc|uso
/: (ecccce: :cc cxicccc: pc( cife(enIe: enzimc: ccn:ccnIe c Icmcnhc cc
ccceic cc /ci|-Cc/.
/ ce:icncccc -cxiccccc cecc((e ce c enI(ccc ce cxicnic. nc :ecuncc
cxiccccc. cccnIece( c n|ve| cc cc(Lcnc {c n 3 cc ccceic).
/ :ecuncic cc -cxiccccc ce(c cceIi|-cc/. F/DH2 e N/DH.
C cci|ccenzimc / fc(mccc. ccm menc: 2 cIcmc: ce cc(Lcnc. enI(c(c ncuI(c
cic|c ce cxiccccc.
/: enzimc: cue inIe(vm nc -cxiccccc e:Icc |ccc|izccc: nc: miIcccnc(ic:. c cue
pe(miIe c ccnjuccccc. ccm e|evccc (encimenIc. cc: p(cce::c: ce cxiccccc cc:
ccmpc:Ic: ene(cIicc: ccm c p(ccuccc ce /IF.

kegu|oo do metobo||smo dos c|dos gordos
kegu|oo de curto duroo - Hormono dependente
Hc(mcnc:: Epinef(inc. Nc(-epinef(inc. G|icccinc
/cIivcccc cc fc:fc(i|cccc cc |ipc:e Hic(c|i:e cc: I/G nc ccipcciIc
lniLiccc cc /ceIi|-Cc/ cc(Lcxi|c:e
Diminuiccc cc ccncenI(cccc ce Mc|cni|c-Cc/ |evcnIc c iniLiccc cc CFI l
Insu||no
/cIivcccc cc /ceIi|-Cc/ cc(Lcxi|c:e {in:u|inc cepencenIe) e:Iimu|cccc
cc :|nIe:e ce ccicc: cc(cc:
lniLiccc cc fc:fc(i|cccc cc |ipc:e

8o|ono energt|co do -ox|doo
Cccc -cxiccccc (e:u|Ic nc fc(mcccc ce:
F/DH2 e N/DH Ccceic I(cn:pc(Iccc(c ce e|ecI(ce: Fc(mcccc.
(e:pecIivcmenIe. ce 1.5 e 2.5 mc|cu|c: ce /IF {IcIc| = 4 /IF)
/ceIi|-cc/ Cic|c ce K(eL: 10 /IF.
Ex.: C cicc pc|m|Iicc {1 C) ccmp|eIc c :uc cxiccccc ccm c fc(mcccc ce {7x4) +
{8x10) = 108 /IF: e. cecuzicc c /IF nece::c(ic pc(c c ccIivcccc. Ie(emc: um ccnhc
ce 107 /IF.
C (encimenIc ene(cIicc pc( cIcmc ce cc(Lcnc . pc(IcnIc. ce pe(Ic ce 7
/IF {107/1) pc(c c cxiccccc cc: ccicc: cc(cc:.

Ox|doo de c|dos gordos soturodos com um nmero |mpor de tomos de corbono
Um ccicc cc(cc ce ccceic impc( cc c(icem. nc |Iimc (cncc cc cic|c c cceIi|-
Cc/ e p(cpicni|-Cc/.
Fc(c cue e:Ie pc::c :e( uIi|izccc pe|c cic|c ce K(eL:. nece::c(ic ccicicnc(-|he
um cIcmc ce cc(Lcnc. c cue feiIc pc( cc(Lcxi|cccc.
C meIi|mc|cni| c::im fc(mccc enIcc (ec((cnjccc c :uccini| Cc/. numc (eccccc
c::i:Iicc pe|c ccLc|cminc {c viIcminc 812).

C :uccini|-Cc/. c|m ce :e( um inIe(mecic(ic nc cic|c ce K(eL:. um p(ecu(:c( cc
heme. Umc ceficincic em viIcminc 812 (e:u|Ic pc( i::c nc cificu|ccce ce :inIeIizc(
heme. i:Ic . nc ce:envc|vimenIc ce cnemic pe(nicic:c.
Nc ciIcp|c:mc pcce IcmLm :e( ce:cc(Lcxi|ccc c ccicc pi(vicc pe|c enzimc
mc|icc. ccm p(ccuccc :imu|Icnec ce N/DFH. C ccicc pi(vicc pcce enI(c( nc
miIcccnc(ic. e :e( ccmp|eIcmenIe cxicccc c CC2 pe|c cic|c ce K(eL:.
Frop|on|| CoA + HCO3
-
+ AIF D-Met||mo|on||-CoA + ADF + F|
F(cp(icni| Cc/ Cc(Lcxi|c:e {LicIinc)

D-Met||mo|on||-CoA L-Met||mo|on||-CoA
MeIi|mc|cni|-Cc/ epime(c:e {(ccemc:e)

L-Met||mo|on||-CoA $ucc|n||-CoA
MeIi|mc|cni|-Cc/ muIc:e {ccLc|cminc)
43

Ox|doo dos c|dos |nsoturodos
Ccc((em ccui. c|m cc: 4 (ecccce: e:Iucccc: pc(c c: ccicc: cc(cc: :cIu(ccc:.
mci: cuc: (ecccce: ccm c inIe(venccc ce i:cme(c:e:.
/ exi:Incic cc |iccccc cup|c impece c fc(mcccc cc |iccccc . cu {2:3). cue nc
cxiccccc cc: ccicc: :cIu(ccc: :e fc(mc nc p(imei(c (eccccc.
E:Ic :iIucccc u|I(cpc::ccc pe|c inIe(venccc ce i:cme(c:e: cue ccnve(Iem c
|iccccc cup|c ci:-3 em I(cn:-2 {umc i:cme(c:e ci:-I(cn: e umc epime(c:e).
/ -cxiccccc pcce enIcc p(c::ecui( cc fc(mc hcLiIuc|.
E:Ie e:cuemc ce cec(cccccc vc|icc pc(c Iccc: c: ccicc: cc(cc: in:cIu(ccc:.
cuc|cue( cue :ejc c |ccc|izcccc cc: cup|c: |icccce:.

III. Cetognese

Loco|: miIcccnc(ic hepcIicc
Umc c(cnce cucnIiccce cc cceIi|-Cc/ p(ccuzicc pe|c -cxiccccc cc: ccicc:
cc(cc: nc: miIcccnc(ic: cc f|cccc ccnve(Iicc em ccicc cceIcccIicc e ccicc
-hic(cxiLuI|(icc. cencminccc: pc( cc(pc: ceIcnicc: {mc|cu|c: hic(c::c|uvei:)
/ cceIcnc p(cvm cc ce:cc(Lcxi|cccc e:pcnIcnec cc ccicc cceIcccIicc.
E:Ie: ccmpc:Ic: pccem :e( u:ccc: pe|c cc(cccc e pe|c: m:cu|c: e:cue|Iicc:
pc(c p(ccuzi( ene(cic. C c(eL(c. cue nc(mc|menIe cepence cc c|icc:e ccmc
fcnIe ce ene(cic. pcce IcmLm uIi|izc( cc(pc: ceIcnicc: cu(cnIe um jejum
p(c|cncccc.
/ ceIccne:e ccc((e nc inIe(ic( cc: miIcccnc(ic:.

1 Etopo: Ccncen:cccc ce cuc: mc|cu|c: ce cceIi|-Cc/

2 Etopo: Fc(mcccc cc ccicc cceIcccIicc {2 vic:)

Nc f|cccc. cue {c cife(encc ce cuI(c: Iecicc:) ccnIm umc ce:cci|c:e. ve(ificc-:e
c fc(mcccc ce ccicc cceIcccIicc ci(ecIcmenIe. pe|c pe(cc ce Cc/.
E:Ie -ceIcccicc ccnve(Iicc em -hic(cxiLuI|(icc e cceIcnc e. ccmc e:Ie:
p(ccuIc: :cc meIcLc|izccc: ccm cificu|ccce nc f|cccc. cifuncem-:e pc(c c
ci(cu|cccc.

C ccicc cceIcccIicc IcmLm fc(mccc nc f|cccc pe|c fc(mcccc ce 3-hic(cxi-
3-meIi|-c|uIc(i|-Cc/.
/ ccncen:cccc ce cuI(c mc|cu|c ce cceIi|-Cc/ ccm c /ceIccceIi|-Cc/ p(ccuz
3-hic(cxi-3-meIi|-c|uIc(i|-Cc/ {HMG-Cc/). ccIc|i:ccc pe|c HMG-Cc/ :inIeIc:e.

2 Acet||-CoA Acetoocet||-CoA + CoA
-CeIcIic|c:e
o) Acetoocet||-CoA c|do ocetooct|co + CoA
De:cci|c:e
b1) Acetoocet||-CoA + Acet||-CoA 3-h|drox|-3-met||-g|utor||-CoA + CoA
HMG-Cc/ :inIeIc:e
b2) 3-h|drox|-3-met||-g|utor||-CoA c|do ocetooct|co + ocet||-CoA
HMG-Cc/ Lic:e
44

3 Etopo: Fc(mcccc ce ccicc -hic(cxiLuI|(icc e cceIcnc

/ -hic(cxiLuIi(cIc ce:ic(ccenc:e {p(e:enIe nc miIcccnc(ic) (ecu|ccc pe|c
(e|cccc miIcccnc(ic| ce N/DF
+
/N/DFH. i:Ic . pe|c pcIencic| (eccx.
C ccicc -hic(cxiLuI|(icc . cucnIiIcIivcmenIe. c cc(pc ceIcnicc p(eccmincnIe.
p(e:enIe nc ceIc:e.

IV. bt|||dode metobo||co dos corpos ceton|cos
/ cceIcnc. fc(mccc em menc( cucnIiccce. exc|ccc nc (e:pi(cccc e e|iminccc
nc u(inc.
C: ccicc: cceIcccIicc e -hic(cxiLuI|(icc :cc inci:pen:cvei: pc(c c c(ccni:mc
humcnc. Umc vez I(cn:pc(Iccc: cI(cv: cc :cncue pc(c c: cuI(c: Iecicc:. :cc
cxicccc: cI(cv: cc cic|c ce K(eL:.
C f|uxc ce cc(pc: ceIcnicc: cc f|cccc (e:u|Ic ce um meccni:mc enzimcIicc
ccIivc nc f|cccc pc(c c p(ccuccc ce cc(pc: ceIcnicc:. cccp|ccc c ccIiviccce
ccn:ice(cve|menIe Lcixc cc: enzimc: (e:pcn:cvei: pe|c :uc uIi|izcccc. / :iIucccc
inve(:c ccc((e nc: Iecicc: exI(c-hepcIicc:.
/pe:c( cc f|cccc e:Ic( ecuipccc ccm meccni:mc enzimcIicc ccIivc pc(c c
p(ccuccc ce ccicc cceIcccIicc c pc(Ii( cc cceIccceIi|-Cc/. c ccicc
cceIcccIicc. umc vez fc(mccc ncc pcce :e( (eccIivccc ci(ecIcmenIe nc
f|cccc. excepIc nc ciIc:c|. cnce e:Ie ccmpc:Ic pe(cu(:c( nc :|nIe:e cc
cc|e:Ie(c|. mc: e:Ic vic muiIc menc: ccIivc cc cue c ceIccne:e. l:Ic exp|icc c
p(ccuccc efecIivc ce cc(pc: ceIcnicc: pe|c f|cccc.
Umc vez cL:c(vicc. (ecce nc miIcccnc(ic ccm c :uccini|-Cc/. :encc c Cc/
I(cn:fe(icc pc(c fc(mc( /ceIccceIi|-Cc/ e ccicc :uccinicc. pe|c ccccc cc
:uccini|-Cc-/ cceIccceIcIc-Cc/ I(cn:fe(c:e.
C cceIccceIi|-Cc/ pcce :e( c|ivccc em cuc: mc|cu|c: ce cceIi|-Cc/.
C ccicc -hic(cxiLuI|(icc ccnve(Iicc c ccicc cceIcccIicc pe|c -hic(cxiLuIi(cIc
ce:ic(ccenc:e. nc p(e:encc ce N/D
+
.
C cceIccceIi|-Cc/ fc(mccc ne:Ic: (ecccce: cincicc pe|c Iic|c:e em /ceIi|-
Cc/. :encc e:Ie cxicccc nc cic|c ce K(eL:.
C cceIi|-Cc/ fc(mccc pcce(c enI(c(. nc cic|c ce K(eL:. ce:ce cue c meIcLc|i:mc
c|ic|cicc p(cpc(cicne ccncicce: pc(c Ic| - impc(Icncic cc ccicc cxc|cccIicc -
cc| cue c Iec( ce c|icc:e nc :cncue Ienhc ce :e mcnIe( c um n|ve| m|nimc.
C cceIccceIcIc (ecu|ccc( neccIivc cc |ipc|i:e nc: ccipcciIc:.
Em exce::c. c: cc(pc: ceIcnicc: ccicificcm c meic ce|u|c( ccm p(eju|zc mc(cccc
cc: :uc: funcce:. :encc ccu:c ce umc :iIucccc ce:icnccc ccicc:e ceIcnicc
{f(ecuenIe nc: ccenIe: ccm cicLeIe: ce:ccmpen:ccc:).

c|do ocetooct|co + NADFH + H
+
c|do -h|drox|butr|co + NADF
+

-hic(cxiLuIi(cIc ce:ic(ccenc:e
c|do ocetooct|co Acetono + CO2
45

kegu|oo:
1 Etopo (tec|do od|poso)
/ ceIccne:e ncc ccc((e nc :e( humcnc. c menc: cue hcjc um cumenIc
ccn:ice(cve| nc n|ve| ce ccicc: cc(cc: ci(cu|cnIe:. cue :u(cem cc |ipc|i:e ce
I(icci|c|ice(ci:. nc Iecicc ccipc:c.
C f|cccc ccpcz ce (eIi(c( ce(cc ce 307 cu mci: cc IcIc|iccce cc: ccicc:
cc(cc: |iv(e:.
2Etopo:
F(cce::c-:e cpc: c ccpIcccc cc: ccicc: cc(cc: |iv(e: e cc :uc ccIivcccc.
Ne:Ic c|Iu(c. pccem Ie( 2 ce:Iinc:: cu :cc e:Ie(ificccc: c I(icci|c|ice(ci: cu :cc
cxicccc: c cceIi|-Cc/. Fc( :uc vez. c /ceIi|-Cc/ cxicccc nc cic|c ce K(eL:. cu
enI(c nc ceIccne:e.
C ccnI(c|c cc-:e. :cL(e c cc(niIinc pc|miIi| I(c:nfe(c:e l. nc memL(cnc
miIcccnc(ic|. E:Ic enzimc (ecu|ccc pe|c Mc|cni|-Cc/. cujc ccncenI(cccc
cumenIc nc e:Iccc ce Lcc c|imenIcccc e c iniLe.
Ne:Ic: ccncicce: cuc:e Iccc: c: ccicc: cc(cc: cue :e enccnI(cm nc f|cccc :cc
e:Ie(ificccc: c cci|c|ice(ci: e I(c:npc(Iccc: pc(c c: Iecicc: exI(c-hepcIicc: vic
VLDL.
CcnIucc. cucncc c ccncenI(cccc ce ccicc: cc(cc: |iv(e: cumenIc cevicc c
fcme. c cceIi|-Cc/ cc(Lcxi|c:e iniLicc ci(ecIcmenIe pe|c cci|-Cc/ e c Mc|cni|-
Cc/ ciminui. e c::im c Cc(niIinc Fc|miIci| I(cn:fe(c:e l ceixc ce e:Ic( iniLicc.
pe(miIincc c -cxiccccc.
E:Ie: cccnIecimenIc: :cc (efc(cccc: em jejum. pe|c ciminuiccc cc (e|cccc
in:u|inc/c|ucccinc. |evcncc cc cumenIc cc |ipc|i:e cc Iecicc ccipc:c. ccm c
|iLe(Icccc ce ccicc: cc(cc: |iv(e: e iniLiccc ce cceIi|-Cc/ cc(Lcxi|c:e nc f|cccc.
3Etopo
Ccn:i:Ie nc fc(mcccc ce cc(pc: ceIcnicc: c pc(Ii( ce cceIi|-Cc/ cu nc :uc
uIi|izcccc nc cic|c ce K(eL:.
Ccm c cumenIc cc ccncenI(cccc :(icc ce ccicc: cc(cc: |iv(e:.
p(cp(ccicnc|menIe mci: ccicc: cc(cc: :cc ccnve(Iicc: c cc(pc: ceIcnicc: e
menc: cxicccc nc cic|c ce K(eL:.
/ ci:I(iLuiccc ce cceIi|-Cc/ enI(e c vic ceIccnicc e c ce cxiccccc (ecu|ccc
ce mccc cue c ene(cic IcIc| ccnve(Iicc em /IF cue (e:u|Ic cc cxiccccc ce
ccicc: cc(cc: pe(mcnece ccn:IcnIe.

/ ccmp|eIc cxiccccc ce umc mc|e ce ccicc pc|m|Iicc envc|ve c p(ccuccc ce 12
mc|e: ce /IF vic -cxiccccc e p(ccuccc ce CC2 nc cic|c ce K(eL:. encucnIc cue
:cmenIe 33 mc|e: ce /IF :cc p(ccuzicc: cucncc c ccicc cceIcccIicc e c p(ccuIc
finc| e :cmenIe 21 mc|e: cucncc c -hic(cxiLuI|(icc. /::im. c ceIccne:e pcce :e(
ccn:ice(ccc ccmc um meccni:mc cue pe(miIe cc f|cccc cxicc( c(cnce:
cucnIiccce: ce ccicc: cc(cc: cenI(c cc :i:Iemc ce fc:fc(i|cccc cxiccIivc. :em
cumenIc( c cc:Ic ce ene(cic.
46

7
77.
.. C
CCo
oot
tto
oob
bbo
oo|
|||
||s
ssm
mmo
oo F
FFr
rro
oot
tte
ee|
||c
cco
oo

C niI(ccnic cImc:f(icc. cnIe: ce :e( uIi|izccc pe|c: cnimci: Iem cue :e( (ecuzicc
ce N2 c NH3 {pc( mic(cc(ccni:mc:. p|cnIc: e ce:cc(cc e|cI(icc) e c::im pcce :e(
incc(pc(ccc em cmincccicc: e p(cIe|nc:.
C: :e(e: humcnc: cpenc: ccn:ecuem :inIeIizc( 11 cc: 20 cmincccicc: nece::c(ic:
pc(c c :|nIe:e ce p(cIe|nc:.
Am|noc|dos essenc|o|s: cmincccicc: cue ncc pccem :e( :inIeIizccc: e Im cue
:e cLIe( cI(cv: cc cieIc.

kenovoo Frote|co (furnover)
/: p(cIe|nc: cc(pc(ci: e:Icc em ccn:IcnIe :|nIe:e e cec(cccccc {ce(cc ce 1-27.
:cL(eIucc ce c(icem mu:cu|c()
8o|ono ozotodo: exi:Ie cucncc um ccu|Ic :cuccve| ccm umc cieIc vc(iccc ince(e c
me:mc cucnIiccce ce czcIc cue exc(eIc. Aporte: d|eto: $odo: ure|o
8o|ono ozotodo negot|vo: exc(eIccc mci: czcIc cue ince(icc. Ccc((e em jejum e
nc enve|hecimenIc. em cue c: ccceic: ce cc(Lcnc cc: cmincccicc: :cc nece::c(ic:
pc(c c c|iccneccne:e e c cmcnic |iLe(Iccc exc(eIccc nc u(eic e ncc
incc(pc(ccc. Aporte < $o|do
8o|ono ozotodo pos|t|vo: c incc(pc(cccc ce cmincccicc: em p(cIe|nc: mcic( cue
c :uc cueL(c. ccc((encc em pe(|ccc: ce c(e:cimenIc cc(pc(c| cc: c(icncc: e
ccc|e:cenIe:. nc c(cvicez e nc c|imenIcccc cpc: c jejum. Aporte > $o|do

Degrodoo prote|co
/ cec(cccccc vc(ic ccm c Iipc ce p(cIe|nc e e:Iccc fi:ic|ccicc. cepencc ce
p(cIec:e: e pepIicc:e: {/IF cepencenIe: cu incepencenIe:).
Exi:Iem 3 Iipc: ce meccni:mc: {2 nc ciIc:c| e c (e:IcnIe nc: |i:c::cmc:)
1. /: p(cIe|nc: inI(cce|u|c(e: ce meic-vicc cu(Ic e cnc(mci: :cc cec(ccccc: nc
ciIc:c|. cpc: icenIificcccc e fixcccc c um c|iccppIicc. c uLicuiIinc. :encc c
p(cIec|i:e ccIc|i:ccc pc( um ccmp|exc p(cIec:e 2S /IF-cepencenIe:
2. /: p(cIe|nc: cc memL(cnc. exI(cce|u|c(e: cu ce meic-vicc |cncc :cc
cec(ccccc: nc: |i:c::cmc: pc( ccIep:inc: {/IF-incepencenIe:) cpc:
enccciIc:e.
3. / cuI(c mccc|iccce ce cec(cccccc p(cIeicc exI(c-|i:c::cmc| envc|ve enzimc:
Cc
2+
-cepencenIe:.
A degrodoo dos protenos no s|stemo d|gest|vo:
E:Icmccc: c hc(mcnc cc:I(inc e:Iimu|c c :ec(eccc ce ccicc c|cnicucc {cnIi:pIicc e
ccenIe ce:ncIu(cnIe) e pep:inccnic {ccIivc zimccnic e c pep:inc hic(c|i:c c:
|icccce: pepIicicc:).
Fcnc(ec:: e:Iimu|ccc pe|c hc(mcnc :ec(eIinc c p(ccuzi( Licc(LcncIc pc(c
neuI(c|izc( HC|. / hc(mcnc cc|eci:Iccinc e:Iimu|c c :ec(eccc ce umc :(ie ce
enzimc:: I(ip:inccnic {fc(mc ccIivc cc I(ip:inc. ccIivccc pc( enIe(cpepIicc:e)
cuimcI(ip:inccnic e p(ccc(LcxipepIicc:e: {ccIivccc: pe|c I(ip:inc em cuimcI(ip:inc e
cc(LcxipepIicc:e:): F(ce|c:Ic:e p(ce|c:Ic:e.
lnIe:Iinc ce|cccc: cminipepIicc:e: {icenIificcm e c|ivcm c pc(Ii( cc c(upc cminc)

Am|noc|dos
Or|gens: Dest|nos:
/|imenIcccc 8ic::|nIe:e p(cIeicc
Dec(cccccc p(cIeicc /mincccc ce ceIcccicc:
De(ivccc: ncc-p(cIeicc: Cxiccccc meIcLc|icc cc e:cue|eIc cc(Lcnccc
{cc ceIccnicc: e c|icccnicc:)
Fecene(cccc cc c|icc:e
47

Degrodoo de om|noc|dos
Ce(cc ce 757 cc: cmincccicc: p(cvenienIe: cc cec(cccccc p(cIeicc :cc
(euIi|izccc: nc :|nIe:e p(cIeicc:
C: (e:IcnIe: pe(cem c :eu c(upc cminc { u(eic) pc(c fc(mc( u-ceIcccicc:. c:
e:cue|eIc: ce cc(Lcnc cc: cmincccicc:. C: u-ceIcccicc: pccem :e( cxicccc: c
CC2 e H2C cu fc(nece( uniccce: ce 3 c 4 C cue pccem :e( ccnve(Iicc: em c|icc:e
pe|c c|iccneccne:e:
/ cec(cccccc cxiccIivc ce cmincccicc: pcce ccc((e( em 3 cife(enIe:
ci(cun:Icncic: meIcLc|icc::
1. Du(cnIe c :|nIe:e e cec(cccccc nc(mc| ce p(cIe|nc: ce|u|c(e:: c|cun:
cmincccicc: |iLe(Iccc: cu(cnIe c cueL(c ce p(cIe|nc: pccem :e( cxicccc: :e
ncc fc(em nece::c(ic: pc(c ncvc: :|nIe:e: p(cIeicc:.
2. Cucncc umc cieIc (icc em p(cIe|nc: e c cucnIiccce ce cmincccicc:
excece c nece::c(ic pc(c :|nIe:e c exce::c ccIcLc|izccc: c: cmincccicc:
ncc pccem :e( c(mczenccc:.
3. Du(cnIe c jejum cu nc: c|ccete: ne|||tu:. cucncc c: cc(Lchic(cIc: e:Icc
inci:pcn|vei: cu c :e( incc((ecIcmenIe uIi|izccc:. c: p(cIe|nc: ce|u|c(e: :cc
uIi|izccc: pc(c c cLIenccc ce ene(cic.

Dest|nos metobo||cos dos grupos om|no:
C exce::c pcce :e( exc(eIccc ci(ecIcmenIe cu :e( ccnve(Iicc em u(eic cu ccicc
(icc pc(c exc(eccc. cepencencc cc c(ccni:mc. C exce::c ce cmcnic ce(ccc
em Iecicc: exI(chepcIicc: vicjc cI cc f|cccc. pc(c :e( ccnve(Iicc nc fc(mc em
cue exc(eIccc.

1. Irons|ernc|o dos grupos om|no
/::im cue c: cmincccicc: checcm cc f|cccc. ccc((e c (emcccc cc: c(upc:
cminc. p(cmcvicc pe|c: cmincI(cn:fe(c:e: cu I(cn:cminc:e:.
C c(upc cminc I(cn:fe(icc pc(c c cc(Lcnc u cc ceIcc|uIc(cIc. |iLe(Icncc c u-
ceIcccicc cnc|ccc cc cmincccicc.
C efeiIc cc: (ecccce: ce I(cn:cmincccc junIc( c: c(upc: cminc ce cife(enIe:
cmincccicc: nc fc(mc ce L-c|uIcmcIc.
C c|uIcmcIc funcicnc ccmc cccc( ce c(upc cminc pc(c c: vic: Lic::inIIicc: cu
vic: ce exc(eccc cue |evcm c e|imincccc ce p(ccuIc: niI(ccenccc:.
C esque|eto corbonodo fc(mccc uIi|izccc em inIe(mecic(ic: meIcLc|icc: {cceIi|-
Cc/. pi(uvcIc. inIe(mecic(ic: cc cic|c ce K(eL:) ccm pcIencic| ce fc(mcccc ce
ccicc: cc(cc:. cc(pc: ceIcnicc: e c|icc:e.

/mincI(cn:fe(c:e
/: c|u|c: ccnIm um n ce cife(enIe: cmincI(cn:fe(c:e:.
MuiIc: :cc e:pec|ficc: pc(c c u-ceIcc|uIc(cIc ccmc c(upc ccepIc(. mc: cife(enIe nc
e:pecificiccce pc(c c: L-cmincccicc:.
Iccc: c: cmincI(c:nfe(c:e: Im c fc:fcIc pi(iccxc| ccmc c(upc p(c:IIicc {ce(ivccc
cc viIcminc 8).
Fc:fcIc pi(iccxc| {FLF)
u-cetog|utoroto + L-om|nooc|dos L-g|utomoto + u-cetoc|do
cmincI(cn:fe(c:e
48

Fc(mc ccIivc cc viIcminc 8
Fc(Iicipc em cive(:c: (ecccce: impc(IcnIe: cc: cmincccicc: - I(cn:cmincccc.
ce:cc(Lcxi|cccc e mccificcccc cc ccceic |cIe(c| ccm pe(mcnncic cc c(upc u-
cminccc(Lcxi|icc. e|imincccc ce H2C cu H2S: (ccemizcccc.
Nc: 3 p(imei(c: (ecccce: (efe(icc:. fc(mc-:e c me:mc ccmpc:Ic inIe(mcic {umc
Lc:e ce Schiff) enI(e c ccenzimc e c :uL:I(cIc cmincccicc.

2. kemoo dos grupos om|no

2.1 Desom|noo ox|donte do g|utomoto
Nc: hepcIcciIc:. c c|uIcmcIc I(cn:pc(Iccc cc ciIc:c| pc(c c miIcccnc(ic. cnce
:cf(e ce:cmincccc cxiccIivc. ccIc|i:ccc pe|c L-c|uIcmcIc ce:ic(ccenc:e.

G|utomoto + NAD(F)
+
u-cetog|utoroto + NH4
+
+ NAD(F)H + H
+

L-c|uIcmcIc ce:ic(ccenc:e

/ ccmLinccc ccccc ce cmincI(cn:fe(c:e: e c|uIcmcIc ce:ic(ccenc:e (efe(icc
ccmc I(cn:ce:cmincccc.
C u-ceIcc|uIc(cIc fc(mccc pcce :e( uIi|izccc nc cic|c ce K(eL: e pc(c :|nIe:e ce
c|icc:e.
/ inIe(ccnve(:cc enI(e c c|uIcmcIc e c u-ceIcc|uIc(cIc ccn:IiIui um impc(IcnIe
pcnIc ce ccnIccIc enI(e c: meIcLc|i:mc: c|icicicc e p(cIeicc.

G|uIcmcIc ce:ic(ccenc:e:
Cpe(c numc impc(IcnIe inIe(:ecccc cc: meIcLc|i:mc: ce cc(Lcnc e czcIc.
/ :uc ccIiviccce inf|uenciccc pc( umc ccmp|exc (ece ce mccu|ccc(e:
c|c:I(icc:
Mccu|ccc(e: pc:iIivc:: GDF. /DF
Mccu|ccc(e: neccIivc:: GIF. /IF

2.2 Desom|noo d|recto de o|guns om|noc|dos

C: c(upc: cminc cc :e(inc. I(ecninc e ci:Ie|nc pccem :e( ccnve(Iicc:
ci(ecIcmenIe em cmcn|ccc. vi:Ic cue e:Ie: cmincccicc: ccnIm c(upc: hic(cxi|c
cu :u|fic(i|c nc: ccceic: |cIe(ci:.
Nc ce:cmincccc cc L-:e(inc {:e(inc-ce:ic(cIc:e) e cc L-ci:Ie|nc {ci:Ieinc-
ce::u|fic(c:e) fc(mc-:e pi(uvcIc e cmcn|ccc.
C c(upc p(c:IIicc ce:Ic: enzimc: c fc:fcIc ce pi(iccxc|.
Nc cc:c cc ce:cmincccc cc I(ecninc fc(mc-:e cmcn|ccc e u-ceIcLuI|(icc.

2.3 Desom|noo por om|noc|dos-ox|doses:

F(cce::c i((eve(:|ve|
1 Ccc((e umc ce:ic(ccencccc cc cmincccicc c imincccicc. em cue c (eccccc
ccIc|i:ccc pc( umc f|cvcp(cIe|nc. e c: 2 H (eIi(ccc: cc :uL:I(cIc vcc (ecuzi( c F/D
cu FMN.
Depci: c imincccicc hic(c|i:c-:e em u-ceIcccicc e cmcn|ccc.
/ ccenzimc (ecuzicc ce(c|menIe (ecxiccce pe|c cxicnic mc|ecu|c( c H2C.
49

Nc enIcnIc cuI(c: cceiIccc(e: ce e|ecI(ce: pccem pe(miIi( c (ecxiccccc cc
F/DH2.
/: cmincccicc:-cxicc:e: :cc enzimc: e:Ie(ece:pec|ficc:. e cpenc: c:
cmincccicc-cxicc:e: L ccIc|i:cm c ce:cmincccc cxiccnIe cc: cmincccicc:
p(cIeicc:.

3. Formoo e tronsporte de omonoco

/ cmcnic Icxicc pc(c c: Iecicc: cnimci:. e c exce::c ccnve(Iicc num
ccmpc:Ic ncc Icxicc cnIe: ce :e( I(cn:pc(Iccc cc: Iecicc: exI(chepcIicc: pc(c c
f|cccc cu (in:. pe|c ci(cu|cccc :cncu|nec.
/ cmcnic |iv(e ccmLinc-:e ccm c c|uIcmcIc. c(icincncc g|utom|no pe|c ccccc
cc c|uIcminc :inIeIc:e. em 2 eIcpc::

L-g|utomoto + AIF v-g|utom|| |os|oto + NH4
+
+ ADF + F| g|utom|no
c|uIcminc :inIeIc:e

/ c|uIcminc em exce::c em (e|cccc c nece::c(ic pc(c Lic::|nIe:e I(cn:pc(Iccc
nc :cncue pc(c c f|cccc e (in: pc(c p(cce::cmenIc.
/ cmcnic cpenc: |iLe(Iccc nc: miIcccnc(ic: cc f|cccc/(in:. cnce c enzimc
c|uIcminc:e ccnve(Ie c c|uIcminc c c|uIcmcIc e NH4
+
.
Nc f|cccc e:Ic (eccccc fc(nece cuI(c fcnIe ce cmcnic cue pcce :e( uIi|izccc
pc(c c :|nIe:e ce u(eic.
C c|uIcmcIc pcce :e( cepci: p(cce::ccc pe|c c|uIcmcIc ce:ic(ccenc:e.
|iLe(Icncc mci: cmcnic e p(ccuzincc e:cue|eIc: ce cc(Lcnc pc(c cLIenccc ce
ene(cic.

L-g|utom|no + H2O L-g|utomoto + NH4
+

G|uIcminc:e

/ o|on|no IcmLm ce:empenhc um pcpe| e:pecic| nc I(cn:pc(Ie ce c(upc cminc
pc(c c f|cccc numc fc(mc ncc Icxicc. pe|c cic|c c|icc:e-c|cninc.
C c|uIcmcIc IcmLm pcce I(cn:fe(i( c :eu c(upc u-cminc cc pi(uvcIc. um
p(ccuIc p(cnIcmenIe ci:pcn|ve| cc c|icc:e cc m:cu|c. pe|c ccccc cc c|cninc
cmincI(cn:fe(c:e.
/ c|cninc fc(mccc pc::c pc(c c :cncue e vicjc cI cc f|cccc.
Nc ciIc:c| cc: hepcIcciIc:. c c|cninc cmincI(cn:fe(c:e I(cn:fe(e c c(upc cminc cc
c|cninc pc(c c u-ceIcc|uIc(cIc. fc(mcncc pi(uvcIc e c|uIcmcIc.
C c|uIcmcIc enI(c nc miIcccnc(ic cnce c (eccccc cc c|uIcmcIc ce:ic(ccenc:e
|iLe(Ic NH4
+
. cu pcce :cf(e( I(cn:cmincccc ccm c cxc|ccceIcIc pc(c fc(mc(
c:pc(IcIc. cuI(c cccc( ce czcIc nc :|nIe:e cc u(eic.
/ uIi|izcccc cc c|cninc pc(c I(cn:pc(Ic( cmcnic cc m:cu|c e:cue|Iicc pc(c c
f|cccc cuI(c exemp|c cc eccncmic inI(|n:ecc cc: c(ccni:mc: vivc:.

/ ccnI(ccccc vicc(c:c cc: m:cu|c:. efecIuccc em ccncicce: cnce(cLic:. p(ccuz
pi(uvcIc e |ccIcIc {c|icc|i:e) e cmcnic {c|ivccem ce p(cIe|nc:). E:Ie: p(ccuIc:
50

(ec(e::cm cc f|cccc. cnce c pi(uvcIc e c |ccIcIc :cc incc(pc(ccc: nc c|icc:e. cue
(ec(e::c cc: m:cu|c:. e c cmcnic ccnve(Iicc em u(eic pc(c exc(eccc.
C c|c|o do g||cose-o|on|no. em ccnjunIc ccm c c|c|o de cor|. (ec|izc e:Ic I(cn:ccccc.
C fc(cc ene(cIicc cc c|iccneccne:e impc:Ic enIcc cc f|cccc :encc Iccc c /IF
ci:pcn|ve| nc m:cu|c uIi|izccc pc(c c ccnI(ccccc mu:cu|c(.

4. Excreo do ozoto e c|c|o do ure|o

Ccc((e em 2 ccmpc(IimenIc:
3 cmincccicc:: c(niIinc: ciI(u|inc: c(cininc
lnIe(mecic(ic: cc cic|c ce K(eL:: cxc|ccceIcIc. fumc(cIc. ceIcc|uIc(cIc.
Gc:Ic ce 3 /IF pc( 2 NH4
+
incc(pc(ccc:

Nc: c(ccni:mc u(ecI|icc: c cmcnic cepc:iIccc nc: miIcccnc(ic: cc: hepcIcciIc:
ccnve(Iicc em u(eic nc cic|c cc u(eic.
/ u(eic pc::c pc(c c cc((enIe :cncu|nec e cepci: pc(c c (im e exc(eIccc nc
u(inc.
C cic|c cc u(eic ccmecc nc miIcccnc(ic cc f|cccc. mc: c: 3 pc::c: :uL:ecuenIe:
ccc((em nc ciIc:c|: c cic|c ccc((e em 2 ccmpc(IimenIc:.
C 1 c(upc cminc cue enI(c nc cic|c cc u(eic ce(ivccc cc cmcnic nc mcI(iz
miIcccnc(ic|.
C f|cccc IcmLm (eceLe cmcnic. cI(cv: cc veic pc(Ic cc inIe:Iinc.
C NH4
+
. cuc|cue( cue :ejc c fcnIe. imecicIcmenIe u:ccc em ccnjunIc ccm CC2
{ccmc HCC3
-
) p(ccuzicc nc (e:pi(cccc miIcccnc(ic| pc(c fc(mc( cc(Lcmci| fc:fcIc
nc mcI(iz. E:Ic (eccccc /IF cepencenIe ccIc|izccc pe|c cc(Lcmci| fc:fcIc
:inIeIc:e l. umc enzimc (ecu|ccc(c.

NH4
+
+ HCO3
-
+ 2AIF Corbomo|| |os|oto + 2ADF
cc(Lcmci| fc:fcIc :inIeIc:e

C cc(Lcnci| fc:fcIc enI(c ccc(c nc cic|c cc u(eic. cue Iem 4 pc::c: enzimcIicc::

1. C cc(Lcnci| fc:fcIc ccc c :eu c(upc cc(Lcmci| c c(niIinc pc(c fc(mc( ciI(u|inc.
ccm |iLe(Icccc ce Fi. / c(niIinc ce:empenhc um pcpe| :eme|hcnIe cc cc
cxc|ccceIcIc nc Cic|c ce K(eL:. cceiIcncc mcIe(ic| em cccc vc|Ic cc cic|c. /
(eccccc ccIc|izccc pe|c c(niIinc I(cn:cc(Lcmci|c:e e c ciI(u|inc (e:u|IcnIe
pc::c cc miIcccnc(ic cc ciIc:c|

Corbomo|| |os|oto + orn|t|no c|tru||no
c(niIinc I(cn:cc(Lcmci|c:e

2. C 2 c(upc cminc inI(ccuzicc pe|c c:pc(IcIc pc( umc ccncen:cccc enI(e c
c(upc cminc cc c:pc(IcIc e c c(upc cc(Lcni| cc ciI(u|inc. fc(mcncc
c(cininc:uccincIc. E:Ic (eccccc ccIc|izccc pe|c c(cininc:uccincIc :inIeIc:e.
(ecue( /IF e Iem c ciI(u|i|-/MF ccmc inIe(mecic(ic

C|tru||no + AIF c|tru|||-AMF + FF| + Asportoto org|n|nosucc|noto + AMF
/(cininc:uccincIc :inIeIc:e

3. / c(cininc:uccincIc enIcc c|ivccc (eve(:|ve|menIe pe|c c(cininc:uccincIc
|ic:e. c(icincnIc c(cininc e fumc(cIc {cue enI(c pc:Ie(ic(menIe nc miIcccnc(ic
pc(c :e(vi( ccmc inIe(mecic(ic cc C/C)
51

Arg|n|nosucc|noto Arg|n|no + Fumoroto
/(cininc:uccincIc |ic:e

4. Fc( fim. c c(cinc:e c|ivc c c(cininc pc(c fc(mc( u(eic e c(niIinc. / c(niIinc
I(cn:pc(Iccc pc(c c miIcccnc(ic pc(c inicic( cuI(c vc|Ic cc cic|c cc u(eic.

Arg|n|no + H2O bre|o + orn|t|no
/(cinc:e

Enzimc: cc cic|c cc u(eic:
Scc :inIeIizccc: em e|evccc: n|vei: ce jejum cu em cieIc: ccm e|evccc Iec(
p(cIeicc.
Numc pecuenc e:cc|c c (ecu|cccc c|c:I(icc ce pe|c: menc: um enzimc-chcve
cju:Ic c f|uxc cI(cv: cc cic|c cc u(eic.
/ 1 enzimc cc cic|c c|c:I(icc. ccIivccc pe|c N-cceIi|c|uIcmcIc. cue
:inIeIizccc cc cceIi|-Cc/ e c|uIcmcIc pe|c N-cceIi|c|uIcmcIc :inIeIc:e {e cue
ccIivccc pe|c c(cininc)

Se c cic|c cc u(eic fc( vi:Ic i:c|cccmenIe c :|nIe:e ce umc mc|cu|c ce u(eic
(ecue( 4 c(upc: fc:fcIc ce c|Ic ene(cic {2/IF pc(c fcze( cc(Lcmci| fc:fcIc + 1/IF pc(c
:|nIe:e ce c(cininc:uccincIc)

2NH4
+
+ HCC3
-
+ 3/IF
4-
+ H2C u(eic + 2/DF
3-
+ 4Fi + /MF
2-
+ 5H
+

CcnIucc. c cic|c cc u(eic IcmLm p(cvccc c ccnve(:cc ce czc|ccceIcIc em
fumc(cIc {vic c:pc(IcIc) e c (ecene(cccc cc cxc|ccceIcIc. cue p(ccuz N/DH nc
(eccccc cc mc|cIc ce:ic(ccenc:e

ke|oes entre o c|c|o do ure|o e o c|c|o de Krebs
C fumc(cIc p(ccuzicc nc (eccccc ce c(cininc:uccincIc |ic:e. inIe(mecic(ic cc
cic|c ce K(eL:. pcce :e( ccnve(Iicc c mc|cIc e c cxc|ccceIcIc nc ciIc:c|. e e:Ie:
inIe(mecic(ic: pccem :e( meIcLc|izccc: nc ciIc:c| cu I(cn:pc(Iccc: pc(c c
miIcccnc(ic. :encc uIi|izccc: nc cic|c ce K(eL:.
C c:pc(IcIc fc(mccc nc miIcccnc(ic pc( I(cn:cmincccc enI(e cxc|ccceIcIc e
c|uIcmcIc pcce :e( I(cn:pc(Iccc pc(c c ciIc:c|. cnce :e(ve ccmc cccc( ce czcIc
nc cic|c cc u(eic nc (eccccc ccIc|izccc pe|c c(cininc:uccincIc :inIeIc:e.
52

E:Ic: (ecccce: ccn:IiIuem c ce:vic c:pc(IcIc-c(cininc:uccincIc. cue fc(nece
|icccce: enI(e c: vic: :epc(ccc: pe|c: cuci: c: c(upc: cminc e c: e:cue|eIc: ce
cc(Lcnc :cc p(cce::ccc:.

G|utomoto + oxo|oocetoto osportoto + u-cetog|utoroto
I(cn:cminc:e

H|peromo|n|m|o:
lnccnvenienIe: cc ccumu|cccc cc icc cmcnic nc: c|u|c: e em ci(cu|cccc.
/ cu:ncic ccncniIc {pc(cic| cu IcIc|) cc: enzimc: cc cic|c cc u(eic ccu:c ce
mc(Ie cu ce cI(c:c c(cve cc ce:envc|vimenIc cc(pc(c| e menIc|.
/ ccumu|cccc ce NH4
+
nc Iecicc ne(vc:c cumenIc c uIi|izcccc cc u-ceIcc|uIc(cIc
pc(c fc(mcccc cc c|uIcmcIc pe|c ce:cc(ccenc:e c|uIcmicc. e ccnve(:cc ce:Ie
em c|uIcminc.

G|utom|no||se

/ c|uIcminc |iLe(Iccc cc m:cu|c ccnve(Iicc em c|cninc pe|c epiI|ic inIe:Iinc|.
/ c|uIcminc pc(cic|menIe cxicccc nc: enIe(cciIc: pc(c fc(nece( ene(cic e
mc|cu|c: pe(cu(:c(c: cc :|nIe:e ce pi(imicinc: e pu(inc:. ccm c: cc(Lcnc: e NH4
+

(e:IcnIe: |iLe(Iccc: nc cc((enIe :cncu|nec nc fc(mc ce c|cninc.

G|utom|no mo|oto p|ruvoto o|on|no

Umc impc(IcnIe vic exi:IenIe nc epiI|ic inIe:Iinc| pc(c c ccnve(:cc cc c|uIcminc
em ciI(u|inc.
Umc cc: enzimc: {c|uIcmcIc (ecuIc:e /IF-cepencenIe) nece::c(ic: pc(c exccIc
ccnve(:cc exi:Ie cpenc: nc: enIe(cciIc:.
/ ciI(u|inc p(ccuzicc nc inIe:Iinc meIcLc|izccc pe|c (im em c(cininc. cue pcce
:e( ccnve(Iicc em c(ecIinc cu |iLe(Iccc nc :cncue.
C f|cccc uIi|izc c c(cininc cc :cncue pc(c ce(c( c(niIinc. cue expcnce c
ccpcciccce cc cic|c cc u(eic em pe(|ccc: ce c(cnce uIi|izcccc p(cIeicc.
E:Ic vic ce c(cnce impc(Icncic pc(c c f|cccc. vi:Ic cue e:Ie ccnIm um :i:Iemc
enzimcIicc cue ccnve(Ie c(niIinc em c|uIcmcIc i((eve(:ive|menIe.
/ cep|ecccc cc c(niIinc pc( e:Ic: (ecccce: iniLe c :|nIe:e ce u(eic nc f|cccc.
e:Icncc c (efc(necimenIc ce c(niIinc cepencenIe cc fcnIe ce c(cininc nc
:cncue.
53

8
88.
..8
88|
||o
oos
sss
ss|
||n
nnt
tte
ees
sse
ee F
FFr
rro
oot
tte
ee|
||c
cco
oo

1. 8|ossntese de c|dos gordos

Loco| de sntese: ciIc:c| cc mcic( pc(Ie cc: Iecicc:. e:pecic|menIe. nc ficccc. (im.
Iecicc ccipc:c. c(eL(c. pu|mcc e c|cncu|c mcmc(ic.
Co|octores nece::c(ic: :cc:
N/DFH
/IF.
Mn
2+
8icIinc
HCC3
-
{ccmc fcnIe ce CC2)
$ubstroto gero|: cceIi|-Cc/
Froduto ||no|: pc|miIcIc {C1).
A b|ossntese de c|dos gordos:
Ccc((e cucncc c cieIc (icc cc(Lchic(cIc: e p(cIe|nc:
Ccc((e ccm n|vei: ciIcp|c:mcIicc: e|evccc: ce ciI(cIc e N/DFH
Ccc((e nc ciIcp|c:mc (ecue(encc c ccmp|exc mu|IienzimcIicc cc :inIc:e cc:
ccicc: cc(cc:.

Dife(encc: enI(e c -Cxiccccc e c 8ic::|nIe:e ce ccicc: cc(cc: e p(incipci: c:pecIc:
cc Lic::|nIe:e:
Ccc((e nc ciIcp|c:mc. e ncc nc miIcccnc(ic:
U:c NADFH ccmc fcnIe ce e|ecI(ce::
C I(cn:pc(Iccc( ce c(upc: cci| c ACF {/cy| Cc((ie( F(cIein). e ncc c ccenzimc /
C c(upc ccIivccc( ncc c c(upc Iic| cc Cc/.
/: enzimc: e:Icc c(ccnizccc: num :i:Iemc mu|IienzimcIicc {s|ntose dos c|dos
gordos)
/ ccceic ce ccicc: cc(cc: c|cncccc pe|c cciccc :ecuencic| ce uniccce: ce 2
cc(Lcnc: p(cvenienIe: cc ocet||-CoA. C cccc( ccIivccc ce::c: uniccce: c
mo|on||-ACF e c (eccccc ce c|cnccmenIc fcvc(ecicc pe|c |iLe(Icccc ce CC2.
C c|cnccmenIc pe|c ccmp|exc cc :inIc:e Ie(minc ccm c fc(mcccc cc po|m|toto.
C: c|cnccmenIc: pc:Ie(ic(e: e fc(mcccc ce cup|c: |icccce: :cc ccIc|i:ccc: pc(
cuI(c: :i:Iemc: mu|IienzimcIicc:.

Comp|exo enz|mt|co - s|ntose de c|dos gordos
CcnIm 2 :uLuniccce: e:Icncc cccc uniccce {mcncme(c) ce:ccL(ccc em 3
ccm|nic: cue. nc :eu ccnjunIc. cp(e:enIcm 7 cenI(c: ccIc||Iicc:.
Dcm|nic 1: uniccce ce |iccccc cc :uL:I(cIc e ce ccncen:cccc.
/ceIi|-Cc/-/CF I(cn:cceIi|c:e {cu cceIi|I(cn:fe(c:e)
-ceIccci|-/CF :inIc:e
mc|cni|-Cc/-/CF I(cn:fe(c:e
Dcm|nic 2: uniccce ce (ecuccc
/CF
ceIccci|-/CF (ecuIc:e
-hic(cxicci|c-/CF ce:ic(cIc:e
enci|-/CF (ecuIc:e
Dcm|nic 3: c uniccce ce |iLe(Icccc cc ccicc pc|m|Iicc
Iice:Ie(c:e

/: :uLuniccce: e:Icc unicc: ce umc fc(mc cnIi-pc(c|e|c pe|c: c(upc: Iici:c
/ c::ccicccc ce cife(enIe: enzimc: ce fc(mc ccvc|enIe. pc(c ccn:IiIui( umc nicc
p(cIe|nc mu|Iifuncicnc|. Iem vcnIccen: ce pe(miIi( umc ccc(cencccc mci: (cpicc.
cumenIc cc eficincic e ciminuiccc ce inIe(fe(ncic: ce cuI(c: (ecccce:
ccmpeIiIivc:.

54

C ACF {/cy| Cc((ie( F(cIein) umc pecuenc p(cIe|nc cue ccnIm c c(upc
p(c:IIicc 4-fc:fcpcnIeIeinc. / hic(c|i:e cc |iccccc Iic:Ie( cue une c /CF cc c(upc
cci| muiIc exe(ccnicc. e c ene(cic |iLe(Iccc ccnI(iLui pc(c cue c 1 (eccccc nc
Lic::|nIe:e cc: ccicc: cc(cc: :ejc Ie(mccincmiccmenIe fcvc(cve|.

1. $ntese do mo|on||-CoA
/ :|nIe:e ce ccicc: cc(cc: feiIc c pc(Ii( ce cceIi|-Cc/. Nc enIcnIc. c p(cce::c
enccene(cIicc. pe|c cue c cceIi|-Cc/ ceve :e( p(evicmenIe ccIivccc. E:Ie
pc(IcnIc cc(Lcxi|ccc pe|c cceIi|-Cc/ cc(Lcxi|c:e. umc enzimc cue Ic| ccmc c: cuI(c:
cc(Lcxi|c:e: pc::ui b|ot|no

Acet||-CoA + HCO3
-
+ AIF Mo|on||-CoA + H2O + ADF + F|
/ceIi|-Cc/ cc(Lcxi|c:e

C c(upc cc(Lcxi|c. ce(ivccc ce Licc(LcncIc. I(cn:pc(Iccc pc(c c LicIinc numc
(eccccc /IF-cepencenIe.
C c(upc LicIinc :e(ve ccmc um I(cn:pc(Iccc( Iempc(c(ic ce CC2. I(cn:fe(incc-c
cepci: cc cceIi|-Cc/ nc 2 eIcpc pc(c c(icinc( mc|cni|-Cc/.
/ cceIi|-Cc/ cc(Lcxi|c:e um pcnIc-chcve nc (eccccc c|cLc| ce :|nIe:e ce
ccicc: cc(cc: e pcce :e( i:c|ccc num e:Iccc mcncm(icc. inccIivc. C:
p(cIcme(c: cc(eccm pc(c fc(mc( pc||me(c: enzimcIiccmenIe ccIivc: pc( cciccc
ce ciI(cIc. {Fc(mc ccIivc: pc|ime(c: fc(mc inccIivc: mcncme(c)
C pc|miIci|-Cc/ iniLe c enzimc ccIivc.
CcnI(c|ccc (ecu|cccc. cI(cv: ce fc:fc(i|cccc c umc fc(mc inccIivc pc( umc
p(cIe|nc-cinc:e. e ce ce:fc:fc(i|cccc c umc fc(mc ccIivc {pc( umc fc:fcIc:e).

2. $ntese propr|omente d|to
C: c(upc: cci| :cIu(ccc: p(ccuzicc: Ic(ncm-:e c :uL:I(cIc nc ccncen:cccc
:uL:ecuenIe ccm um c(upc mc|cni| ccIivccc.
Em cccc pc::ccem pe|c cic|c c ccceic cci| cumenIc 2 cc(Lcnc:.
Cucncc c ccceic cIince 1 cc(Lcnc:. c p(ccuIc {pc|miIcIc) ceixc c cic|c.
55

/nIe: cc: (ecccce: ce ccncen:cccc cue ccn:I(cem c ccceic ce ccicc: cc(cc:
ccmecc(em. c: 2 c(upc: Iic| nc ccmp|exc enzimcIicc :cc cc((ecccc: ccm c:
c(upc: cci| cc((ecIc:.

1. C c(upc cci| cc cceIi|-Cc/ I(cn:fe(icc pe|c /ceIi|-Cc/-/CF I(cn:cceIi|c:e
pc(c c p(cIe|nc I(cn:pc(Iccc(c ce /ci| cu /CF fc(mcncc cceIi|-/CF.

Acet||-CoA + ACF-$H Acet|| ACF + CoA
/ceIi|-Cc/-/CF I(cn:cceIi|c:e

2. C c(upc cci| cc /ceIi|-Cc/ cepci: I(cn:fe(icc pc(c um c(upc Iic| ce umc
ci:Ie|nc cc cenI(c ccIivc cc enzimc ccncen:ccc(c cu -ceIccci|-/CF :inIc:e.
ficcncc |iv(e c |ucc( |iv(e nc /CF.

Acet|| ACF Acet|| s|ntose + ACF-$H
I(cn:cci|c:e + :inIc:e SH

3. C c(upc mc|cni| I(cn:fe(icc pc(c c c(upc SH cc /CF pe|c mc|cni|-Cc/-/CF
I(cn:fe(c:e.

Mo|on|| CoA + ACF-$H Mo|on|| ACF + CoA
Mc|cni| Cc/ I(cn:cci|c:e

4. Ccncen:cccc cc: c(upc: cceIi|-ci:Ieinc e mc|cni|-/CF ccIivccc: pc(c fc(mc(
cceIccceIi|-/CF. ccm fc(mcccc ce umc mc|cu|c ce CC2. E:Ic (eccccc
ccIc|izccc pe|c -ceIccci|-/CF :inIc:e

Acet|| s|ntose + Mo|on|| ACF Acetoocet||o ACF + CO2
-CeIccci|c /CF :inIc:e

/ ce:cc(Lcxi|cccc {cue ccc((e pc(cue um cc: inIe(mecic(ic: c mc|cni|-Cc/. e ncc
cuI(c cceIi|-/CF) fcvc(ece c (eccccc ci(ecIc cevicc c um cLcixcmenIc :uL:Icncic|
cc ene(cic |iv(e.

5. C cceIccceIi|-/CF fc(mccc ccc(c (ecuzicc pc(c fc(mc( D--hic(cxiLuIi(i|-
/CF. C ccIc|izccc( ceIccci|-/CF (ecuIc:e

Acetoocet||o ACF + NADFH NADF + -3 h|drox|but|r|| ACF
-CeIccci|c /CF (ecuIc:e

. (eIi(ccc ccuc cc D--hic(cxiLuIi(i|-/CF. c(icincncc umc |iccccc cup|c nc
p(ccuIc. I(cn:-A
2
-LuIenci|-/CF -hic(cxicci|c-/CF ce:ic(cIc:e.

-3 h|drox|but|r|| ACF Croton||o ACF + H2O
-hic(cxicci|c-/CF ce:ic(cIc:e

7. / |iccccc cup|c cc p(ccuIc cnIe(ic( (ecuzicc pe|c N/DFH pc(c fc(mc( LuIi(i|-
/CF. pe|c ccccc cc enci|-/CF (ecuIc:e.

Croton||o ACF + NADFH NADF + but|r||-ACF
enci|-/CF (ecuIc:e

56

8. / ccceic ce LuIi(i| I(cn:fe(icc cc /CF pc(c c enzimc ccncen:ccc(c.
ceixcncc |ucc( nc /CF pc(c c uniccce ce Mc|cni|-Cc/ :ecuinIe.
. C LuIi(i|c |icc(-:e-c ccc(c c cci: cc(Lcnc: cc ncvc uniccce ce mc|cni|-/CF
pc(c fc(mc( umc ccceic ce cc(Lcnc:.

7 Cic|c: ce ccncen:cccc e (ecuccc c(icincm c c(upc pc|miIci| ce 1 cc(Lcnc:.
cincc |icccc cc /CF. Umc Iic-e:Ie(c:e I(cn:fc(mc c pc|miIci|-Cc/ em pc|miIcIc.
De:Iinc: cc pc|miIci|-Cc/
S|nIe:e ce I(icci|c|ice(ci:
S|nIe:e ce fc:fc|ipicc:
S|nIe:e cc: ccicc: cc(cc: cc ccceic |cncc e in:cIu(ccc:

keoco g|obo| do po|m|toto
Acet||-CoA + 7 Mo|on||-CoA + 14 NADFH + 7 H
+
c|do po|mt|co + 7 CO2 +
14 NADF
+
+ 8 CoA + H2O

/ Lic::|nIe:e ce ccicc: cc(cc:. ccmc c pc|miIcIc. (ecue( enIcc /ceIi|-Cc/ e c inpuI"
ce ene(cic cu|micc em 2 fc(mc::
AIF: (ecue(icc pc(c |icc( c CC2 c cceIi|-Cc/. c(icincncc mc|cni|-Cc/.
NADFH: (ecuz c: |icccce: cup|c::
F(cvenienIe cc:
Vic cc: fc:fcpenIc:e: {c|icc:e -F ce:ic(ccenc:e e fc:fcc|iccncIc ce:ic(ccenc:e)
Enzimc mc|icc {cxicc c mc|cIc c pi(uvcIc)
De:ic(ccenc:e i:cciI(icc N/DF cepencenIe
Nc: hepIcciIc: c (e|cccc {N/DFH]/{N/DFH
+
] nc ciIc:c| muiIc e|evccc. fc(necencc
um cmLienIe (ecuIc( pc(c c :|nIe:e ce ccicc: cc(cc: e cuI(c: Licmc|cu|c:.

Or|gens do Acet||-CoA e NADFH
Cuc:e Iccc: c: ocet||-CoA uIi|izccc nc :|nIe:e ce ccicc: cc(cc: fc(mccc nc
miIcccnc(ic pe|c cxiccccc cc p|ruvoto e cc ccIcLc|i:mc cc: esque|etos de
corbono cc: cmincccicc:.
/ memL(cnc miIcccnc(ic| inIe(nc impe(mecve| c cceIi|-Cc/. Iencc ce :e( c:
c(upc: cceIi| ecuivc|enIe: I(cn:pc(Iccc: cI(cv: cc memL(cnc pc( um "shutt|e"
inci(ecIc.
C cceIi|-Cc/ inI(cmiIcccnc(ic| (ecce ccm c oxo|oocetoto pc(c fc(mc( c|troto. nc
(eccccc cc cic|c ce K(eL: ccIc|izccc pe|c ciI(cIc :inIc:e.
C ciI(cIc pc::c pc(c c ciIc:c| pe|c tronsportodor de c|troto nc memL(cnc
miIcccnc(ic| inIe(nc.
Nc ciIc:c|. c c|ivccem cc ciI(cIc pe|c ciI(cIc |ic:e (ecene(c c cceIi|-Cc/ numc
(eccccc /IF-cepencenIe.
Ccmc c oxo|oocetoto ncc Iem um I(cn:pc(Iccc( cue c pe(miIc (ec(e::c( c mcI(iz
miIcccnc(ic|. (ecuzicc c mo|oto {pe|c mc|cIc ce:ic(ccenc:e) cue I(cn:pc(Iccc
nc tronsportodor mo|oto/u-cetog|utoroto.
Nc mcI(iz miIcccnc(ic| (ecxicccc c cxc|ccceIcIc.
E:Ie meccni:mc Iem cuI(c: vcnIccen: pc(cue CO2 e NADFH pc(c c :|nIe:e ce
ccicc pc|m|Iicc pccem :e( p(ccuzicc: c pc(Ii( cc ccicc cxc|cccIicc ciIc:c|icc.
C N/DH (ecuz c ccicc cxc|cccIicc c ccicc mc|icc. vic Mc|cIc ce:ic(ccenc:e
ciIc:c|icc. e c ccicc mc|icc . enIcc ce:cc(Lcxi|ccc pe|c enzimc Mc|icc N/DF-
cepencenIe. pc(c p(ccuzi( N/DFH. ccicc pi(vicc e CC2. /::im. c N/DFH
p(ccuzicc c pc(Ii( ce N/DH ce(ccc nc c|icc|i:e.
C cic|c ccmp|eIccc pe|c (eIc(nc cc ccicc pi(vicc c miIcccnc(ic. cnce pcce
:e( cc(Lcxi|ccc pc(c (ecene(c( c ccicc cxc|cccIicc.

57

A|ongomento dos c|dos gordos
Nc (eI|cu|c enccp|c:mcIicc
/ :ecuncic ce (ecccce: :eme|hcnIe c cue ccc((e nc ciIc:c|. ccm c
Mc|cni|-Cc/ ccmc fcnIe ce cuc: uniccce: ce cc(Lcnc e c N/DFH fc(necencc
c pcce( (ecuIc(.
C :uL:I(cIc p(efe(encic| ce c|cnccmenIc nc FE c c|do po|m|t|co p(ccuzicc
nc ciIc:c| cue fci ccIivccc c Fo|m|t||-CoA pe|c /ci|-Cc/ :inIeIc:e.
C: inIe(mecic(ic: nc: (ecccce: :uL:ecuenIe: :cc :Ie(e: ce Cc/. cc inv: ce
|icccc: c umc p(cIe|nc {enzimc: :epc(ccc:)
Nc mcic(ic cc: Iecicc: e::e :i:Iemc ccnve(Ie c ccicc pc|m|Iicc {C1) c c|do
ester|co {C18).
C c(eL(c pc::ui um cu mci: :i:Iemc: ce c|cnccmenIc ccicicnci:. cue
:inIeIizcm c|m cc ccicc e:Iec(icc. cuI(c: ccicc: ce ccceic |cncc {cI C24)
nece::c(ic: pc(c c: |ipicc: ce(eL(ci:. E:Ie: :i:Iemc: IcmLm u:cm Mc|cni|-Cc/

Nc miIcccnc(ic
C cceIi|-Cc/ c fcnIe ce uniccce: ce 2 cc(Lcnc: ccicicnccc:. e cmLc:.
N/DH e N/DFH. :e(vem ccmc ccenIe: (ecuIc(e:.
E:Ie :i:Iemc funcicnc cc ccnI(c(ic cc vic ce cxiccccc ce ccicc: cc(cc:.
excepIc cue c enci|-Cc/ (ecuIc:e N/DFH-cepencenIe {u|Iimc eIcpc cc
c|cnccmenIc) :uL:IiIui c /ci|-Cc/ ce:ic(ccenc:e F/D-cepencenIe {p(imei(c
eIcpc cc cxiccccc).
C p(cce::c Iem pcucc cfiniccce pc(c c ccicc pc|m|Iicc {C1). :e(vincc.
p(incipc|menIe. pc(c c e|cnccccc ce e:pcie: ce ccceic mci: cu(Ic.

kegu|oo do sntese de c|dos gordos

In||unc|o do estodo nutr|c|ono| do orgon|smo:
Ve|cciccce ciminui:
Ccncicce: ce cieIc (icc em cc(cu(c:
Cucncc hc ceficincic cc|c(icc
Cucncc hc (ecuccc ce in:u|inc.
Fe|cccc inve(:c enI(e c ccncenI(cccc ce ccicc: cc(cc: |iv(e: nc :c(c e c |ipccne:e
hepcIicc.
58

keoco ||m|tonte: ccIc|izccc pe|c /ceIi|-Cc/ cc(Lcxi|c:e
Fc(mc ccIivc: pc|ime(izccc ce:fc:fc(i|ccc {in:u|inc cepencenIe)
Fc(mc ccIivc: ce:pc|ime(izccc fc:fc(i|ccc {cepencenIe cc c|icccinc e epinef(inc)
/cIivccc(e:: n|vei: e|evccc: ce ciI(cIc
lniLicc(e:: n|vei: e|evccc: ce mc|cni|-Cc/ e ce pc|miIci|-Cc/

C ciI(cIc ce:empenhc um pcpe| cenI(c| nc cive(:cc cc meIcLc|i:mc ce|u|c( cc
cxiccccc ce nuI(ienIe: pc(c c :eu c(mczencmenIc ccm ccicc: cc(cc:.
Cucncc hc cumenIc cc ccncenI(cccc ce /ceIi|-Cc/ e /IF miIcccnc(ici:. c
ciI(cIc e I(cn:pc(Iccc pc(c fc(c cc c|u|c.
C ciI(cIc Ic(nc-:e enIcc ccIivccc( c|c:Ie(icc cc cceIi|-Cc/ cc(Lcxi|c:e
/c me:mc Iempc c ciI(cIc iniLe c fc:fcf(uIccinc:e (ecuzincc c c|icc|i:e.
/ cceIi|-Cc/ cc(Lcxi|c:e e (ecu|ccc pc( mccificcccc ccvc|enIe.
/ fc:fc(i|cccc ce:eccceccc pe|c: hc(mcnc: c|uccccn e epinef(inc ce:ccIivcm-
nc
Se c cci|-Cc/ :e ccumu|c ccmc (e:u|Iccc cc cumenIc cc |ipccne:e cu inf|uxc
ce ccicc: cc(cc: |iv(e: nc Iecicc. hcve(c iniLiccc cc :|nIe:e ce ccicc: cc(cc:.

Exi:Ie icuc|menIe umc (e|cccc inve(:c enI(e c ccncenI(cccc ce ccicc: cc(cc:
|iv(e: e c p(cpc(ccc cc pi(uvcIc ce:ic(ccenc:e ccIivc e inccIivc. umc vez cue
e:Ic (ecu|c c ci:pcniLi|iccce ce cceIi|-Cc/ pc(c c |ipccne:e. / cxiccccc cc
/ceIi|-Cc/. cevicc cc cumenIc ce ccicc: cc(cc: |iv(e:. pcce cumenIc( c (e|cccc
/ceIi|-Cc//Cc/ e N/DH/N/D
+
nc miIcccnc(ic. iniLincc c pi(uvcIc ce:ic(ccenc:e.

/ in:u|inc e:Iimu|c c |ipccne:e pc( vc(ic: meccni:mc:. nc mecicc em cue
cumenIc c I(cn:pc(Ie ce c|icc:e cc c|u|c e. ccn:ecuenIemenIe c ci:pcniLi|iccce
ce ccicc pi(vicc pc(c c :|nIe:e ce ccicc: cc(cc:.
/ c|ucccinc Iem um pcpe| inve(:c. umc vez cue iniLe c /ceIi|-Cc/ cc(Lcxi|c:e e.
pc(IcnIc. c |ipccne:e.

Se c :|nIe:e ce ccicc: cc(cc: e c -cxiccccc ccc((e::em :imu|IcnecmenIe.
ccn:IiIui(icm um cic|c fIi|. / -cxiccccc e iniLicc pe|c mc|cni|-Cc/. I(cvcncc c
ccIiviccce cc cc(niIinc cci|-I(cn:fe(c:e l. EnIcc. c p(imei(c inIe(mecic(ic cc :|nIe:e ce
ccicc: cc(cc: vci L|ccuec( c -cxiccccc cc n|ve| cc :i:Iemc ce I(cn:pc(Ie cc
memL(cnc miIcccnc(ic| inIe(nc.

c|dos gordos |nsoturodos (dessoturoo)
Dessoturoo: inI(ccuccc ce cup|c: |icccce:. ccIc|i:ccc pc( ce::cIu(c:e:.
/: ce::cIu(c:e: :cc cxicc:e: exi:IenIe: nc FE. uIi|izcncc c N/DFH cu N/DH.
ciIcc(cmc 85 e cxicnic mc|ecu|c(. ccIucncc :cL(e c cci|-Cc/ cc ccicc c
I(cn:fc(mc(.
C: e|ecI(ce: p(cvenienIe: cc N/DFH :cc I(cn:fe(icc: cI(cv: cc ciIcc(cmc pc(c c
cxicnic. ccIivcncc-c e c::im e|e i(c cxicc( c ccicc cc(cc.
59

C :e( humcnc Iem ce::cIu(c:e: cue ccIucm nc: pc:icce: 5. e . c cue :icnificc
cue
impc::|ve| inI(ccuzi( |icccce: cup|c: enI(e c
cc(Lcnc e c cc(Lcnc m.
C: ccicc: cc(cc: in:cIu(ccc: ce:empenhcm
vc(ic: funcce: impc(IcnIe: nc (ecu|cccc cc
nc::c c(ccni:mc {ex. mcnuIenccc cc f|uicez
cc memL(cnc. (ecuccc cc cc|e:Ie(c|
p|c:mcIicc).

c|dos gordos essenc|o|s: p(ecu(:c(e:
nece::c(ic: pc(c c :|nIe:e ce cuI(c: p(ccuIc:
{ex.: |inc|eicc. |inc|enicc e c(ccuiccnicc)

2. 8|oss|ntese de Ir|oc||g||cero|s
C: ccicc: cc(cc: :inIeIizccc: cu ince(icc: pe|c c(ccni:mc Im 2 ce:Iinc: pc::|vei:.
ccn:ccnIe c nece::iccce cc c(ccni:mc:
lncc(pc(cccc em I(icci|c|ice(ci: pc(c c c(mczencmenIc ce ene(cic meIcLc|icc
lncc(pc(cccc nc: ccmpcnenIe: fc:fc||picicc: cc: memL(cnc:

C: I(icci|c|ice(ci: e c: c|ice(cfc:fc|ipicc: {Ic| ccmc c fc:fcci|eInc|cminc) pc(Ii|hcm 2
pe(cu(:c(e:: ccicc: cc(cc: e c|ice(c| 3-F.

1. Act|voo do G||cero|:
/ fc(mc ccIivc cc c|ice(c| c ciice(c|-3-fc:fcIc. cue pcce :e( fc(mccc ce 2 mccc::
o. Fecuccc cc ci-hic(cxicceIcnc-fc:fcIc. ccIc|i:ccc pe|c G|ice(c|-3-fc:fcIc
ce:ic(ccenc:e

D|h|drox|cetono |os|oto + NADH G||cero| 3-|os|oto + NAD
+

G|ice(c| 3 fc:fcIc ce:ic(ccenc:e

b. Fc:fc(i|cccc cc c|ice(c| pe|c ccccc cc G|ice(c|cinc:e {f|cccc e (im)

G||cero| + AIF G||cero| 3-|os|oto + ADF
G|ice(c|cinc:e

C: Iecicc: cue pc::uem pcucc cu nenhumc c|ice(c|cinc:e. ccmc c cc:c cc
Iecicc ccipc:c. cLIm c c|ice(c|-3-fc:fcIc cc: inIe(mecic(ic: c|icc||Iicc:. l:Ic :icnificc.
cue c: ccipcciIc: cevem Ie( c|icc:e pc(c (ec|izcccc cc Lic::|nIe:e ce I(icci|c|ice(ci:.
/::im. c c|icc:e mc:I(c :e( c mcic( (ecu|ccc( ce:Ic :|nIe:e. pci: cI(cv: cc :eu
meIcLc|i:mc cue p(cvm c mcic( pc(Ie cc c|ice(c|-3-fc:fcIc.

2. Act|voo dos c|dos gordos:
C: ccicc: cc(cc: :cc ccIivccc: c /ci|-Cc/. pe|c ccccc cc /ci|-Cc/ :inIeIc:e {c
me:mc enzimc (e:pcn:cve| pe|c ccIivcccc nc -cxiccccc)

c|dos Gordos + CoA + AIF Ac||-CoA + AMF
/ci|-Cc/ :inIeIc:e

60

3. Formoo do c|do |os|ot|d|co ou d|oc||g||cero| 3-Fos|oto

/ fc(mcccc ce ccicc fc:fcI|cicc {inIe(mecic(ic
chcve) (e:u|Ic ce cuc: cci|ccce: :ecuencici: cc
c|ice(c|-3-fc:fcIc. nc p(e:encc ce c|ice(c|fc:fcIc
cci|I(cn:fe(c:e: l e ll.
/ci|cccc ce 2 c(upc: hic(cxi|c cc c|ice(c| 3-
Fc:fcIc pc( 2 mc|cu|c: ce cci| Cc/ pc(c c(icinc(
cicci|c|ice(c| 3 Fc:fcIc cu fc:fcIiccIc.
C cicci|c|ice(c|-3-Fc:fcIc c pe(cu(:c( ce
I(icci|c|ice(ci: mc: IcmLm ce
c|ice(cfc:fc||picc:.

4. Formoo de tr|oc||g||cero|s

C ccicc fc:fcI|cicc I(cn:fc(mc-:e num
1.2-cicci|c|ice(c|. pe(cencc c c(upc
fc:fcIc. (eccccc ccIc|i:ccc pe|c
fc:fcIiccIc fc:fcIc:e.
/ ccmLincccc cc 1.2-cicci|c|ice(c|
ccm um cuI(c cci|-Cc/ {pc(
I(cn:e:Ie(ificcccc) fc(mc um
I(icci|c|ice(c|. cujc (eccccc
ccIc|izccc pe|c cicci|c|ice(c|-
cci|I(cn:fe(c:e.
/ fc:fcIiccIc fc:fcIc:e e c
cicci|c|ice(c|-cci|I(cn:fe(c:e fc(mcm
um ccmp|exc enzimcIicc
{I(icci|c|ice(c|-:inIeIc:e)

kegu|oo do b|ossntese de tr|oc||g||cero|s:
/ Lic::inIe:e e c cec(cccccc ce I(icci|c|ice(ci: :cc (ecu|ccc: (ecip(cccmenIe. e
c vic fcvc(ecicc cepence cc: (ecu(:c: meIcLc|icc: e cc: nece::iccce:
mcmenIcnec:.
/ Icxc ce :|nIe:e ce I(icci|c|ice(ci: |c(ccmenIe c|Ie(ccc pe|c ccccc ce cive(:c:
hc(mcnc:.
/ in:u|inc p(cmcve c ccnve(:cc ce cc(Lchic(cIc: em I(icci|c|ice(ci:.
/: pe::cc: ccm cicLeIe: me||iIu:. cevicc c fc|Ic ce ccccc cu :ec(eccc ce
in:u|inc. ncc :c ncc ccn:ecuem uIi|izc( c c|icc:e ce mccc cp(cp(iccc ccmc
IcmLm ncc :cc ccpcze: ce :inIeIizc( ccicc: cc(cc:. e Im pc( i::c e|evccc:
Icxc: ce -cxiccccc e fc(mcccc ce cc(pc: ceIcnicc:.

61

Absoro dos tr|oc||g||cero|s exogenos
/pc: c :uc ince:Icc. c: cc(cu(c: c|imenIc(e: :cc emu|:icnccc:. ccnciccc
nece::c(ic pc(c c ccccc :uL:ecuenIe cc |ipc:e pcnc(ecIicc.
/ emu|:cc ccmecc c (ec|izc(-:e nc e:Icmccc e Ie(minc nc inIe:Iinc.
Nc cuccenc. c emu|:cc |ip|cicc pc:Ic em ccnIccIc ccm c |ipc:e pcnc(ecIicc e
ccm c: :ci: Li|ic(e:.
Ccm c hic(c|i:e cc: I(icci|c|ice(ci:. c: p(ccuIc: vcc :e( cc(ecccc: em mice|c: pe|c
ccccc cc: :ci: Li|ic(e:.
C p(ccuIc p(incipc| cc cice:Icc inIe:Iinc| ce ||picc: c 2-mcnccci|c|ice(c|.
cL:c(vicc ccmc Ic| nc: c|u|c: cc mucc:c inIe:Iinc|.
Umc enzimc. ne::c: c|u|c:. ccIc|i:c c cci|cccc ce::e: mcnccci|c|ice(ci:. ccm cci|-
Cc/. pc(c fc(mc( 1.2-cicci|c|ice(ci:. c: cuci: pccem :e( cci|ccc: c I(icci|c|ice(ci:.
C: I(icci|c|ice(ci: fc(mccc: i(cc fc(mc( c: cui|cmic(cn:.

3. 8|ossntese de |os|o|p|dos ou g||cero|os|o||p|dos
Nc ce(c|. c fc(mcccc ce fc:fc|ipicc: c pc(Ii( ce :imp|e: p(ecu(:c(e: (ecue(:
1. S|nIe:e cc mc|cu|c e:cue|eIc {c|ice(c| cu e:fincc:inc)
2. Liccccc cc{:) ccicc{:) cc(cc{:) cc e:cue|eIc em |iccccc :Ie( cu cmicc.
3. /ciccc ce um c(upc ce ccLecc" hic(cfi|icc pc( umc |iccccc fc:fcci:Ie(
4. /|Ie(cccc cc muccncc cc c(upc ccLecc pc(c c(icinc( c p(ccuIc finc| {nc|cun:
cc:c:)

Loco|: ccc((e p(imc(icmenIe nc :upe(f|cie cc (eI|cu|c enccp|c:mcIicc e nc memL(cnc
inIe(nc cc miIcccnc(ic ce Iccc: c: c|u|c:

C: p(imei(c: pc::c: cc :|nIe:e ce c|ice(cfc:fc|ipicc: :cc ccmun: c vic cc:
I(icci|c|ice(ci:: 2 c(upc: cci| :cc e:Ie(ificccc: nc C1 e C2 cc c|ice(c|-3 Fc:fcIc. pc(c
fc(mc( |os|ot|doto.
C fc:fcIiccIc IcmLm pcce :e( cLIicc pc( fc:fc(i|cccc ce um cicci|c|ice(c| pc(
umc cinc:e e:pec|ficc.
/ cobeo po|or cc: c|ice(cfc:fc|ipicc: |icccc pc( umc |iccccc fc:fcci:Ie(. nc
cuc| cccc um cc: 2 hic(cxi|c: c|ccc|icc: {1 c(upc pc|c( e um c|ice(c|) fc(mc um
:Ie( ccm c fc:fcIiccIc.
Nc p(cce::c Lic::inIIicc um cc: hic(cxi|c: p(imei(c ccIivccc pc( umc |iccccc ce
um nuc|ecIicc. ciIicinc cifc:fcIc {CDF). cLIencc-:e CMF pc( cIccue ce cuI(c
c(upc hic(cxi|c.
C CDF pcce :e( |icccc cc cicci|c|ice(c| fc(mcncc c fc:fcIiccIc ccIivccc CDF-
cicci|c|ice(c|. cu cc c(upc hic(cxi|c cc c(upc pc|c(.

Fos|ot|d||co||no

1. Cc|inc |iv(e fc:fc(i|ccc pe|c /IF pe|c cc|inc cinc:e
2. / fc:fc(i|cc|inc. cI(cv: cc ciIicinc I(ifc:fcIc {CIF). ccnve(Iicc c CDF-cc|inc pe|c
fc:fc(i|cc|inc ciIici|I(cn:fe(c:e |iLe(Icncc Fi.
3. / |iccccc pi(cfc:fc(i| ce c|Ic ene(cic cc CDF-cc|inc muiIc in:Icve| e (eccIivc
pc(c cue c uniccce ce fc:fc(i|cc|inc :ejc I(cn:fe(icc pc(c um cenI(c nuc|ecfi|icc
p(cmcvicc pe|c c(upc hic(cxi|c cc pc:iccc 3 cc 1.2-cicci|c|ice(c| pe|c cc|inc
fc:fc(i|I(cn:fe(c:e.

62

Fc:fccc|inc ciIici|I(cn:fe(c:e
Formo oct|vo: |icccc cc (eI|cu|c enccp|c:mcIicc.
Formo |noct|vo: nc ciIc:c| {:e(ve ccmc (e:e(vc ce:Ic: enzimc:)
/ I(cn:|cccccc cc enzimc cc ciIc:c| pc(c c FE (ecu|ccc pe|c c/MF e cci|-Cc/.
/ fc:fc(i|cccc cc enzimc. cinc:e c/MF-cepencenIe. ccu:c c |iLe(Icccc cc
memL(cnc {inccIivcccc)
/ci|-Cc/ p(cmcvem c |iccccc cc FE.

Nc f|cccc. c fc:fcIici|cc|inc pcce :e( fc(mccc cI(cv: cc meIi|cccc (epeIicc cc
|os|ot|d||etono|om|no pe|c fc:fcIici|eIcnc|cminc N-meIi|I(cn:fe(c:e. C: c(upc: meIi|c :cc
(emcvicc: cc S-ccenc:i|meIicninc.
Fos|ot|d||etono|om|no
1. / eIcnc|cminc fc:fc(i|ccc pe|c /IF pe|c eIcnc|cminc cinc:e. :encc
pc:Ie(ic(menIe ccIivccc c CDF-eIcnc|cminc pe|c fc:fceIcnc|cminc
ciIici|I(cn:fe(c:e.
2. / CDF-eIcnc|cminc I(cn:fe(icc pc(c c 1.2-cicci|c|ice(c| pe|c eIcnc|cminc
fc:fcI(cn:fe(c:e. fc(mcncc fc:fcIici|eIcnc|cminc.

Nc: miIcccnc(ic: cc f|cccc fc:fcIici|eIcnc|cminc IcmLm :inIeIizccc c pc(Ii( cc
ce:cc(Lcxi|cccc cc fc:fcIici|:e(inc.

Fos|ot|d||ser|no
:inIeIizccc c pc(Ii( cc I(ccc cc ccLecc pc|c( cc fc:fcIici|eIcnc|cminc pe|c :e(inc.
umc (eccccc /IF incepencenIe e (eve(:|ve|
/ (eccccc iniciccc pe|c cIccue" cc |iccccc fc:fccie:Ie( cc
fc:fcIici|eIcnc|cminc pe|c c(upc hic(cxi|c cc :e(inc.

Fos|ot|d|||nos|to|
:inIeIizccc pe|c vic cc CDF-cicci|c|ice(c| c inc:iIc| |iv(e.
Feccccc ccIc|izccc pe|c fc:fcIici|inc:iIc| :inIc:e {nc FE).

63

4. 8|ossntese de es||ngo|p|dos
Fercursores: pc|miIci|-Cc/ e :e(inc
Etopos:
1. S|nIe:e ce e:finccninc c pc(Ii( cc po|m|to||-CoA e ser|no
2. Liccccc ce um ccicc: cc(cc em |iccccc cmicc pc(c c(icinc( N-oc||es||ngon|no
3. De::cIu(cccc ce e:finccninc pc(c fc(mc( N-oc||es||ngos|no {cerom|do)
4. Liccccc ce um c(upc pc|c( pc(c p(ccuzi( um e:fincc||picc. Ic| ccmc um
cerebros|do cu es||ngom|e||no

Nc fc(mcccc cc: ce(eL(c:icc: c: cccc(e: enI(cm nc fc(mc cc: :eu: ce(ivccc: ce
nuc|ecIicc: ccIivccc:.

5. 8|ossntese do co|estero|
Cc|e:Ie(c|:
E:Ic p(e:enIe nc: Iecicc: e nc: |ipcp(cIe|nc: p|c:mcIicc:. |iv(e cu ccmLinccc ccm
ccicc: cc(cc: ce ccceic |cncc {:Ie(e: ce cc|e:Ie(c|)
:inIeIizccc c pc(Ii( ce um pe(cu(:c( nicc {/ceIi|-Cc/)
e|iminccc cc c(ccni:mc nc L||i: ccmc cc|e:Ie(c| cu ccicc: Li|ic(e:.
pe(cu(:c( ce Iccc: c: e:Ie(cice: cc c(ccni:mc {ex.: cc(Iiccice:. hc(mcnc: :exuci:.
ccicc: Li|ic(e: e viIcminc D)
um |ipicc cnfipcIicc:
Fe:pcn:cve| pe|c f|uicez cc: memL(cnc:
CcmpcnenIe cc ccmccc mci: exIe(nc cc: |ipcp(cIe|nc: p|c:mcIicc:.

Loco|: (eI|cu|c enccp|c:mcIicc e nc ciIc:c| ce Iccc: c: c|u|c:.
Fercursor {nicc): /ceIi|-Cc/

Etopos do b|ossntese

1. $ntese do c|do Mevo|on|co (Mevo|onoto)
2 Mc|cu|c: ce ocet||-CoA ccncen:cm-:e fc(mcncc Acetoocet||-CoA. (eccccc
ccIc|i:ccc pe|c Iic|c:e. umc enzimc ciIc:c|icc.
Nc f|cccc. c ccicc cceIcccIicc :inIeIizccc nc miIcccnc(ic pe|c vic cc
ceIccne:e e cifunce-:e pc(c c ciIc:c| cnce ccIivccc c Acetoocet||-CoA pe|c
/ceIccceIi|-Cc/ :inIeIc:e. nece::iIcncc pc(c Ic| ce Cc/ e /IF.
C /ceIccceIi|-Cc/ ccncen:c-:e ccm mci: umc mc|cu|c ce /ceIi|-Cc/. (eccccc
ccIc|i:ccc pe|c HMG-Cc/ :inIeIc:e. fc(mcncc-:e HMG-CoA
/ pc(Ii( cc HMG-Cc/ fc(mc-:e c cicc Mevc|cnicc pc( (ecuccc em 2 eIcpc:.
(eccccc ccIc|i:ccc pe|c HMG-Cc/ (ecuIc:e. N/DFH-cepencenIe.
E:Ic c eIcpc (ecu|ccc(c cc :|nIe:e cc cc|e:Ie(c|. i((eve(:|ve|. Se c Lic::|nIe:e fc(
:u:pen:c. fc(mc(-:e-c cc(pc: ceIcnicc:.

64

2. Formoo de |sopreno oct|vodo
C cicc Mevc|cnicc {C) fc:fc(i|ccc pe|c /IF fc(mcncc-:e vc(ic: |nIe(mecic(ic:
fc:fc(i|ccc: ccIivc:.
Fc( meic ce umc ce:cc(Lcxi|cccc. fc(mc-:e c uniccce i:cp(encice ccIivc. c
i:cpenIeni|pi(cfc:fcIc {C5)

3. $ntese do esquo|eno
C l:cpenIeni|pi(cfc:fcIc umc e:I(uIu(c ccIivccc em C5. c pc(Ii( cc cuc| :e i(c
fc(mc( umc e:I(uIu(c em C1C. c Ge(cni|pi(cfc:fcIc.
/c Ge(cni|pi(cfc:fcIc junIc-:e cuI(c mc|cu|c ce l:cpenIi|pi(cfc:fcIc fc(mcncc umc
e:I(uIu(c em Cl5. c Fc(ne:i|pi(cfc:fcIc {pcce cc( c(icem c uLicuincnc cu cc cc|icc|)
2 mc|cu|c: ce Fc(ne:i|pi(cfc:fcIc :cf(em ccncen:cccc. fc(mcncc c e:cuc|enc
{C30)

4. $ntese do co|estero|
C E:cuc|enc ccnve(Iicc c Lcnc:Ie(c| pc( (ecccce: ce cic|izcccc {N/DFH e C2
cepencenIe:) pc( ccccc ce umc cic|c:e.
E:Ic eIcpc ccc((e nc: memL(cnc: cc FE e envc|ve muccncc: nc nc|ec e:Ie(cice
e nc ccceic |cIe(c|. c::im ccmc c pe(cc ce 3 c(upc: meIi|.
C: inIe(mecic(ic: cc E:cuc|enc cI cc cc|e:Ie(c| e:Icc |icccc: c umc p(cIe|nc -
F(cIe|nc I(cn:pc(Iccc(c ce E:Ie(ci: {SFC - SIe(cic F(cIein Cc((ie() - cue Iem pc(
funccc ce p(cmcve( c ccnIccIc ce:Ie: :uL:I(cIc: |ipc::c|vei: ccm um meic
ccuc:c.

65

Fr|nc|po|s der|vodos do Co|estero|
c. Sci: e ccicc: Li|ic(e:
C ccicc cc|icc e c: vc(ic: ccicc: Li|ic(e: :c|uLi|izcm c: |ipicc: c|imenIc(e:.
fcci|iIcncc c :uc cice:Icc.
C ccicc cc|icc e c ccicc ce:cxicc|icc enccnI(cm-:e nc L||i:. ccnjucccc: ccm c
c|icinc cu ccm c Icu(inc. fc(mcncc cc: ccicc: c|icccc|icc. c|iccce:cxicc|icc e
Icu(ccc|icc. ccm ice: mcncvc|enIe: {Nc
+
. K
+
) fc(mcm c: :ci: Li|ic(e:.

Co|estero| Co|oto Co|||-CoA G||coco|oto

L. Hc(mcnc: e:Ie(cice:
Frogesterono: :ec(ecccc cc n|ve| cc cc(pc cmc(e|c. cc p|ccenIc e cc: ccp:u|c:
:up(c((enci: - inIe(vm nc niccccc e ce:Icccc.
M|nero|ocort|co|des: :cc :ec(ecccc: cc n|ve| cc: ccp:u|c: (enci: e pe(miIem c
(ecL:c(ccc cc :ccic e cc c|c(c nc (im.
Androgn|os {ex.: Ie:Ic:Ie(cnc): :ec(ecccc cc n|ve| cc Ie:I|cu|c :encc (e:pcn:cve|
pc( vc(ic: cc(ccIe(e: :exuci: mc:cu|inc:.
Estrogn|os {ex.: e:I(ccic| e e:I(cnc) :ec(ecccc: nc: cvc(ic: e nc p|ccenIc. :cc
(e:pc:cvei: pc( vc(ic: cc(ccIe(e: :exuci: femininc: - cp(e:enIcm nc :uc e:I(uIu(c um
c(upc fenc|icc.
G||cocort|co|des {ex.: cc(Ii:c| e cc(Ii:cnc) :cc hc(mcnc: cc(Iicc-:up(c((enci: cue
e:Iimu|cm c meIcLc|i:mc p(cIeicc e c necc|icccne:e cc n|ve| cc f|cccc.
kegu|oo
/ enzimc (ecu|ccc(c e c HMG-Cc/ (ecuIc:e.
Hc umc ciminuiccc mc(ccnIe nc ccIiviccce ce:Ic enzimc nc: inciv|cuc: em jejum. c
cue exp|icc c :|nIe:e (ecuzicc ce cc|e:Ie(c| cu(cnIe c jejum.
Act|vodores:
ln:u|inc
Hc(mcnc: Ii(ciceic:
In|b|dores:
G|icccinc
G|icccc(Iiccice: {ex: cc(Ii:c|)
lDL-cc|e:Ie(c| ccpIccc vic (ecepIc(e: ce lDL {(ecepIc(e: cpc 8-100 e cpc E).

A nve| ce|u|c| :cc ccn:|ce|ccc: c: :egu|nte: c|cce::c: cue ccnt|c|cn c cc|cncc cc
cc|e:te|c|.
C oumento cevicc c:
CcpIcccc cc cc|e:Ie(c| cc: |ipcp(cIeinc: {(icc: em :Ie(e: ce cc|e:Ie(c|) vic
(ecepIc(e: {ex: (ecepIc(e: ce LDL)
S|nIe:e cc cc|e:Ie(c|
Hic(c|i:e cc: :Ie(e: ce cc|e:Ie(c| pe|c enzimc Cc|e:Ie(c| e:Ie(ec:e.
/ d|m|nu|o cevicc c:
F|uxc ce cc|e:Ie(c| cc memL(cnc pc(c c: |ipcp(cIeinc: ce Lcixc pcIencic| em
cc|e:Ie(c|. p(cmcvicc pe|c LC/I {LeciIinc Cc|e:Ie(c| /ci| I(cn:fe(c:e)
E:Ie(ificcccc cc cc|e:Ie(c| pe|c /C/I {/ci|-Cc/ Cc|e:Ie(c| /ci| I(cn:fe(c:e)
UIi|izcccc cc cc|e:Ie(c| pc(c c :inIe:e ce hc(mcnc:. ccicc: e :ci: Li|ic(e:.

66

Ironsporte ||p|d|co
/ ene(cic ci:pcn|ve| nc: ccicc: cc(cc: nece::iIc ce :e( ci:I(iLu|cc cI(cv: cc
c(ccni:mc. c pc(Ii( cc |ccc| ce cL:c(ccc. Lic::|nIe:e cu c(mczencmenIc pc(c c:
Iecicc: funcicnci: cue c uIi|izem.

Fc(mc: ce I(cn:pc(Ie ce |ipicc: nc :cncue
1. L|poprote|nos p|osmt|cos. nc: cuci: c: I(icci|c|ice(ci: e cuI(c: ||picc: :cc
I(cn:pc(Iccc: ccmc ccI|cu|c: ce ||picc: (eccLe(Ic: pc( p(cIe|nc::
2. cicc: Gc(cc: Liv(e: |icccc: c A|bum|no sr|co: {Iecicc ccipc:c f|cccc)
3. Cc(pc: CeIcnicc:: cicc /ceIcccIicc e -Hic(cxiLuI|(icc {f|cccc m:cu|c:.
cc(cccc. c(eL(c)

L|poprotenos p|osmt|cos: ccmp|exc: mcc(cmc|ecu|c(e: e:f(icc:. ccn:IiIu|cc: pc(:
Um cc(e {nc|ec cenI(c|) ccnIencc |p|dos neutros cu opo|ores {:Ie(e: cc
cc|e:Ie(c| e I(icci|c|ice(ci:).
/c (ecc( hc umc ccmccc ce protenos e |p|dos ont|pt|cos {fc(mcncc umc mcnc
ccmccc |ip|cicc cue fczem c inIe(fcce enI(e c: fc:e: ccuc:c e |ipicicc) ccmpc:Ic
ce cc|e:Ie(c| |iv(e {ncc e:Ie(ificccc) e fc:fc||picc:. :cL(eIucc LeciIinc
{fc:fcIici|cc|inc).
Ne::c ccmccc :upe(ficic| enccnI(cm-:e c: p(cIe|nc: - opoprotenos.
CucnIc menc( c pc(I|cu|c. mcic( c cen:iccce p(cIeicc nc envc|Ic(ic exIe(nc e
menc( c Iec( |ipicicc nc nc|ec.

C|oss|||coo
Lipcp(cIe|nc Contedo Apoproteno Or|gem Mecon|smo de ||bertoo
do contedo
Cui|cmic(c I(icci|c|ice(ci:
cc cieIc
{exccenc:)
/1. /2. /4.
848
lnIe:Iinc Hic(c|i:e pe|c |ipcp(cIeinc
|ipc:e
Cui|cmic(c
(emcne:cenIe
Cc|e:Ie(c| cc
cieIc
848. E EnccciIc:e meciccc pc(
(ecepIc(e: {f|cccc)
VLDL
{ve(y |cw cen:iIy
|ipcp(cIein:)
I(cci|c|ice(ci:
encccenc:
8100. C. E F|cccc e
inIe:Iinc
Hc(c|ie:e pe|c |ipcp(cIeinc
|ipc:e
lDL
{inIe(mecicIe DL)

Cc|e:Ie(c|
encccenc
8100. E VLDL EnccciIc:e meciccc pc(
(ecepIc(e: {f|cccc)
LDL
{|cw DL)
8100 lDL EnccciIc:e meciccc pc(
(epecIc(e: {f|cccc e cuI(c:
Iecicc:)
HDL
{hich DL)
/1. /2 F|cccc e
inIe:Iinc
I(cn:fe(ncic cc: :Ie(e: ce
cc|e:Ie(c| pc(c cuI(c:
|ipcp(cIe|nc: {LDL/lDL)

Apo||poprote|nos
/|cumc: cpcp(cIe|nc: :cc inIec(ci: e ncc pccem :e( (emcvicc: {cpc 8-100 e
8-48). encucnIc cuI(c: :cc pe(if(icc:. e:Icncc |iv(e: pc(c :e(em I(cn:fe(icc: c
cuI(c: |ipcp(cIeinc:.
/: /pcp(cIe|nc: pe(Iencem c c|c::e: ce:icnccc: pc( |eI(c: {/. 8. C. D e E).
C: memL(c: ce cccc c|c::e (eceLem nme(c: (cmcnc: {L ||. lll. lV e V).
/: cpc|ipcp(cIe|nc::
Ccnfe(em e:IcLi|iccce e:I(uIu(c| c: |ipcp(cIeinc:
DeIe(mincm c ce:Iinc meIcLc|icc cc: pc(I|cu|c: cue c: ccnIm
Scc cc-fccIc(e: enzimcIicc::
Se(vem ce |iccccc pc(c inIe(ccccc ccm c: (ecepIc(e: ce |ipcp(cIeinc: nc:
Iecicc:.

67

/pcp(cIeinc: cc :(ie /
/: cpc /-l. /-ll e /-lV :cc enccnI(ccc: nc HDL.
/ /-l e /-ll :cc :inIeIizccc: nc f|cccc e nc inIe:Iinc
/ cpc /-lV fc(mc-:e cpenc: nc inIe:Iinc.
/ /-l e::encic| pc(c c inIec(iccce cc: HDL e ccIivc c LC/I {cue e:Ie(ificc c
cc|e:Ie(c|).

/pcp(cIeinc: cc :(ie 8
/ cpc 8-100 p(ccuzicc nc f|cccc. cnce incc(pc(ccc c VLDL.
/ cpc-100 c nicc cpcp(cIelnc cc LDL e inci:pen:cve| c :|nIe:e e c :ec(eccc
cc VLDL.
c |iccnIe pc(c c (ecepIc( ce LDL - e:Ie umc p(cIe|nc ce :upe(f|cie ce|u|c( cue
fixc e inIe(nc|izc c LDL.
/ cpc 8-48 {Iem 487 cc: cmincccicc: cc /pc-100)e::encic| c :|nIe:e cc:
cui|cm|c(cn: cc n|ve| cc: inIe:Iinc:.

/pcp(cIeinc: cc :(ie C
Scc :inIeIizccc: nc f|cccc e e:Icc em Iccc: c: |ipcp(cIe|nc: - menc: nc LDL.
Iccc: c: cpc C iniLem c (emcccc cc: (emcne:cenIe: ce cui|cm|c(cn: e ce VLDL
pe|c ficccc.
/ cpc C-ll ccIivc c |ipcp(cIe|nc |ipc:e {LFL). cue cice(e {hic(c|i:c) c: I(icci|c|ice(ci:
{cc(cu(c) cc: cui|cm|c(cn: e cc: VLDL.
/ cpc C-lll. pe|c ccnI(c(ic. iniLe c LFL.

/pcp(cIeinc cc :(ie D: :cc enccnI(ccc: em :uf(cccce: cc HDL.

/pcp(cIeinc E
:inIeIizccc nc f|cccc. neu(cnic:. c|ic e mcc(cfccc:.
Ccc((e em Iccc: c: |ipcp(cIe|nc:. ccm excepccc cc LDL.
Fc(Iicipc nc ccpIcccc ce::c: |ipcp(cIe|nc: pe|c ficccc {vic (ecepIc( pc(c c LDL).
/u:ncic ce cpc E |evc c cIe(c:c|e(c:e p(eccce.

/pcp(cIeinc {/) minc(
umc c|iccp(cIe|nc enc(me. hcmc|ccc cc p|c:minccnic. :inIeIizccc pe|c ficccc.
Nc p|c:mc. fc(mc umc |iccccc ccvc|enIe ccm c cpc 8-100 cc: LDL. ce(cncc umc
ncvc |ipcp(cIe|nc. c Lp {c).
EnccnI(c-:e |icccc c cpc 8-100 cc LDL pc( umc |iccccc ci::u|ficicc.
/ cpc {c) pcce |icc(-:e c mcI(iz exI(cce|u|c( cc pc(ece vc:cu|c( iniLincc c
fiL(inc|i:e encccenc. pc( ccmpeIi( pc(c c: (ecepIc(e: ce p|c:minccnic. :eu
hcmc|ccc e:I(uIu(c|.
/::im. c cepc:iccc ce fiL(inc fcvc(ecicc nc: |ccci: ce |e:cc e p|ccc:
cIe(c:c|e(cIicc:.
/ cpc {c) e:IcLe|ece |icccce: ccm c |iccncc L cc mc|cu|c ce fiL(inc. impecincc
c::im c fixcccc cc p|c:minccnic e c :uc ccIivcccc pe|c: (e:pecIivc: ccIivccc(e:
Iecicu|c(e: {I-F/). impecincc c fiL(inc|i:e.
/ hcmcci:Ie|nc cumenIc c cfiniccce cc Lp {c) pc(c c |iccncc L cc mc|cu|c ce
fiL(inc. :encc pc( i::c um fccIc( ce (i:cc.
/ Lp {c) (ecuz c ccIivcccc cc IGF- {Iumc( c(cwIh fccIc( ). pc( (ecuccc cc :eu
ccIivccc(. c p|c:minc. l:Ic Ie(c ccmc ccn:ecuncic umc mcic( mic(cccc e
p(c|ife(cccc cc: c|u|c: mu:cu|c(e: |i:c: vc:cu|c(e:. fencmenc c(ucic| nc
cIe(ccne:e.
Fc( cuI(c |ccc. c cpc {c) umc impc(IcnIe p(cIe|nc ce fc:e ccucc pc:iIivc.
cumenIccc em |e:ce:. cc| c impc(Icncic nc (epc(cccc cc: Iecicc:. Ccmc e:Ic
p(e:enIe nc: p(imei(c: eIcpc: cc p(cce::c cu(cIivc. impece c fiL(inc|i:e e pc::iLi|iIc
cue c: ccccu|c: ce fiL(inc (evi:Icm c fe(icc.
c mci: cIe(ccnicc cc: |ipcp(cIe|nc:. e:Icncc cuc:e :emp(e e|evccc nc:
pccienIe: ccm enfc(Ie:. mc: IcmLm impc(IcnIe nc (epc(cccc Iecicu|c(.
68

Enz|mos envo|v|dos
Lipcp(cIe|nc Lipc:e {LFL) F|c:mcIicc

E:Ic p(e:enIe nc :upe(f|cie ccpi|c( ce Iccc c c(ccni:mc.
Hic(c|i:c c: I(icci|c|ice(ci: p(e:enIe: nc VLDL e nc: cui|cm|c(cn:. ce(cncc ccicc:
cc(cc: |iv(e: e c|ice(c| c n|ve| Iecicuc|.
C: ccicc: |iv(e: cifuncem pc(c c: Iecicc: :uLjccenIe:. mc: umc pecuenc pc(Ie
(ec(e::c c ci(cu|cccc cnce :e |icc c p(cIe|nc c|Luminc.
/ in:u|inc e:Iimu|c c :|nIe:e e c ccIiviccce cc LFL.
Nc cicLeIe: me||iIu:. c ciminuiccc cu cu:ncic cc efeiIc cc in:u|inc. p(ejucicc c
cepu(cccc ce I(icci|c|ice(ci:. c cue cc(cvc c hipe(I(icci|c|ice(icmic.
C: e:I(ccenic: c(icm umc ci:I(iLuiccc cife(enciccc cc Lipcp(cIe|nc |ipc:e
p|c:mcIicc. cefinincc c pcc(cc cc(pc(c| femininc.
/ :uc ccIiviccce cepencenIe cc /pc C-ll e cc in:u|inc.

Ir|oc||g||cero| 2 c|dos gordos + 2-Monooc||g||cero|
Lipcp(cIeinc |ipc:e

A LFL do coroo fem um Km bo|xo poro os fr|oc||g||cer|s, oo posso que o LFL do
fec|do od|poso fem um Km 10 vezes super|or. Forqu?
/ pc::ccem cc e:Iccc c|imenIccc pc(c um e:Iccc ce jejum cccmpcnhccc
pe|c ciminuiccc cc: I(icci|c|ice(ci: exi:IenIe: nc p|c:mc.
Ne::e cc:c. c LFL cc cc(cccc ccnIinuc :cIu(ccc ccm :uL:I(cIc. pc(cue exi:Ie umc
ciminuiccc cc :cIu(cccc cc LFL cc Iecicc ccipc:c.
Exi:Ie ccmc cue umc (eci(ecccc cc: I(icci|c|ice(ci:. cu :ejc. c: I(icci|c|ice(ci: cue
:e(icm ce:Iinccc: cc Iecicc ccipc:c. em ccncicce: nc(mci:. vcc pc(c c cc(cccc.
exccIcmenIe pc( ccu:c cc: Km cife(enIe:. c|m ce cue c: I(icci|c|ice(ci:
c(mczenccc: pccem :e( (euIi|izccc:.

Lipc:e Sinu:cicc| hepcIicc
Hic(c|i:c I(icci|c|ice(ci: e fc:fc|ipicc: cc lDL nc: :inu:cice: hepcIicc:. ccnve(Iencc-c
em LDL.
Ccnve(Ie c HDL em HDL3

Lipc:e cicc Li:c::cmicc
Hic(c|i:c c: (emcne:cenIe: cc: cui|cm|c(cn:. VLDL {lDL) e c HDL2 ccpIccc pe|c
f|cccc.

LC/I {LeciIinc Cc|e:Ie(c| /ci| I(cn:fe(c:e)
CcIc|i:c c I(cn:fe(ncic cc cci| {em ce(c| ccicc |inc|eicc. ccicc c(ccuiccnicc) cc
|eciIinc pc(c c cc|e:Ie(c| |iv(e - cmLc: ccmpcnenIe: cc mcncccmccc |ipicicc
exIe(nc. ce(cncc :Ie( ce cc|e:Ie(c| e |i:c|eciIinc.
/ (eccccc I(cn:fc(mc c mc|cu|c pc|c( cc cc|e:Ie(c| {exIe(nc) num :Ie( cpc|c(.
C: :Ie(e: ce cc|e:Ie(c| Im cci: ce:Iinc::
1. Mic(cm pc(c c (ecicc cpc|c( cenI(c| cc HDL cu:
2. Scc I(cn:fe(icc: pc(c VLDL e LDL.
/ /pc /-l ccfccIc( pc(c c ccccc cc LC/I.

Lec|t|no + Co|estero| L|so|ec|t|no + ster de co|estero|
LC/I

/C/I {/ci|-Cc/ Cc|e:Ie(c| /ci| I(cn:fe(c:e)
Iem ccmc funccc c fc(mcccc ce :Ie(e: ce cc|e:Ie(c|

Co|estero| + O|e||-CoA O|eoto de co|estero| + CoA$H
/C/I
69

Frotenos de Irons|ernc|o
CEIF {p(cIe|nc ce I(cn:fe(ncic ce :Ie(e: ce cc|e:Ie(c|)
p(ccuzicc nc f|cccc e ci(cu|c em c::ccicccc ccm LC/I nc HDL.
/cIuc nc I(cn:fe(ncic ce :Ie(e: ce cc|e:Ie(c| cc: HDL pc( I(ccc cc: I(icci|c|ice(ci:
cc: VLDL. LDL e em menc( e:cc|c pc(c c: cui|cm|c(cn:.
um ccIivccc( inci(ecIc cc LC/I. umc vez cue (emcve c :eu p(ccuIc ce iniLiccc.
ce :c|ienIc( cue e:Ic I(ccc :c :e (ec|izc :e exi:Ii(em I(icci|c|ice(ci:.
Se c cucnIiccce ce I(icci|c|ice(ci: ince(icc fc( e|evccc vcc exi:Ii( I(icci|c|ice(ci:
:uficienIe: pc(c c I(ccc enI(e c: |ipcp(cIeinc: e ccn:ecuenIemenIe c cc|e:Ie(c| vci
pc::c( cc: HDL pc(c c: cuI(c: |ipcp(cIeinc:. C(c Ic| p(cce::c ncc Lenficc umc
vez cue c cc|e:Ie(c| I(cn:pc(Iccc pe|c: HDL ce:Iinc-:e c :uc e|imincccc nc ficccc.
Fc( cuI(c |ccc exi:Ie c cumenIc ce pc(I|cu|c: LDL cen:c: e pecuenc:.
DeficiI: ne:Ic p(cIe|nc |evcm cc cumenIc cc HDL.
FLIF {p(cIe|nc: ce I(cn:fe(ncic ce fc:fc|ipicc:): pc(Iicipcm nc I(ccc ce fc:fc||picc: e
I(icci|c|ice(ci: cc: cui|cm|c(cn: e cc: VLDL pc( :Ie(e: ce cc|e:Ie(c| cc HDL3.

1. Ironsporte de Lp|dos Exogenos - Qu||omcrons
/pc: c cice:Icc. c co|estero| e c: tr|oc||g||cero|s :cc cL:c(vicc: e incc(pc(ccc: nc
cc(e cc: qu||omcrons noscentes.
/ Apo 8-48. Apo A-I e Apo A-IV :cc ce(ccc: nc (eI|cu|c enccp|c:mcIicc (ucc:c cc
enIe(cciIc e ccmLincm-:e ccm c cc|e:Ie(c|. fc:fc||picc: e I(icci|c|ice(ci: {cc(cu(c:)
cL:c(vicc:.
C: cui|cm|c(cn: fc(mccc: vcc pc(c ve:|cu|c: :ec(eIc(ic:. :cc |cncccc: nc ||cuicc
exI(cce|u|c( e ccpIccc: pe|c s|stemo ||n|t|co {cevicc cc :eu Icmcnhc).
ScmenIe c: compostos hdrosso|ve|s :cc cL:c(vicc: pe|c: cop||ores songuneos
me:enI(icc: e |evccc: cc f|cccc pe|c :i:Iemc pc(Ic. c: ccmpc:Ic: |ipc::c|vei:
:cc cL:c(vicc: pe|c :i:Iemc |infcIicc.
C: cui|cmic(c: inIe(ccem ccm c HDL. ce cuem (eceLem Apo E {nece::c(ic pc(c
cue c: cui|cm|c(cn: (emcne:cenIe: pc::cm :e( ccpIccc: pe|c f|cccc) e c Apo C-II
{e::encic| pc(c c hic(c|i:e cc: I(icci|c|ice(ci:). /: opo C :cc cevc|vicc: c HDL.
/ Apo C-II ccIivc c LFL. cue hic(c|i:c c: I(icci|c|ice(ci:. c: cui|cmic(c pe(cem 07
cc: I(icci|c|ice(ci: fc(mcncc-:e c: qu||om|cros remonescentes.
C: ccicc: cc(cc: :cc ccpIccc: e uIi|izccc: pe|c: Iecicc:.
C: cui|cmic(c: (emcne:cenIe: pe:cm ce(cc ce meIcce (e|cIivcmenIe cc
cui|cmic(c. :encc c :uc f(ccccc em cc|e:Ie(c| e :Ie(e: ce cc|e:Ie(c| mcic( cevicc
c pe(cc cc: I(icci|c|ice(ci:.
C: cui|cmic(c: (emcne:cenIe: :cc ccpIccc: pe|c f|cccc {:encc cec(ccccc: pe|c
|ipc:e |i:c::cmicc hepcIicc). ScmenIe ce:Ie mccc c cc|e:Ie(c| {e c: cemci: |ip|cc:
ince(icc: nc cieIc) cIince c f|cccc.
C cc|e:Ie(c| ccpIccc pe|c f|cccc. pccencc :ecui( pc( I(: ccminhc::
1. Exc(eIccc nc L||i: ccmc cc|e:Ie(c| |iv(e:
2. Ccnve(Iicc c ccicc: Li|ic(e: e |cncccc nc L||i::
3. Sec(eIccc: pc(c c :cncue cenI(c cc: VLDL.

2. Ironsporte de |p|dos endogenos
c. Si:Iemc cc cpc 8-100. cue I(cn:pc(Ic ||picc: cc f|cccc pc(c c: Iecicc: pe(if(icc::
L. Si:Iemc cc cpc /-l. cue (ec|izc c I(cn:pc(Ie (eve(:c {cc pe(ife(ic pc(c c f|cccc).
Iecicc /cipc:c f|cccc C|u|c pe(if(icc

70

o. Ironsporte de Lp|dos Endogenos - VLDL e LDL. $|stemo do Apo 8-100

ExcecenIe: cc|c(icc:. ccn:umc ce c|ccc| e cicLeIe: me||iIu: |evcm c ||pognese.
ccm c ccnve(:cc ce c|icc:e em ccicc: cc(cc:. fc:fc||picc: e cc|e:Ie(c| e ce:Ie:
em tr|oc||g||cero|s {cc(cu(c:) e steres de co|estero|.
E:Ie: ||picc: :cc (eunicc: ccm opo 8-100. C-II. C-III e E. ce(cncc c VLDL. cue
|iLe(Iccc pe|c f|cccc ccm ce:Iinc cc Iecicc ccipc:c.
/: VLDL inIe(ccem ccm c: HDL (eceLencc mci: opo E e opo C-II. E:Ic ccIivc c LFL.
cue hic(c|i:c I(icci|c|ice(ci:. (ecuzincc c VLDL ce Icmcnhc e Ic(ncncc-:e mci:
cen:c c mecicc cue vci pe(cencc I(icci|c|ice(ci:.
Nc finc| ce::e p(cce::c. (e:Icm VLDL remonescentes cu IDL
Fc(Ie cc: lDL :cc ccpIccc: pe|c f|cccc e cec(ccccc:. c (e:IcnIe ccnve(Iicc em
LDL. Nc p(cce::c. lDL pe(ce tr|oc||g||cero|s. Apo E e Apo C-II. c cue (ep(e:enIc um
cumenIc nc ccncenI(cccc ce cc|e:Ie(c|.
/ opo 8-100 c nicc cpcp(cIe|nc cue pe(mcnece nc LDL.
/ LDL ccnIm c mcic( pc(Ie cc cc|e:Ie(c| ci(cu|cnIe. 75 7 cc cuc| nc fc(mc ce
:Ie(e: ce cc|e:Ie(c|.
/ LDL ccpIccc vi(Iuc|menIe pc( Iccc: c: c|u|c: cc c(ccni:mc. mc: c: :eu:
p(incipci: c(ccc:-c|vc :cc c: c|cncu|c: :up(c-(enci: {ccm mcic( ccncenI(cccc ce
(ecepIc(e:). c: ccnccc: e c f|cccc. cI(cv: cc receptor poro os LDL.
C (ecepIc( pc(c c LDL umc c|iccp(cIe|nc cmnip(e:enIe nc nc::c c(ccni:mc cue
(eccnhece c /pc 8-100.
CcpIcccc cc LDL:
FecepIc(e: ce LDL ci:pc:Ic: nc (ecicc memL(cnc( (eve:Iicc pc( c|cI(inc
EnccciIc:e cc: (ecepIc(e: ce LDL - ve:|cu|c (eve:Iicc
Ve:|cu|c: (eve:Iicc: pe(cem c (eve:IimenIc - ce:pc|ime(izcccc cc c|cI(inc
FecepIc(e: ce:|iccm c: LDL
Fu:cc cc: ve:|cu|c: ce:(eve:Iicc: ccm |i:c::cmc:
FecepIc(e: cc: LDL :cc (ecic|ccc: pc(c c: memL(cnc:
Dec(cccccc cc: p(cIe|nc: LDL: hic(c|i:e cc: :Ie(e: ce cc|e:Ie(c|.
Fecu|cccc cc enccciIc:e cc ccmp|exc (ecepIc(-LDL
Cc|e:Ie(c| inI(cce|u|c( :up(ime c ccIiviccce ce hic(cximeIi|c|uIc(i|-Cc/
(ecuIc:e {feecLcck neccIivc)
Cc|e:Ie(c| inI(cce|u|c( e:Iimu|c c ccIiviccce cc cci|c-Cc/-cc|e:Ie(c|
cci|I(cn:fe(c:e.
Cc|e:Ie(c| iniLe c :|nIe:e cc: (ecepIc(e: cc: LDL.

/ meic vicc p|c:mcIicc cc LDL ceIe(minccc pe|c ccIiviccce cc (ecepIc( ce
LDL. N|vei: e|evccc: ce LDL imp|iccm Lcixc ccIiviccce cc :eu (ecepIc( e
cIe(c:c|e(c:e.
/ ccpIcccc ce cc|e:Ie(c| meciccc pc( (ecepIc(e: ce LDL ncc ce(c |ccn c|u|c:
{p(ecu(:c(c: cc cIe(c:c|e(c:e). ScmenIe c LDL cxicccc {|ipcpe(cxiccccc e
cxiccccc cc cpc 8-100) cIe(ccnicc.
N|vei: e|evccc: ce LDL-cc|e:Ie(c| cumenIcm c hipcIe:e cc LDL cxicccc - e:Ic c
(e|cccc enI(e hipe(cc|e:Ie(c|mic e cIe(ccne:e.

71

b. Ironsporte keverso de Co|estero|- HDL. $|stemo do Apo A-I

/ opo A-I cnIi-cIe(ccnicc.
/ HDL nc:ce nc: Iecicc: pc( ccc|e:cncic ce |os|o|p|dos ccm opo A-I {:inIeIizccc
nc f|cccc e inIe:Iinc). opo A-II. opo A-IV e opo C.
HDL nc:cenIe" cu ci:ccice" {HDL1) :cc heIe(ccnec: em Icmcnhc. ncc ccnIm
cc|e:Ie(c|
Cucncc HDL1 pc::c c ccnIe( e:Ie: ||picc:. c :eu inIe(ic( c((eccncc c HDL. ccncc-
|he um c:pecIc e:f(icc
Cucncc inIe(ccem ccm c|u|c: cc c(ccni:mc. c: HDL1 ccpIcm mc|cu|c:
excecenIe: ce cc|e:Ie(c| pc::cncc c HDL3.
/ HDL3 (icc em :Ie(e: ce cc|e:Ie(c| c(ccc: c LC/I. cue (eIi(c cc|e:Ie(c| |iv(e cc
:upe(f|cie cc HDL - |iLe(Icncc e:pccc pc(c c ccpIcccc ce mci: mc|cu|c: ce
cc|e:Ie(c| cc: Iecicc:.
/ HDL3 I(ccc :Ie(e: ce cc|e:Ie(c| pc( I(icci|c|ice(ci: cc: VLDL. ccm c cux||ic cc:
p(cIe|nc: ce I(cn:fe(ncic.
En(icuecicc ccm IG. ccnve(Ie-:e em HDL2 cue pcce Ie( cuc: vic: meIcLc|icc::
1. ccpIccc pe|c f|cccc. pccencc :e( cec(ccccc
2. |iLe(Iccc nc fc(mc ce HDL3 ce ncvc {mecicnIe (emcccc. nc :inu:cice
hepcIicc. cc: I(icci|c|ice(ci:)

kemoo do Co|esfero| dos Ce|u|os
Sc|cc cc cc|e:Ie(c| cc: c|u|c:
Mecon|smos:
1. Difu:cc. em cue c cc|e:Ie(c| cifunce cc memL(cnc p|c:mcIicc pc(c c HDL:
2. I(cn:pc(Ie meciccc pc( /pcp(cIe|nc:. em cue c HDL inIe(cce ccm (ecepIc(e: ce
memL(cnc pc(c cpcp(cIe|nc: e (eIi(c c cc|e:Ie(c| cc memL(cnc.

/ cifu:cc cc cc|e:Ie(c| cc c|u|c pc(c c HDL :c ccc((e cucncc exi:Ie um c(ccienIe
ce ccncenI(cccc enI(e c :upe(f|cie ce|u|c( e HDL (ecepIc(c cc cc|e:Ie(c|.
72

Cucncc c HDL e:Ic :cIu(ccc ce cc|e:Ie(c|. c cifu:cc ccnIinuc :cmenIe :e hcuve(
LC/I {|eciIinc cc|e:Ie(c| cci|-I(cn:fe(c:e). cue mccificc c: p(cp(ieccce: cc HDL.
cumenIcncc c :uc ccpcciccce ce (emcve( c cc|e:Ie(c| cc: c|u|c:.
/ LC/I e:Ie(ificc c cc|e:Ie(c| p(e:enIe nc :upe(f|cie cc |ipcp(cIe|nc. :encc c :Ie(
ce:|ccccc cc pe(ife(ic pc(c c nc|ec cc HDL.
De::e mccc. c: |os|o|p|dos {|eciIinc) ficcm |iv(e: nc pe(ife(ic pc(c incc(pc(c( mci:
cc|e:Ie(c| |iv(e.
/ (eIi(ccc ce cc|e:Ie(c| cc: c|u|c: IcmLm cepencenIe cc Apo A-I. cue :e
ci::ccic cc HDL e :e cp(cximc cc :upe(f|cie ce|u|c(. (eIcmcncc c HDL cpc: c
ccpIcccc cc cc|e:Ie(c|.
/ /pc /-l ncc cpenc: (emcve cc|e:Ie(c| e fc:fc||picc: {|eciIinc) ci(ecIcmenIe cc
memL(cnc p|c:mcIicc. mc: IcmLm e:Iimu|c c mcLi|izcccc cc :Icck ce cc|e:Ie(c|
cue e:Ic ci:pcn|ve| pc(c c e:Ie(ificcccc meciccc pe|c /C/I {/ci| Cc|e:Ie(c| /ci|-
I(cn:fe(c:e). umc enzimc |ccc|izccc nc (eI|cu|c enccp|c:mcIicc.
l:Ic eviIc c ccmu|c inI(cce|u|c( ce cc|e:Ie(c|. me:mc nc: mcc(cfccc: e c|u|c:
e:pumc:c:.
/::im. c HDL (cpiccmenIe cep|eIc c: :Icck: ce|u|c(e: ce cc|e:Ie(c|.
/ Apo A-I. A-II, Apo A-IV e Apo E fczem c me:mc efeiIc. pc(m Im meIcce cc
pcIncic cc /pc /-l.
/ inIe(ccccc HDL-c|u|c meciccc pe|c /pc /-l mcLi|izc c cc|e:Ie(c| IcnIc cc
memL(cnc cucnIc cc :Icck inI(cce|u|c(.
/ CEHI {cc|e:Ie(i|-e:Ie( hic(c|c:e neuI(c) hic(c|i:c c: :Ie(e: ce cc|e:Ie(c| mc: e::e
cc|e:Ie(c| ncc ficc ci:pcn|ve| pc(c (e-e:Ie(ificcccc pe|c /C/I. :encc enIcc
fcci|menIe (emcvicc cc c|u|c.
C cc|e:Ie(c| ci(cu|c ccnIinucmenIe enI(e c (eI|cu|c enccp|c:mcIicc e c memL(cnc
p|c:mcIicc. pc::cncc cnIe: pe|c ccmp|exc ce Gc|ci.
/: c|u|c: ccumu|cm mci: {cu menc:) cc|e:Ie(c| ce ccc(cc ccm c :uc
nece::iccce ce :inIeIizc( memL(cnc:. Cucncc c memL(cnc ficc :cIu(ccc ccm
cc|e:Ie(c| c exce::c ci(icicc cc (eI|cu|c enccp|c:mcIicc. :encc cuc(cccc :cL c
fc(mc ce :Ie(e: ce cc|e:Ie(c|.

Meccnicc cc ccpIcccc cc cc|e:Ie(c|
/ /pc /-l ci::ccic-:e cc :upe(f|cie cc HDL ci(cu|cnIe. mic(cncc pc(c c inIe(:I|cic
ce|u|c(. cnce :e |icc cc cc|e:Ie(c| |iv(e.
/ (ecuccc nc ccncenI(cccc |ccc| ce cc|e:Ie(c| pe(miIe cue e:Ie :e cifuncc cc
memL(cnc p|c:mcIicc pc(c c inIe(:I|cic. c fcvc( cc :eu c(ccienIe ce
ccncenI(cccc.
/ /pc /-l (ecm :inIeIizccc :ec(eIccc pe|c f|cccc ccmc cpc|ipcp(cIe|nc pcL(e
{cu ce:p(cvicc) em ||picc:.
Fc(Ie ce:Ic /pc /-l inIe(cce ccm (ecepIc(e: cc p(cp(ic f|cccc. :encc ccnve(Iicc
em HDL1. mc: c|cumc: mc|cu|c: ci(cu|cm pc(c Iecicc: exI(c-hepcIicc:.
ccumu|cncc-:e nc ||cuicc inIe(:Iicic|.
/I(cv: ce inIe(cccce: (eve(:|vei: ccm (ecepIc(e: ce memL(cnc. e::c /pc /-l
(emcve c cc|e:Ie(c| |iv(e e fc:fc||picc:. ccnve(Iencc-:e p(imei(cmenIe em F( -
HDL. e pc:Ie(ic(menIe em HDL1.
/ e:Ic c|Iu(c cc: cccnIecimenIc:. ccmeccm-:e c fc(mc( c: h|ice:. ce mccc cue
c
HDL1 vci pe(cencc c :uc ccpcciccce ce inIe(cci( ccm c: (ecepIc(e: ce :upe(f|cie
ce|u|c(.
E:Ic: pc(I|cu|c: enI(cm. c :ecui(. nc cc:ccIc ce cmccu(ecimenIc.
/ LC/I pc::c c e:Ie(ificc( c cc|e:Ie(c| |iv(e. ccn:I(uincc c nc|ec |ipicicc cc: :Ie(e:
cc cc|e:Ie(c| cc HDL3.
/|cun: ce:Ie: :Ie(e: :cc enIcc I(cn:fe(icc: pc(c c: VLDL e LDL pc( ccccc cc CEIF
{p(cIe|nc ce I(cn:fe(ncic ce :Ie(e: cc cc|e:Ie(c|). CuI(c: :cc I(cn:fe(icc: pc(c c
f|cccc e Iecicc: cue :inIeIizcm hc(mcnc: e:Ie(cice:.
73

/ (emcccc ce:Ie: nc|ec: |ip|cicc: cefc(mc c pc(I|cu|c. c cue pe(miIe c
ci::ccicccc cc: /pc /-l |ccc|izccc: nc pe(ife(ic. ce(cncc ncvc: p(ecu(:c(e: pc(c c
vic ce (emcccc cc cc|e:Ie(c|.
De:Ie mccc. c /pc /-l u:ccc m|Iip|c: veze: pc(c mcLi|izc(

74

.
.. 8
88|
||o
oos
sss
ss
n
nnt
tte
ees
sse
ee G
GG|
|||
||c
cc|
||d
dd|
||c
cco
oo

/: vic: cncLc|icc:. cue u:cm ene(cic cu|micc nc fc(mc ce /IF e N/DH cu N/DFH
pc(c :inIeIizc( ccmpcnenIe: ce|u|c(e: c pc(Ii( ce pe(cu(::c(e: mc|ecu|c(e:. :cc
ce(c|menIe (ecuIc(c:.

1. G||coneognese
G||coneognese: vic cue |evc c fc(mcccc ce cc(Lchic(cIc: c pc(Ii( ce pe(cu(::c(e:
ncc c|icicicc: e ccicc |ccIicc. Ccc((e nc F|cccc e Fim. em :iIuccce: ce fcme
p(c|cncccc.
Loco|: ciIc:c|. Ic| ccmc c c|icc|i:e.
/: vic: ncc :cc icnIicc:. pci: c: 3 (ecccce: i((eve(:|vei: cc c|icc:e impecem cue c
necc|icccne:e :ejc umc :imp|e: (eve(:cc cc p(cce::c. Nc c|iccneccne:e e:Ic:
eIcpc: :cc u|I(cpc::ccc: pc( um ccnjunIc c pc(Ie ce enzimc:. ccIc|izcncc (ecccce:
cue :cc :uficienIemenIe exe(ccnicc: pc(c :e(em efecIivcmenIe i((eve(:|vei: nc :enIicc
cc :|nIe:e ce c|icc:e.
Nc c|uccneccne:e. cccc um cc: pc::c: i((eve(:|vei: cc c|icc|i:e. :uL:IiIu|cc pc(
(ecccce: Ie(mccincmiccmenIe fcvc(cvei:.

Feccccc G|icc|i:e G|iccneccne:e
G|icc:e G|icc:e -F Hexccinc:e G|icc:e -fc:fcIc:e
F(uIc:e -F F(uIc:e 1. 8i:F Fc:fcf(uIccinc:e F(uIc:e 1.-Li:fc:fcIc:e
Fc:fcenc|pi(uvcIc pi(uvcIc Fi(uvcIc cinc:e Fi(uvcIc-cc(Lcxi|c:e
Fc:fcenc|pi(uvcIc-cc(Lcxicinc:e

$ubstrotos do Neog||cognese
/mincccicc: c|icccnicc: cu c|iccfc(mccc(e:: /mincccicc: cue pc( ce:cmincccc cu
I(cn:cmincccc c(icincm ccicc pi(vicc cu inIe(mecic(ic: cc Cic|c ce K(eL:.
Ac|cc c-cetcg|utc||cc: ccicc c|uIcmicc. c|uIcminc. hi:Iicinc. c(cininc e p(c|inc:
Ac|cc Cxc|ccct|cc: ccicc c:pc(Iicc e c:pc(ccinc:
Ac|cc F||uv|cc: c|cninc. I(ipIcfcnc. :e(inc. I(ecninc. ci:Ie|nc e c|icinc:
Succ|n||-CcA: I(ecninc. vc|inc. i:c|eucinc e meIicninc:
Ac|cc Func||cc: Ii(c:inc e feni|c|cninc.

Lipicc:
G||cero|:
um cc: p(ccuIc: cc meIcLc|i:mc cc Iecicc ccipc:c e :c c: Iecicc: cue
pc::uem c enzimc ccIivccc(c. c G|ice(c|cinc:e. pccem uIi|izc-lc.
E:Ic (ecue( /IF e ccIc|i:c c ccnve(:cc ce G|ice(c| c G|ice(c|-3-fc:fcIc.
E:Ie vci :e( cxicccc c Di-hic(cxicceIcnc fc:fcIc pe|c N/D
+
. nc p(e:encc cc
G|ice(c|-3-F-ce:ic(ccenc:e.
E:Ic enzimc enccnI(ccc enI(e cuI(c: Iecicc:. e:pecic|menIe nc ficccc e nc: (in:.
c|dos Gordos:
C: ccicc: cc(cc: ccm um nme(c pc( ce cc(Lcnc: c(icincm /ceIi|-Cc/.
Fc( cuI(c |ccc. c /ceIi|-Cc/ ccIivc c Fi(uvcIc-cc(Lcxi|c:e e iniLe c Fi(uvcIc-
ce:ic(ccenc:e.
C: ccicc: cc(cc: ccm um nme(c impc( ce cc(Lcnc:. pc(c c|m ce c(icinc(em
/ceIi|-Cc/. IcmLm c(icincm F(cp(icni|-Cc/. cue cepci: :e I(cn:fc(mc em
Succini|-Cc/.
Nctc: c /ceIi|-Cc/ ncc pcce :e( u:ccc ccmc pe(cu(:c( c|iccneccnicc. vi:Ic cue c
(eccccc cc pi(uvcIc ce:ic(ccenc:e i((eve(:|ve|.
CuI(c: cccc(e:: f(uIc:e. c cc|ccIc:e e c mcnc:e.
1. Converso do p|ruvoto o |os|oeno|p|ruvoto

75

/ :|nIe:e ce |os|oeno|p|ruvoto c pc(Ii( cc p|ruvoto c mci: exicenIe em Ie(mc:
ene(cIicc:. pc( Ie( um AG bostonte pos|t|vo.
Fc(c u|I(cpc::c( c Lc((ei(c Ie(mccincmicc. e:Ic (eccccc vci :e( cccp|ccc c umc
descorbox||oo. umc e:I(cIcic u:ccc f(ecuenIemenIe pe|c c|u|c pc(c impe|i(
um ecui||L(ic nc :enIicc cc fc(mcccc ce p(ccuIc:.
C pi(uvcIc. Ic| ccmc. c fc:fcenc|pi(uvcIc Iem 3 cc(Lcnc:. i:Ic imp|icc um
corbox||oo p(vic (ecue(encc ene(cic {ci:pncic ce /IF)
C pi(uvcIc I(cn:pc(Iccc pc(c c miIcccnc(ic cu ce(ccc c pc(Ii( cc o|on|no. nc
miIcccnc(ic. pc( I(cn:cmincccc.
Fc( ccccc cc pi(uvcIc cc(Lcxi|c:e. c pi(uvcIc ccnve(Iicc c cxc|ccceIcIc.
E:Ic (eccccc cncp|e(cIicc. i:Ic . (epce c: inIe(mecic(ic: cc cic|c ce K(eL:.

F|ruvoto + HCO3
-
+ AIF Oxo|oocetoto + ADF + F|

Enzimc miIcccnc(ic|
Ccnve(Ie c pi(uvcIc c cxc|ccceIcIc
Fecue( LicIinc cue I(cn:pc(Ic CC2 {(eIi(ccc cc HCC3
-
) :encc c c(upc p(c:IIicc
cc enzimc {LicIinc-enzimc LicIinc-enzimc-CC2)
c 1 enzimc (ecu|ccc(c cc vic c|icc||Iicc. :encc mccu|ccc ccvc|enIemenIe e
c|c:I(iccmenIe.
Depence cc ccIivcccc pe|c /ceIi|-Cc/ {cu cuI(c mc|cu|c cci|-Cc/)
Fecue( Mn
2+
cu Mc
2+

C cxc|ccceIcIc fc(mccc c pc(Ii( cc pi(uvcIc pcce Ie( cci: ce:Iinc: pc::|vei:: cu
enI(c nc cic|c ce K(eL: cu vci fc(mc( fc:fcenc|pi(uvcIc. cepencencc cc: n|vei: ce
/IF
Nc cc:c cc fc(mcccc ce fc:fcenc|pi(uvcIc. c cxc|ccceIcIc Iem cue cI(cve::c( c
memL(cnc nc fc(mc ce cuI(c ccmpc:Ic {mc|cIc cu c:pc(IcIc). pci: ncc
ccn:ecue pc::c-|c.
Se hcuve( N/DH :uficienIe nc ciIcp|c:mc c cxc|ccceIcIc I(cn:cminccc c
/:pc(IcIc.
Cc:c ccnI(c(ic. (ecuzicc c mc|cIc pe|c mc|cIc ce:ic(ccenc:e miIcccnc(ic|. ccm
cxiccccc ce N/DH.

+
L-mo|oto + NAD
+

Mc|cIc ce:ic(ccenc:e

C mc|cIc :ci cc miIcccnc(ic pe|c I(cn:pc(Iccc( mo|oto-u-cetog|utoroto cc
memL(cnc miIcccnc(ic| inIe(nc
C mc|cIc :e(ve ccmc I(cn:pc(Iccc( ce ecuivc|enIe: (ecuIc(e: cc miIcccnc(ic
pc(c c ciIc:c| c::ecu(cncc c I(cn:fc(mcccc ce 1.3-Li:fc:fcc|ice(cIc c
c|ice(c|ceicc 3-F.
Nc ciIc:c| (ecxicccc c cxc|ccceIcIc. ccm p(ccuccc ce N/DH ciIc:c|icc.

Mo|oto + NAD
+
+

mc|cIc ce:ic(ccenc:e

C cxc|ccceIcIc enIcc ccnve(Iicc c fc:fcenc|pi(uvcIc pe|c fc:fcenc|pi(uvcIc-
cc(Lcxinc:e.
umc (eccccc Mc
2+
cepencenIe cue (ecue( GIF ccmc cccc( ce fc:fcIc.

Oxo|oocetoto + GIF Fos|oeno|p|ruvoto + CO2 + GDF
Fc:fcenc|pi(uvcIc cc(Lcxicinc:e

76

Ecucccc ce(c|:
F|ruvoto + AIF + GIF + HCO3
-
Fos|oeno|p|ruvoto + ADF + F| + GDF + CO2
AG0 = 0.5 Kj/mc|

Fc(c fc(mc( umc mc|cu|c ce pi(uvcIc c fc:fcenc|pi(uvcIc :cc nece::c(ic: 2
ecuivc|enIe: ce fc:fcIc {1 /IF e 1 GIF). encucnIc cue nc c|icc|i:e. pc(c ccnve(Ie( c
fc:fcenc|pi(uvcIc em pi(uvcIc. cpenc: 1 /IF fc(mccc.

LccIcIc ccmc pe(cu(:c( c|iccneccnicc

C |octoto, p(cvenienIe cc: e(iI(cciIc: cu m:cu|c: em :iIuccce: cnce(cLic:.
IcmLm pcce :e( pe(cu(:c( c|iccneccnicc.
/ ccnve(:cc ce |octoto p|ruvoto nc ciIc:c| cc: hepcIcciIc: c(icinc N/DH. e c
expc(Icccc ce ecuivc|enIe: (ecuIc(e: {ccmc mc|cIc) cc miIcccnc(ic ncc
nece::c(ic.
Depci: cc pi(uvcIc p(ccuzicc pe|c (eccccc cc |ccIcIc ce:ic(ccenc:e. :e(
I(cn:pc(Iccc pc(c c miIcccnc(ic. ccnve(Iicc em cxc|ccceIcIc pe|c pi(uvcIc
cc(Lcxi|c:e.
E:Ie cxc|ccceIcIc. ccnIucc. ci(ecIcmenIe ccnve(Iicc em |os|oeno|p|ruvoto pc(
umc i:cenzimc miIcccnc(ic| cc fc:fcenc|pi(uvcIc cc(Lcxicinc:e. e c
fc:fcenc|pi(uvcIc I(cn:pc(Iccc pc(c fc(c cc miIcccnc(ic pc(c ccnIinuc( c vic
c|iccneccnicc.

77

2. Converso do |rutose 1,-b|s|os|oto em |rutose -|os|oto

/: (ecccce: ccIc|izccc: pe|c fc:fcf(uIccinc:e l e pe|c hexccinc:e :cc :uL:IiIu|cc:
nc c|iccneccne:e pc( (ecccce: hic(c||Iicc:.
Ne:Ie pcnIc. em vez ce fc:fc(i|c( /DF c /IF {c inve(:c cc c|icc|i:e. mc:
ce:fcvc(ecicc Ie(mccincmiccmenIe em ccncicce: fi:ic|ccicc:). ccc((e c
|iLe(Icccc cc fc:fcIc pc( hic(c|i:e.
E:Ic c enzimc chcve. nc :enIicc cue c :uc p(e:encc ceIe(minc :e um Iecicc
cu ncc ccpcz ce (e::inIeIizc( c c|icccnic cu c|icc:e c pc(Ii( cc ccicc pi(vicc e
cc: I(ic:e-fc:fcIc.
Ju|cc-:e cue ncc exi:Ie nc: m:cu|c: cc(c|ccc: e |i:c.

/ f(uIc:e 1.-Li:fc:fcIc:e {Mc
2+
cepencenIe) p(cmcve c hic(c|i:e i((eve(:|ve| cc
fc:fcIc C1.

Frutose 1,-b|s|os|oto + H2O Frutose -|os|oto + F|
F(uIc:e 1.-Li:fc:fcIc:e

3. Converso de g||cose -|os|oto o g||cose ||vre

C inve(:c cc (eccccc cc hexccinc:e (ecue(ic I(c:nfe(ncic ce 1 c(upc fc:fc(i| cc
c|icc:e -F pc(c c /DF. fc(mcncc /IF. (eccccc ene(ciccmenIe ncc fcvc(cve|.
/ (eccccc ccIc|izccc pe|c c|icc:e -fc:fcIc:e ncc (ecue( :|nIe:e ce /IF. mc: c
:imp|e: hic(c|i:e ce um :Ie( fc:IcIc:

G||cose -|os|oto + H2O G||cose + F|
G|icc:e -fc:fcIc:e

G|icc:e -fc:fcIc:e
Enzimc Mc
2+
cepencenIe
Exi:Ie cpenc: nc f|cccc. (im e c|u|c: cc epiI|ic inIe:Iinc|. c cue |he: pe(miIe
fc(nece( c|icc:e cc (e:Ic cc c(ccni:mc.
EnccnI(c-:e nc memL(cnc cc FeI|cu|c enccp|c:mcIicc.

keoco g|obo| do g||coneognese
2p|ruvoto + 4AIF + 2 GIF + 2NADH + 4H2O G||cose + 4ADF + 2GDF + F| + 2NAD
+
+ 2H
+

keoco g|obo| do g||co||se
G||cose + 2ADF + 2F| + 2NAD
+
2p|ruvoto + 2AIF + 2NADH + 2H
+
+ 2H2O

8o|ono energt|co
/ :|nIe:e ce c|icc:e (ecue( /IF. :encc e:Ie cc:Ic ce ene(cic nece::c(ic pc(c
c::ecu(c( c i((eve(:iLi|iccce cc c|iccneccne:e.
C /IF ce cue c: c|u|c: hepcIicc: nece::iIcm pc(c c :|nIe:e ce c|icc:e
fc(necicc em c(cnce pc(Ie pe|c cxiccccc cc: ccicc: cc(cc:.
/: ccncicce: meIcLc|icc: :cL c: cuci: c f|cccc ceve :inIeIizc( c c|icc:e.
fcvc(ecem. ce(c|menIe. um cumenIc nc ci:pcniLi|iccce ce ccicc: cc(cc: nc
:cncue p(cvenienIe: cc: (e:e(vc: ccipc:c:.

78

kegu|oo
/ c|icc|i:e e c necc|icccne:e :cc ccnI(c|ccc: pe|c: meccni:mc: pc(c cue :ejc
pc::|ve| cue cpenc: umc cc: vic: funcicne. Iencc em ccnIc cue ccc((em em
ccmpc(IimenIc: ce|u|c(e: icuci: {excepIc nc 1 (eccccc ce c|iccneccne:e)
Cccc umc cc: enzimc: (ecu|ccc(c: e:Ic :ujeiIc c 2 Iipc: ce (ecu|cccc: c|c:I(icc
e :|nIe:e p(cIeicc {e:Ic cepencenIe ce cive(:c: hc(mcnc: ccmc in:u|inc.
c|ucccinc. epinef(inc)
C: efecIc(e: e:Iimu|cm c c|iccneccne:e Iencem c (ep(imi( c c|icc|i:e e vice-ve(:c
cc inf|uencic(em c ccIiviccce e cucnIiccce: cccue|c: enzimc:.

C pi(uvcIc pcce :e( ccnve(Iicc c /ceIi|-Cc/ {ccmp|exc cc ce:ic(ccenc:e
pi(uvicc) cu c cxc|ccceIcIc {pi(uvcIc cc(Lcxi|c:e).
/ pi(uvcIc-cinc:e iniLicc pe|c AIF e o|on|no. cc ccnI(c(ic cc cc(Lcxicinc:e. cue
iniLicc pe|c ADF e e:Iimu|ccc pe|c AIF e ocet||-CoA. Iencc e:Ie um efeiIc
c|c:I(icc neccIivc nc pi(uvcIc ce:ic(ccenc:e. CU SEJ/:
Cucncc c: nece::iccce: ene(cIicc: cc c|u|c e:Icc :cIi:feiIc:. c fc:fc(i|cccc
cxiccIivc I(cvc. c N/DH ccumu|c e iniLe c Cic|c ce K(eL:
/cumu|c-:e /ceIi|-Cc/ cue iniLe c ccmp|exc cc ce:ic(ccenc:e pi(vicc.
(ecuzincc c fc(mcccc ce cceIi|-Cc/ c pc(Ii( cc pi(uvcIc
E:Iimu|c c c|iccneccne:e pe|c ccIivcccc cc pi(uvcIc cc(Lcxi|c:e. fczencc
ccm cue c exce::c ce pi(uvcIc :ejc ccnve(Iicc em c|icc:e.

/ e|evcccc ce ocet||-CoA e cc ccIiviccce cc Fi(uvcIc-Cc(Lcxi|c:e (e:u|Ic numc
mcic( :|nIe:e ce c|do ctr|co. um efecIc( neccIivc cc fc:fcf(uIccinc:e e pc:iIivc
cc f(uIc:e l.-Li:fc:fcIc:e.
/ iniLiccc ce:Ic enzimc p(cvccc umc ciminuiccc nc ccncenI(cccc ce Frutose-I,-
b|s|os|oto. um ccIivccc( cc pi(uvcIc-cinc:e.
79

/::im. (ecuz-:e c f|uxc cc c|do |os|oeno|p|rv|co c ccicc pi(vicc. pe|c ccccc cc
pi(uvcIc-cinc:e. cumenIcncc pc( cuI(c |ccc c: e:fc(cc: ccmLinccc: cc pi(uvcIc-
cc(Lcxi|c:e e cc fc:fcenc|pi(uvcIc-cc(Lcxicinc:e. nc ccnve(:cc cc ccicc pi(vicc
c ccicc fc:fcenc|pi(uvicc.

/ cxiccccc cc: ccicc: cc(cc: fc(nece /IF pc(c c p(cce::c e p(cmcve c :|nIe:e
ce c|icc:e. cI(cv: cc cumenIc cc ccncenI(cccc nc e:Iccc e:Iccicnc(ic ce
/ceIi|-Cc/ miIcccnc(ic|.

/ fc:fcf(uIccinc:e e:Iimu|ccc pe|c AMF/ADF e iniLicc pe|c AIF e c|do ctr|co.
E:Ie: Im umc ccccc cpc:Ic :cL(e c f(uIc:e-|.-Li:fc:fcIc:e.
/::im. cucncc hc um bo|xo nve| energt|co. incicccc pc( ccncenI(ccce: Lcixc:
ce /IF e e|evccc: ce /MF. c f(uIc:e-|.-Li:fc:fcIc:e iniLicc e c g||co||se
|ovorec|do.
Fc( cuI(c |ccc. cucncc c nve| energt|co e|evodo. incicccc pc( e|evccc:
ccncenI(ccce: ce /IF. f(uIc:e-|.-Li:fc:fcIc:e ccIivccc encucnIc c
fc:fcf(uIccinc:e iniLicc. |ovorecendo o neog||cognese.

Em :umc. um oumento dos nve|s de AIF. ccm c ccn:ecuenIe decrsc|mo dos
nve|s de AMF. fcvc(ece c necc|icccne:e cI(cv: cc iniLiccc cc fc:fcf(uIccinc:e
e cc pi(uvcIc-cinc:e. e cc ccIivcccc cc f(uIc:e-l.-Li:fc:fcIc:e.

C pcpe| e:pecic| cc |godo nc mcnuIenccc cc n|ve| ce c|icc:e nc :cncue (ecue(
meccni:mc: (ecu|cIc(ic: ccicicnci: pc(c ccc(cenc( c p(ccuccc e ccn:umc ce
c|icc:e.
Cucncc c: n|vei: ce c|icc:e nc :cncue ciminuem c hc(mcnc g||cog|no :inc|izc cc
f|cccc pc(c p(ccuzi( e |iLe(Ic( mci: c|icc:e.
/ (ecu|cccc hc(mcnc| cc c|icc|i:e e cc c|iccneccne:e nc f|cccc meciccc pe|c
|rutose 2,-b|s|os|oto. um efecIc( c|c:I(icc pc(c c fc:fcf(uIccinc:e l e c F(uIc:e 2.-
Li:fc:fcIc:e.
/ f(uIc:e 2.-8i:fc:fcIc :inIeIizccc pe|c fc:fcf(uIccinc:e 2 e fc:fc(c|i:e pe|c f(uIc:e
2.-Li:fc:fcIc:e.
Cucncc c Frutose 2,-b|s|os|oto :e |icc cc :|Iic c|c:I(icc nc fc:fcf(uIccinc:e l.
cumenIc c cfiniccce cc enzimc pc(c c |rutose -|os|oto e (ecuz c cfiniccce pc(c
c: iniLicc(e: c|c:I(icc:. /IF e ciI(cIc.
/ f(uIc:e 2.-Li:fc:fcIc ccIivc enIcc c fc:fcf(uIccinc:e e e:Iimu|c c c|icc|i:e nc
f|cccc.
/c me:mc Iempc. c f(uIc:e 2.-Li:fc:fcc|ice(cIc iniLe c f(uIc:e 1.-Li:fc:fcIc:e.
cL(cnccncc c c|iccneccne:e.

/: hc(mcnc: ccnI(c|cm c ve|cciccce cc: enzimc:. mc: IcmLm c :uc :|nIe:e.
c|Ie(cncc-:e. e::encic|menIe. c ve|cciccce ce I(cn:c(iccc. Fc( exemp|c. c in:u|inc.
e:Iimu|c c :inIe:e ce fc:fcf(uIccinc:e e cc pi(uvcIc-cinc:e. encucnIc c c|ucccinc
Im umc ccccc cpc:Ic.

80

C|c|os |te|s
Em cccc um cc: 3 pcnIc: cnce c: (ecccce: c|icc||Iicc: Im umc vic
c|iccneccnicc c|Ie(ncIivc. c cpe(cccc :imu|Icncic ce cmLc: c: vic: :e(ic um
ce:pe(c|cic.
E:Ie p(cce::c ncc eccncmicc e ccc((e ccm pe(cc ce ene(cic
Em ci(cun:Icncic: nc(mci: e:Ie cic|c ncc ccc((e c umc e:cc|c :icnificcIivc. vi:Ic
cue p(evenicc pe|c: meccni:mc: (ecu|cIc(ic:
CcnIucc. e:Ie cic|c ccc((e pc( veze: fi:ic|cciccmenIe pc(c mcnIe( cu cumenIc( c
Iempe(cIu(c cc cc(pc.

C|c|o do G||cose-Loctoto ou c|c|o de Cor|
C |ccIcIc I(cn:pc(Iccc cc m:cu|c e:cue|Iicc e e(iI(cciIc. cnce ccc((e
fe(menIcccc |ccIicc. pc(c c f|cccc e (in:. cnce cecc((e c ccnve(:cc ce pi(uvcIc c
c|icc:e.
E:Ie ccnjunIc ce I(cn:fc(mccce: {cic|c ce Cc(i) envc|ve I(cn:fe(ncic: cc |ccIcIc
{mu:cu|c f|cccc) pc( I(ccc ccm mcIe(ic| ene(cIicc {c|icc:e) em :enIicc inve(:c
{f|cccc mu:cu|c).

C|c|o do G||cose-A|on|no
/ c|cninc. p(cvenienIe em c(cnce cucnIiccce cc mu:cu|c em :iIuccce: ce jejum.
pcce :e( I(cn:fc(mccc em c|icc:e pe|c f|cccc. cI(cv: cc cic|c cc G|icc:e-/|cninc
C cumenIc ce c|cninc em ci(cu|cccc (e:u|Ic cc ceccmpc:iccc cc: p(cIe|nc:
mu:uc|c(e: c pc( cc cmincccc cc pi(uvcIc fc(mccc nc c|icc|i:e.
Nc f|cccc. c c|cninc cc cece( c c(upc cminc pc(c c fc(mcccc cc u(eic.
I(cn:fc(mc-:e em pi(uvcIc
C pi(uvcIc. cI(cv: cc c|iccneccne:e. (ecene(c G|icc:e--F e c|icc:e |iv(e. pc(
:uc vez I(cn:fe(icc cc f|cccc pc(c c m:cu|c {e cuI(c: Iecicc: cc(enciccc:)

81

2. G||cognese
Ccc((e nc f|cccc e nc: mu:cu|c: e:cue|Iicc:
Nc |godo. c c|icccnic :e(ve ccmc um (e:e(vcIc(ic ce c|icc:e (cpiccmenIe
ccnve(I|ve| em c|icc:e :cncu|nec pc(c ci:I(iLuiccc pc(c cuI(c: Iecicc:.
Nc mscu|os c c|icc:e p(ccuzicc c pc(Ii( cc c|icccnic meIcLc|izccc cI(cv: cc
c|icc|i:e pc(c p(ccuzi( /IF

Cond|es |ovorve|s g||cognese:
Exce::c ce c|icc:e
Exce::c ce ccpcciccce: ene(cIicc: ce|u|c(e:
E:Iimu|cccc hc(mcnc| pe|c in:u|inc

Object|vos metobo||cos:
/(mczencmenIc ce umc fc(mc cce::|ve|
Eccncmic ce (e:e(vc ene(cIicc

Fercursor: c|icc:e -fc:fcIc c|icc:e 1-fc:fcIc

G||cose + AIF G||cose -Fos|oto
Hexccinc:e {cu c|icccinc:e)

G||cose -|os|oto G||cose 1-|os|oto
Fc:fcc|iccmuIc:e

/pc: c fc(mcccc ce c|icc:e 1-fc:fcIc. e:Ic ccnve(Iicc em UDF-c|icc:e {fc(mc
ccIivc cc c|icc:e).
/ UDF-c|icc:e efecIivc c incc(pc(cccc ce (e:|cuc: ce c|icc:e nc: pc|i::ccc(icc:
em fc(mcccc. c :uc unicc c cuI(c: mcnc::ccc(icc: cu I(cn:fc(mcccc em
ce(ivccc: ce hexc:e:.

G||cose 1-|os|oto + bIF bDF-g||cose + FF|
UDF-c|icc:e pi(cfc:fc(i|c:e

EmLc(c e:Ic (eccccc :ejc (eve(:|ve|. Ience c ccmp|eIc(-:e nc ci(ecccc cc :|nIe:e
cc UDF-c|icc:e. cevicc c (cpicc hic(c|i:e cc pi(cfc:fcIc em c(Icfc:fcIc pe|c
pi(cfc:fcIc:e inc(ccnicc {(eccccc Lc:IcnIe excene(cIicc).

Etopos do g||cognese
1 etopo: c(e:cimenIc |inec( cc: ccceic: cc c|icccnic pc( cciccc ce :uce::ivc ce
(e:|cuc: c|icc:i|c. unicc: enI(e :i pc( |icccce: u-1.4.
/ UDF-c|icc:e c I(cn:pc(Iccc( cc c(upc c|icc:i|c ccIivccc.
/ ccIivcccc cc c|icc:e (e:u|Ic cc e:Ie(ificcccc cc :eu C1 pe|c cifc:fcIc cc
mc|cu|c UDF.
C c(upc c|icc:i|c cc mc|cu|c ce UDF-c|icc:e I(cn:fe(icc pc(c c (e:|cuc Ie(minc|
cc c|icc:e p(e:enIe nc: exI(emiccce: ncc (ecuIc(c: cc ccceic ce cmi|c:e
{c|icccnic iniciccc()
c c|icccnic :inIc:e cue ccIc|izc c fc(mcccc cc |iccccc c|icc:|cicc enI(e c
c(upc CH cc C4 Ie(minc| cc ccceic e c C1 cc c(upc c|icc:i|c I(cn:pc(Iccc cc
UDF-c|icc:e.
Simu|IcnecmenIe ccm c fc(mcccc ce cccc |iccccc c|icc:icicc u-1.4 (emcvicc
umc mc|cu|c ce UDF.
/ c|icccnic :inIc:e ccIc|izc c (eccccc :e c pc|i::ccc(icc p(-exi:IenIe jc ccnIive(
mci: ce 4 (e:|cuc: ce c|icc:e.
82

Fc(Ie cc ene(cic |iv(e cc fc:fc(i|cccc cc c|icc:e ccn:e(vccc nc |iccccc
c|icc:icicc u-1.4.

2etopo: (cmificcccc cc c|icccnic cI(cv: cc: |icccce: c|icc:icicc: u-1.
Cucncc c ccceic cIince 8 c 12 (e:|cuc: ce c|icc:e inIe(vm umc enzimc
(cmificcnIe. c c|icc:i| {1 ) I(cn:fe(c:e.
E:Ic enzimc. cpc: c c|ivccem ce umc |iccccc c|icc:icicc u-1.4. ccIc|izc c
I(cn:fe(ncic ce um Ie(minc| ccm cu 7 (e:|cuc: ce c|icc:e cc exI(emiccce ncc
(ecuIc(c ce um (cmc ce c|icccnic pc(c c c(upc hic(cxi|c C ce um (e:|cuc ce
c|icc:e numc pc:iccc mci: inIe(nc cc ccceic. c(icncc um ncvc (cmc.
Dccui (e:u|Ic c fc(mcccc ce umc |iccccc c|icc:icicc u-1. {enI(e c C1 cc p(imei(c
(e:|cuc ce c|icc:e cc f(ccmenIc I(cn:fe(icc e c C ce (e:|cuc ce c|icc:e cc
ccceic cue c (eceLe) e ccn:ecuenIemenIe umc zcnc ce (cmificcccc cc
c|icccnic. :epc(ccc pc(. pe|c menc:. 4 (e:icuc: ce c|icc:e ce cuI(c (cmificcccc
p(-exi:IenIe.
/ mc|cu|c Iem ccc(c cuc: exI(emiccce: ncc (ecuIc(c: e. pc(IcnIc. cuc: zcnc:
ce c(e:cimenIc.

Vontogens do rom|||coo do g||cogn|o:
/umenIc c :uc :c|uLi|iccce
/umenIc c cucnIiccce ce exI(emiccce: ncc (ecuIc(c: cnce ccIucm c
c|icccnic :inIc:e e. IcmLm. c c|icccnic fc:fc(i|c:e.

/ incc(pc(cccc ce mc|cu|c: ce c|icc:e -fc:f:Ic nc c|icccnic cu:Ic umc |iccccc
ce c|Ic ene(cic. nece::c(ic c (ecene(cccc cc UIF pe|c enzimc nuc|ec:icc-
cifc:fccinc:e:
bDF + AIF bIF + ADF
Nuc|ec:icc-cifc:fccinc:e

/ c|icccnic :inIeIc:e :c ccn:ecue ccicicnc( c|icc:e c ccceic: ce c|icccnic p(-
exi:IenIe: cu c||ne|. i:Ic . ncc ccpcz ce ccmecc( c :|nIe:e ce umc ncvc mc|cu|c
ce c|icccnic. / :|nIe:e cc c|icccnic iniciccc pe|c cciccc ce umc mc|cu|c ce
c|icc:e c um (e:|cuc ce Ii(c:inc ce umc p(cIe|nc cencminccc g||cogen|no.

keoces do sntese do g||cogn|o Enzimc:

G|icc:e -F G|icc:e 1-F Fc:fcc|iccmuIc:e
G|icc:e 1-F + UIF UDF-c|icc:e + FFi UDF-G|icc:e pi(cfc:fcIc:e
FFi + H20 2 Fi
UDF-c|ucc:e + {c|icccnic] n
{c|icccnic] n+1 + UDF
G|icccnic :inIc:e
UDF+/IF UIF+/DF Nuc|ec:icc cifc:fccinc:e
Scmc: G||cose--|os|oto + AIF + [G||cogn|o]n + H2O [G||cogn|o]n+1 + ADF + 2 F|

kegu|oo do metobo||smo do g||cogn|o
/ c|icccenc|i:e e c|icccne:e cecc((em :cL(e ccnI(c|c (icc(c:c. pci: cucncc umc
vic ccIivccc. cuI(c inccIivccc. cI(cv: ce cci: Iipc: ce ccnI(c|c: c|c:I(icc e
ccvc|enIe.
IcnIc c c|icccenc|i:e ccmc c c|icccne:e :cc (ecu|ccc: pc( enzimc: p(cp(ic:: c
c|icccnic fc:fc(i|c:e e c c|icccnic :inIc:e. (e:pecIivcmenIe.
/: cuc: enzimc: cp(e:enIcm-:e :cL cuc: fc(mc:. ccIivc e inccIivc.
83

4/IF
SinIc:e cinc:e
SinIc:e fc:fcIc:e
G|icccnic
SinIc:e c
G|icccnic
SinIc:e L
4/DF
G|icccnic
UDF-G|icc:e
4Fi 4H2C
UDF
C funcicncmenIc ce cccc umc cc: :ecuncic: cepence cc e:Iccc ce ccIivcccc
ce::c: (e:pecIivc: enzimc: (ecu|ccc(e:. :ujeIic: c mccu|cccc c|c:I(icc e
mccificcccc ccvc|enIe.

G|icccnic :inIc:e
ccn:IiIu|cc pc( 4 :uLuniccce: pc|ipepIicicc:.
/p(e:enIc-:e :cL cuc: fc(mc: pc::|vei::
G|icccnic :inIc:e D cu L {fc:fc(i|ccc. inccIivc)
G|icccnic :inIc:e l cu c {ce:fc:fc(i|ccc. ccIivc)
/ c|icccnic :inIc:e L fc:fc(i|ccc cc n|ve| ce um (e:|cuc ce :e(inc em cccc umc
cc: :uLuniccce:. :encc vi(Iuc|menIe inccIivc
Depence cc ccIivcccc pe|c c|icc:e -fc:fcIc. cue ccIuc ccmc mccu|ccc(
pc:iIivc. :encc c ccIivcccc L|ccueccc pe|c p(e:encc ce UDF.
/ fc(mc ce:fc:fc(i|ccc incepencenIe cc c|icc:e -fc:fcIc e. pc(IcnIc. ccIivc nc
:uc cu:ncic.
C Km cc c|icccnic :inIc:e pc(c c UDF-c|icc:e ciminui nc p(e:encc ce G|icc:e -
fc:fcIc.
/ ce:fc:fc(i|cccc cc c|icccnic-:inIc:e iniLicc pe|c c|icccnic. cue ccIuc ccmc
mccu|ccc( c|c:I(icc neccIivc.
/: cuc: fc(mc: cc c|icccnic :inIc:e :cc inIe(ccnve(Iicc: pe|c :inIc:e-cinc:e e
:inIc:e-fc:fcIc:e.
/ :inIc:e-fc:fcIc:e I(cn:fc(mc :inIc:e L nc c. pc( (emcccc cc: c(upc: fc:fc(i|c cc
|iccccc :Ie( ccm (e:|cuc: ce :e(inc cc enzimc. cue ce:Ie mccc ccIivccc.
/ enzimc inccIivccc pc( fc:fc(i|cccc. ccIc|i:ccc pe|c :inIc:e-cinc:e c cu:Ic ce
/IF.
/ :inIc:e cinc:e fcz pc(Ie ce um ccnjunIc ce enzimc: {p(cIe|nc:-cinc:e:) cue
ccIc|izcm e:Ie Iipc ce (eccccc IcmLm exi:Iincc :cL umc fc(mc ccIivc e
inccIivc.

/ (ecu|cccc c|c:I(icc cc meIcLc|i:mc cc c|icccnic c::ccic-:e cc: meccni:mc:
cue ccnI(c|cm c c|icc|i:e e cic|c ce K(eL:.
Em Ie(mc: ce(ci:. um cumenIc cc ci:pcniLi|iccce cc ccIiviccce ce|u|c( em c|icc:e
{:cL c fc(mc ce c|icc:e -fc:fcIc) cu cuI(c: nuI(ienIe: {pc( e|evcccc cc cc(cc
ene(cIicc) ccIivc c c|icccnic :inIc:e e iniLe c fc:fc(i|c:e. Dccui (e:u|Ic um
/ceni|cic|c:e l
/ceni|cic|c:e /
/MF
/MFc
F(cIe|nc cinc:e l
F(cIe|nc cinc:e /
G|icccnic
:inIc:e c
G|icccnic
:inIc:e L
Cc|cic
/c(enc|inc
G|ucccinc
84

cumenIc cc cepc:iIc cc c|icc:e ccmc c|icccnic. nc: m:cu|c: e:cue|Iicc: e
f|cccc.
/ c|icc:e cc |icc(-:e c fc:fc(i|c:e hepcIicc fcvc(ece c :uc inccIivcccc pe|c
fc:fcIc:e: e:Ic enzimc. cc ci::ccic(-:e cc fc:fc(i|c:e L ficc |iv(e pc(c ccIuc( nc
c|icccnic :inIc:e. ccIivcncc-c.
Cucncc ciminui c cc(cc ene(cIicc ce|u|c( cu Lcixc c c|icemic. e:Iimu|ccc c
fc:fc(i|c:e e iniLicc c :inIc:e: c|icccnic enIcc ceccmpc:Ic em c|icc:e 1-fc:fcIc
e cccui em c|icc:e -fc:fcIc. c :e( uIi|izccc nc :ecuncic c|icc||Iicc {cc mu:cu|c)
cu ccm ce:Iinc c ci(cu|cccc {pe|c f|cccc. cpc: ce:fc:fc(i|cccc)
C meccni:mc ce (ecu|cccc ccvc|enIe e:Ic cepencenIe ce e:I|mu|c: hc(mcnci:.
pe|c fc(mcccc ce c/MF e :uL:ecuenIe ccIivcccc cc p(cIe|nc cinc:e. /I(cv: ce
um cc:ccIc ce (ecccce: ce fc:fc(i|cccc. c :inc| hc(mcnc| cmp|iccc e:Ic cc
n|ve| cc: enzimc: (ecu|ccc(c: cc c|icccenc|i:e e c|icccne:e.
85

1
110
00.
.. H
HHo
oor
rrm
mmo
oon
nno
oos
ss

Hormonos: :uL:Icncic Liccu|micc :ec(ecccc em cucnIiccce: m|nimc: pe|c: c|u|c:
ce ceIe(minccc: Iecicc: cu fc(mccc c pc(Ii( ce pe(cu(:c(e: ci :inIeIizccc:. cue ci(cu|c
cc: f|uicc: Lic|ccicc: e ccIuc em |ccci: e:pec|ficc: ce fc(mc c p(ccuzi( umc (e:pc:Ic
ccecuccc e eficcz.

C|oss|||coo:
o. ko|o de oco
Hc(mcnc: enccc(inc:: hc(mcnc: cue p(cvm ce um Iecicc cu c|cncu|c e cue vicjcm
umc ci:Icncic ccn:ice(cve| em ci(cu|cccc cI cIici(em umc c|u|c-c|vc cue ccnIenhc
(ecepIc(e: e:pe|ficc: pc(c e::c hc(mcnc:. /cIivc c|u|c: ci:IcnIe:.
Hc(mcnc: pc(cc(inc:: p(ccuzicc: numc c|u|c e vicjcm umc ci:Icncic (e|cIivcmenIe
pecuenc cI inIe(cci(cm ccm c: (ecepIc(e: :iIuccc: nc: c|u|c: vizinhc:. /cIivc
c|u|c: vizinhc:.
Hc(mcnc: cuIcc(inc:: p(ccuzicc: pe|c me:mc c|u|c cue funcicnc ccmc c|vc pc(c
umc hc(mcnc. /cIivc c p(cp(ic c|u|c.
Hc(mcnc: mi:Ic:

b. Noturezo qum|co
Hc(mcnc: hic(c::c|vei: {hc(mcnc: pepIicicc:. p(cIcc|cncinc: e ce(ivccc: ce ccicc:
cminccc:): |iccm-:e c (ecepIc(e: cc memL(cnc ce|u|c(. (eve(:ive|menIe ccm
ccIivcccc ce um :ecuncc men:ccei(c inI(cce|u|c( {/MFc. GIFc. Cc
2+
) cue
ce:enccceic umc (e:pc:Ic (cpicc cc n|ve| cc ccIiviccce enzimcIicc.

Hc(mcnc: |ipc::c|vei: {e:Ie(cice:. eicc:cccice:): enI(cm nc c|u|c e |iccm-:e c
(ecepIc(e: inI(cce|u|c(e:. C ccmp|exc |iccncc-(ecepIc( |icc-:e cc /DN e ce:enccceic
um Iipc ce (e:pc:Ic mci: |enIc cc n|ve| cc exp(e::cc cenIicc pc( mccificcccc cc
:|nIe:e p(cIeicc.

Coscoto hormono|

Hc(mcnc: inIe(mecic(ic:
Hc(mcnc: inIe(mecic(ic:
lmpu|:c e|ecI(ccu|micc
lmpu|:c e|ecI(ccu|micc
E:Iimu|c inIe(nc cu cmLienIc|
SNC
Si:Iemc LimLicc
FiIuiIc(ic /nIe(ic(
HipcIc|cmc
G|cncu|c /|vc
U|Iimc hc(mcnc
Fe:pc:Ic hc(mcnc|
86

Fecu|cccc cc:ccIc hc(mcnc| {feecLcck)
FeecLcck |cncc: c hc(mcnc finc| |icc-:e c um (ecepIc( cc: c|u|c: cc piIuiIc(ic
cnIe(ic(. hipcIc|cmc e SNC. iniLincc c p(ccuccc pc( pc(Ie ce::c: cc|u|c: ce :inci:
hc(mcnci: inIe(mcic: cc cc:ccIc.
FeecLcck cu(Ic: c hc(mcnc piIuiIc(ic iniLe c p(ccuccc ce mci: hc(mcnc: pe|c
hipcIc|cmc.
FeecLcck u|I(c-cu(Ic: c hc(mcnc hipcIc|micc iniLe c p(cp(ic hipcIc|cmc ce p(ccuzi(
e::c me:mc hc(mcnc inIe(mcic.
/ (ecu|cccc cc :ec(eccc hc(mcnc| IcmLm pcce :e( efecIuccc pc(
ce:en:iLi|izcccc {inIe(nc|izcccc: inccIivcccc)

kegu|oo ce|u|or
/: c|u|c: :cc (ecu|ccc: pc( muiIc: hc(mcnc:. fccIc(e: ce c(e:cimenIc.
neu(cI(cn:mi::c(e: e ce(Ic: Icxinc: cI(cv: cc inIe(ccccc ce:Ie: |iccncc: ccm c:
:eu: (ecepIc(e: |ccc|izccc: nc :upe(f|cie cc: c|u|c:.
/: hc(mcnc: pepIicicc: e c: hc(mcnc: ce(ivccc: ce cmincccicc: ccIucm cc n|ve|
ce|u|c( pc( e:Ie meccni:mc ce (ecepccc.
CuI(c impc(IcnIe meccni:mc c cumenIc cc pe(mecLi|iccce cc memL(cnc
ce|u|c( c hc(mcnc: e:Ie(cice: cue (eccem ccm (ecepIc(e: inI(cce|u|c(e:.
/ memL(cnc p|c:mcIicc e c meic ciIcp|c:mcIicc :cc c: |ccci: cnce ccc((e c
inIe(ccccc inicic| enI(e |iccncc e (ecepIc(.
Ge(c|menIe c cue :ucece c inIe(ccccc |iccncc-(ecepIc( :e cc( nc memL(cnc
ciIcp|c:mcIicc. ccmc c cc:c cc: hc(mcnc: pc|ipepIicicc:.
C (ecepIc( ce(c|menIe (eccnhece c e:I(uIu(c cc hc(mcnc cumenIcncc c :uc
e:pecificiccce e cfiniccce pc(c c hc(mcnc.
/ fc(mcccc ce um ccmp|exc hc(mcnc| (ecepIc( p(cvccc c fc(mcccc ce um cu
mci: men:ccei(c:.

keceptores
Scc e:I(uIu(c: cc c|u|c c|vc cue (eccnhecem c hc(mcnc
Scc ccn:IiIu|cc: pc( um |ccc| ce (eccnhecimenIc e um |ccc| ce ccIivcccc

De(ivccc: ce cmincccicc:: epinef(inc. nc(-epinef(inc. hi:Icminc
De(ivccc: cc ccicc c(ccuiccnicc: p(cIcc|cncinc:
FepIicicc:: G|icccinc. in:u|inc. cc:I(inc. :ec(eIinc. cc|eci:Iccininc. cc(IiccI(cfinc.
hc(mcnc e:Iimu|ccc(c cc fc||cu|c. hc(mcnc e:Iimu|ccc(c cc Ii(cice. |uIeinizcnIe
pc(cIc(mcnc. vc:cp(e::inc.

2 Mensoge|ros
/MF c|c|icc {c/MF): fc(mccc c pc(Ii( cc /IF pc( ccccc cc enzimc cceni|cIc cic|c:e.
lnf|uencic c pe(mecLi|iccce ce|u|c( e ccIivc enzimc:. e:pecic|menIe cinc:e: p(cIeicc:
cue ce|e cepencem.
GMF c|c|icc {cGMF): fc(mccc c pc(Ii( cc GIF e ccIivccc pe|c cucni|cIc cic|c:e cc
memL(cnc ce|u|c(.
Cc|cic

Ironsduco de s|no|
/MFc
/: memL(cnc: ce|u|c(e: fc(mcm c :upe(f|cie (ecepIc(e: p(cIeicc: pc(c c:
hc(mcnc:. cue muiIc: veze: :cc I(cn:memL(cnc(e:.
/: hc(mcnc: |iccm-:e c e::e: (ecepIc(e: e:pec|ficc: e fc(mcm um comp|exo
hormono-receptor
E:Ic inIe(ccccc hc(mcnc-(ecepIc( p(cvccc umc c|Ie(cccc ccnfc(mccicnc|.
C (ecepIc( |icc-:e cc tronsductor de s|no|
/ mcic(ic cc: I(cn:cucIc(e: ce :inc| cc memL(cnc ce|u|c( :cc protenos |icccc: c
GIF (efe(icc: ccmc Frotenos G e cue :cc ccn:IiIu|cc: pc( 3 :uLuniccce: {u. e v).
87

/ :uLuniccce u cnce :e |icc c GIF e cue (ecce ci(ecIcmenIe nc ccm umc
enzimc. ccmc c oden||oto c|c|ose. ccIivcncc-c.
Exi:Iem cci: Iipc: ce :uLuniccce u {ccIivccc(c e iniLicc(c) e ccmc Ic| exi:Iem
(ecepIc(e: cue ccIivcm c cceni|cIc cic|c:e e cuI(c: cue iniLem.
/ |iccccc enI(e c (ecepIc( e c p(cIe|nc G ccIivc-c ce mccc c cue e|c |iLe(Ie c
GDF e :e |icue cc GIF. ccu:cncc c :epc(cccc cc: :uLuniccce: u cc: cuI(c:.
/ :uLuniccce u |icc-:e c cceni|cIc cic|c:e e ccIivc-c ce mccc c cue e:Ic ccIc|ize
c :|nIe:e ce /MFc c pc(Ii( ce /IF

C /MFc p(ccuzicc pe|c ccIiviccce cc cceni|cIc cic|c:e ccIivc c p(cIe|nc cinc:e c
cuc| pc::c c fc:fc(i|c( c: enzimc:-c|vc
Cucncc c GIF hic(c|izccc c GDF. umc (eccccc ccIc|izccc pe|c p(cp(ic
:uLuniccce u. e:Ic ceixc ce ccIivc( c /ceni|cIc cic|c:e. ce::cncc c p(ccuccc ce
/MFc.
/: :uLuniccce: cc p(cIe|nc G (ec::ccicm-:e e c hc(mcnc ci::ccic-:e cc (ecepIc(.
/ p(cIe|nc G pcce :e( iniLicc(c cu ccIivccc(c cc ce /ceni|cIc cic|c:e.

AIF AMFc + FF|
/ceni|cIc cic|c:e
88

Cc|cic
C cumenIc cc cc|cic ciIcp|c:mcIicc ceve-:e c
2 hc(mcnc:: c vc:cp(e::inc cu /DH e c
cc(enc|inc.
IcnIc c vc:cp(e::inc ccmc c cc(enc|inc
ccmLincm-:e ccm (ecepIc(e: e:pec|ficc: cue.
nc cc:c cc cc(enc|inc :cc c: (ecepIc(e:
cc(en(cicc:.
/ enI(ccc ce Cc|cic cI(cv: ce ccnci: icnicc:
pcce :e( cfecIccc pe|c inIe(ccccc hc(mcnc-
(ecepIc(. cue incuz c cLe(Iu(c ce::e: ccnci:.
CuI(c meccni:mc ce cumenIc( c ccncenI(cccc
ce cc|cic inI(cce|u|c( (e:u|Ic cc ccIivcccc cc
fc:fc|ipc:e C IcmLm pe|c inIe(ccccc
hc(mcnc-(ecepIc( {:eme|hcnIe c cc p(cIe|nc G
e p(cIe|nc cinc:e)
/ fc:fc|ipc:e C ccIc|izc c hic(c|i:e cc fc:fcIici|-
inc:iIc|-4.5-Li:fc:fcIc. cue :e i(c cinci( em cci:
men:ccei(c:. c inc:iIc|-1.4.5-I(ifc:fcIc {lF3) e c
1.2-cicci|c|ice(c|.
C lF3 |iLe(Ic cc|cic cc (eI|cu|c enccp|c:mcIicc. cue pc( :uc vez. i(c ccIivc( c
cinc:e.
C 1.2-cicci|c|ice(c| ccIivc ci(ecIcmenIe c F(cIe|nc cinc:e C.
C cumenIc cc: n|vei: ce cc|cic fcz ccm cue c cinc:e :e cp(cxime cc memL(cnc
ce|u|c( :encc ccIivccc pc( umc ccmLincccc ce Cc|cic. D/G e fc:fcIici|:e(inc.
Umc vez ccIivccc c p(cIe|nc cinc:e C fc:fc(i|c enzimc: e:pecificc: nc ciIc:c| cue
umc vez fc:fc(i|ccc: ficcm ccIivc:.

Ep|ne|r|no ou Adreno||no
Em :iIuccce: ce stress. c mecu|c :up(c-(enc| |iLe(Ic pc(c c ci(cu|cccc c(cnce:
cucnIiccce: ce cc(enc|inc e c|cumc nc(-cc(enc|inc.
Iem umc ccccc p(ic(iIc(ic nc: mscu|os
/ cc(enc|inc ccmLinc-:e ccm umc p(cIe|nc e:pec|ficc exi:IenIe nc inIe(ic( cc
memL(cnc. c receptor odregn|co.
C ccmp|exc cc(enc|inc-(ecepIc( nc p(e:encc ce GIF ccmLinc-:e ccm c p(cIe|nc
G cue I(ccuz c :inc| ccpcz ce ccIivc( c oden||c|c|ose.
E:Ic: hc(mcnc: e:Iimu|cm c p(ccuccc ce cAMF.
Funo:
E:Iimu|c c c|icccenc|i:e {IcmLm hepcIicc)
lniLe c inf|uxc ce c|icc:e
Ccn:umc p(efe(encic| cc: ccicc: cc(cc:. nc :ecuencic cc e:Iimu|cccc cc |ipc|i:e
E:Iimu|c c :ec(eccc cc c|icccinc e iniLe c cc in:u|inc.

G|ucogon ou g||cog|no
Nc ficccc. c c|icccinc Icmc c |ucc( cc cc(enc|inc.
Em :iIuccce: ce h|pog||cm|o. c: c|u|c: u {cc: i|hu: ce Lcnce(hcn:) cc pcnc(ec:
:ec(eIcm c|icccinc cue. cc ccmLinc(-:e c um (ecepIc( e:pec|ficc cc memL(cnc
ce|u|c(. e:Iimu|c c cceni|cic|c:e.
Funes:
E:Iimu|c( c c|icccenc|i:e
lniLe c c|icccne:e
lniLe c c|icc|i:e
/cIivc c c|iccneccne:e
lniLe c :|nIe:e ce ccicc: cc(cc: nc f|cccc {pc( ciminui( c fc(mcccc cc pi(uvcIc e
iniLu( c cc cceIi|-Cc/ cc(Lcxi|c:e)
/cIivc c |ipc|i:e nc ccipcciIc
/ c|icccinc umc hc(mcnc h|perg||cem|onte.
Hc(mcnc
FecepIc(
Fc:fcIici|inc:iIc|-4.5-Li:fc:fcIc
lF3 D/G
FeI|cu|c
Cc
2+

Cinc:e / Cinc:e l Cinc:e /
89

Insu||no
:inIeIizccc pe|c: c|u|c: cc: i|hu: cc pcnc(ec:
FecepIc( ce in:u|inc
C (ecepIc( ce in:u|inc fc(mccc pc( 2 subun|dodes u exI(cce|u|c(e: cnce :e |icc c
in:u|inc
/ |iccccc cc in:u|inc p(cvccc umc :ecuncic ce ccIivcccc enI(e c|Ie(ccce:
ccnfc(mccicnci: e |os|or||oo {cuIcfc:fc(i|cccc) ce (e:|cuc: ce t|ros|no
|ccc|izccc: nc: subun|dodes {inI(cce|u|c(e:)
/ ccIivcccc cc Ii(c:inc cinc:e Ic(nc-c ccpcz ce |os|or||or protenos ciIc:c|icc: {cc
n|ve| cc: (e:|cuc: ce :e(inc e I(ecninc) ce mccc c efecIivc( c :inc| cc in:u|inc.
Fc( cuI(c |ccc. c in:u|inc pcce incuzi( c ce:fc:fc(i|cccc ce cuI(c: p(cIe|nc: cI(cv:
ce um pc::|ve| 2 men:ccei(c. pcucc ccnhecicc.
Um pc::|ve| 2 Men:ccei(c vci ccIivc( c fc:fcp(cIeinc fc:fcIc:e ce:fc:fc(i|cccc
ce nume(c:c: enzimc: ccIivcncc-c: cu iniLincc-c:
/ ccIiviccce cc :ecuncc men:ccei(c cu cc Ii(c:inc cinc:e ceve :e( (e:pcn:cve|
pe|c mov|mento dos tronsportodores de g||cose cc ciIc:c| pc(c c :upe(f|cie cc:
c|u|c: ce mccc c cue c c|icc:e :ejc (eIi(ccc cc cc((enIe :cncu|nec.

NOIA: Iccc: c: c|u|c: Im (ecepIc(e: pc(c c in:u|inc {inc|u:ive c f|cccc). nc enIcnIc c
f|cccc ncc Iem (ecepIc(e: ce c|icc:e. / c|icc:e enI(c |iv(emenIe nc: c|u|c: cc
f|cccc.

/cccc cc in:u|inc
Ii(c:inc cinc:e Cc:ccIc ce fc:fc(i|ccce: inI(cce|u|c(e: EfeiIc: c |cncc p(czc:
Dife(encicccc ce|u|c( e c(e:cimenIc:
E:Iimu|cccc cc |ipcp(cIeinc |ipc:e.
2 Men:ccei(c Cc:ccIc ce ce:fc:fc(i|ccce: inI(cce|u|c(e: EfeiIc: c cu(Ic p(czc:
lniLiccc cc |ipc:e hc(mcnc cepencenIe nc: ccipcciIc: pc( ccIivcccc cc fc:fcIc:e
/cIivcccc ce fc:fcIc:e::
fc:fc(i|c:e cc c|icccnic {inccIivc).
:inIeIc:e cc c|icccnic {ccIivc):
fc:fcf(uIccinc:e 1 {ccIivc):
ccmp|exc ce ce:ic(ccenc:e pi(vicc {ccIivc):
cceIi|-Cc/ cc(Lcxi|c:e {ccIivc)
I(cn:fe(ncic ce p(cIe|nc: pc(c c :upe(f|cie ce|u|c(: I(cn:pc(Iccc( ce c|icc:e.
(ecpIc(e: ce LDL.

Funes:
/ in:u|inc fcci|iIc c I(cn:pc(Ie cc c|icc:e pc(c c inIe(ic( cc c|u|c.
E:Iimu|c c c|icccnic :inIc:e fcvc(ecencc. c::im. c c|icccne:e. umc vez ccIivc c
fc:fcp(cIe|nc fc:fcIc:e l.
E:Iimu|c c :|nIe:e ce ccicc: cc(cc:
E:Iimu|c c :|nIe:e ce p(cIeinc:
umc hc(mcnc hipcc|icemicnIe.
90

1
111
11.
.. E
EEn
nnz
zz|
||m
mmo
oo|
||o
oog
gg|
||o
oo

Enz|mos
Scc p(cIe|nc: c|cLu|c(e: cue exe(cem funcce: ce ccIc|i:e e/cu (ecu|cccc
Fc::uem umc e:pecificiccce muiIc e|evccc - cccc enzimc e:Ic ceneIiccmenIe
ceIe(minccc" cc en(c|cmenIc cc :ecuncic p(imc(ic em 3 cimen:ce: ccm umc
p(eci:c c(ienIcccc.
/: ccceic: |cIe(ci: pc|c(e: enccnI(cm-:e c :upe(f|cie {fcvc(ece c ccnIccIc ccm
ccuc)
/: ccceic: ncc pc|c(e: enccnI(cm-:e nc inIe(ic( cc mc|cu|c enzimcIicc. cujc:
c(upc: ncc icnizcvei: e:IcLe|ecem inIe(cccce: cue e:IcLi|izcm c ccnfc(mcccc
Ie(mccincmicc mci: fcvc(cve| cc mc|cu|c enzimcIicc.
/ :ecuncic e ci:pc:iccc e:pccic| cc: cmincccicc: :cc ccncicce: nece::c(ic:
pc(c c pcce( ccIc||Iicc. c cue exp|icc c c(cnce cife(encc ce cimen:ce: enI(e c
enzimc e c :uL:I(cIc.
Scc c: uniccce: funcicnci: cc meIcLc|i:mc ce|u|c(

Fropr|edodes
Scc ccIc|i:ccc(e: Lic|ccicc: exI(emcmenIe eficienIe:
/ce|e(cm c: (ecccce: cu|micc: ccm ciminuiccc cc Iempc ce cLIenccc cc
e:Iccc ce ecui||L(ic. :em c c|Ie(c(em - Lcixc cincc mci: c ene(cic ce ccIivcccc*.
/ ene(cic ce ccIivcccc (e:u|IcnIe cc |iccccc enI(e E e S:
1. Aumento o ve|oc|dode ce (eccccc pc( incuccc ce Ic(cce: e Ien:ce: nc
mc|cu|c ce :uL:I(cIc:
2. Or|g|no o|teroes de con|ormoo nc mc|cu|c ce enzimc
3. Aumento os mov|mentos cc mc|cu|c ce :uL:I(cIc.

EnccnI(cm-:e inIccIc: nc finc| cc (eccccc. ncc pc(Iicipcncc nem ccmc (eccenIe
nem ccmc p(ccuIc:
Ncc c|Ie(cm c ccn:IcnIe ce ecui||L(ic e ncc pc(Iicipcm nc e:IecuicmeI(ic cc
(eccccc.
Ncc pe(miIe cue pc::e c ccc((e( umc (eccccc cue ncc :ejc pc::|ve| cc pcnIc ce
vi:Ic Ie(mccincmicc
/cIucm em ccncenI(ccce: muiIc Lcixc::
/cIucm em ccncicce: :ucve: ce Iempe(cIu(c e pH:
Fc::uem Iccc: c: cc(ccIe(|:Iicc: cc: p(cIe|nc::
Fccem Ie( c :uc ccIiviccce (ecu|ccc:
E:Icc cuc:e :emp(e cenI(c cc: c|u|c:. ccmpc(IimenIc|izccc:.

* / energ|o de oct|voo (ecue(icc pc(c inicic( um p(cce::c cu|micc pcce :e(
fc(necicc cI(cv: ce umc e|evcccc cc Iempe(cIu(c cue cumenIe c cciIcccc
mc|ecu|c(. mc: nc cc:c cc: :e(e: vivc:. ncc :e pcce(c cumenIc( c Iempe(cIu(c cc:
:i:Iemc:. pci: Ic(nc(-:e inccmpcI|ve| ccm c vicc. {ccuecimenIc ce:ncIu(cccc
p(cIeicc)
Lccc|izcccc inI(cce|u|c( cc: enzimc: {enzimc: inI(cce|u|c(e:):
Nc|ec - mcnuIenccc. (encvcccc e uIi|izcccc cc cpc(e|hc cenIicc
91

MiIcccnc(ic: - (ecccce: ce cxiccccc-(ecuccc cue ce(cm ene(cic e nc: cc cic|c ce
K(eL:
FiLc::cmc: - Lic::|nIe:e ce p(cIe|nc:
Li:c::cmc: - enzimc: hic(c||Iicc:
CiIcp|c:mc - inIe(venienIe: c: vic: c|icc||Iicc:. :|nIe:e ce ccicc: cc(cc:. eIc.
MemL(cnc p|c:mcIicc - I(cn:cuccccc ce ene(cic.

Lccc|izcccc exI(cce|u|c(
Ex.: F|c:mc:
enzimc: funcicnci: {exc(eIccc: nc(mc|menIe pe|c f|cccc)
enzimc: ncc funcicnci: {cumenIcm c :|nIe:e p(cIeicc e c pe(mecLi|iccce e
pc(Iicipcm nc nec(c:e e |i:e ce|u|c()

C|oss|||coo
Ox|dorredutoses: Scc enzimc: cue ccIc|i:cm (ecccce: ce I(cn:fe(ncic ce e|ecI(ce:.
cu :ejc: (ecccce: ce cxi-(ecuccc. Ex: c: De:ic(ccenc:e: e c: Cxicc:e:
Irons|eroses: Enzimc: cue ccIc|i:cm (ecccce: ce I(cn:fe(ncic ce c(upc: funcicnci:
ccmc c(upc: cminc. fc:fcIc. cci|. cc(Lcxi|. eIc. Ex.: c: Cuinc:e: e c: I(cn:cminc:e:
H|dro|oses: CcIc|i:cm (ecccce: ce hic(c|i:e ce |icccce: ccvc|enIe:. Ex: /: pepIicc:e:
L|oses: CcIc|i:cm c cueL(c ce |icccce: ccvc|enIe: e c (emcccc ce mc|cu|c: ce
ccuc. cmcnic e cicxicc ce cc(Lcnc. Ex: De:ic(cIc:e: e De:cc(Lcxi|c:e:.
Isomeroses: CcIc|i:cm (ecccce: ce inIe(ccnve(:cc enI(e i:cme(c: cpIicc: cu
cecmI(icc:. Ex: Epime(c:e: e muIc:e:.
L|goses (ou $|ntetoses): CcIc|i:cm (ecccce: ce fc(mcccc ce ncvc: mc|cu|c: c pc(Ii(
cc |iccccc enI(e cuc: jc exi:IenIe:. c cu:Ic ce ene(cic {/IF). Ex: cminccci|-IFN/
:inIeIc:e:.

As enz|mos podem ser:
lnIei(cmenIe p(cIeicc:
Ccn:IiIu|cc: pc(:

/pcenzimc {inccIivc) + ccn:IiIuinIe: ncc p(cIeicc: = Hc|cenzimc {ccIivc)

Co|octores: :cc pecuenc: mc|cu|c: c(ccnicc: cu inc(ccnicc: cue pccem :e(
nece::c(ic: pc(c c funccc ce umc enzimc. pccencc cu ncc e:Ic( |icccc:
pe(mcnenIemenIe c mc|cu|c cc enzimc. mc: nc cu:ncic ce|e:. c enzimc
inccIivc.

Grupo prott|co: fc(mcm |icccce: ccvc|enIe ccm c cpcenzimc

Coenz|mos: :cc ccmpc:Ic: c(ccnicc:. cuc:e :emp(e ce(ivccc: ce viIcminc:. cue
ccIucm em ccnjunIc ccm c: enzimc:. Fccem ccIuc( :ecuncc 3 mcce|c::
Liccncc-:e c enzimc ccm cfiniccce :eme|hcnIe c cc :uL:I(cIc:
Liccncc-:e ccvc|enIemenIe num |ccc| p(cximc cu nc p(cp(ic :|Iic ccIc||Iicc cc
cpcenzimc:
/cIucncc ce mcnei(c inIe(mecic(ic cc: cci: exI(emc: ccimc ciIccc:.
Ies met||cos
Co|octor Enz|mo
Fc(Ie p(cIeicc {cpcenzimc)
Ccn:IiIuinIe ncc p(cIeicc
CcfccIc(
Ccenzimc
F(c:IIicc
Ncc F(c:IIicc
lcc meIc|icc
Fc:fc||picc
92

Cu
2+
Ce(u|cp|c:minc: Fenc|cxicc:e: U(icc:e
Mc
2+
Enc|c:e
Mn
2+
/(cinc:e
Mc
2+
XcnIinc-cxicc:e
Zn
2+
/nic(c:e cc(Lcnicc
K
+
Fi(uvcIc-cinc:e
Nc
+
/IFc:e memL(cnc(ic

I|mogn|os: p(cIe|nc: ccIc||Iicc: cue p(eci:cm ce :e ce:cccificccc: pc(c pc::c( c
fc(mc ccIivc.

F(cp(ieccce: ce umc enzimc c :e( e:Iucccc:
Frote|cos - pe:c mc|ecu|c(. pcnIc i:ce|cI(icc. n ce i:cenzimc:
Estruturo|s - ccmpc:iccc. :ecuncic ce cmincccicc:. c((cnjc e:pccic|
Enz|mt|cos - e:pecificiccce. n ce cenI(c: ccIivc:. meccni:mc ce (eccccc
8|o|og|cos - funcce:. ci:I(iLuiccc. efeiIc: ce ceficincic cu exce::c

Centro oct|vo: umc (ecicc cc enzimc cnce :e cc c |iccccc enI(e c enzimc {E) e c
:uL:I(cIc {S). :encc ccn:IiIu|cc pc(:
Am|noc|dos de contocto e coto|t|cos: fccu|Icm e pc::iLi|iIcm. (e:pecIivcmenIe c
|iccccc e (eccccc ccm c :uL:I(cIc.
Am|noc|dos de suporte: |ccc|izccc: fc(c cc cenI(c ccIivc. mcnIm c e:I(uIu(c cc
enzimc.

A||cn|c e:cec||cc cc: ccce|c: cc||cect|c|cc: F:cec|||c|ccce cc|c cccc :uc:t|ctc

/ |iccccc enI(e c enzimc e c :uL:I(cIc pcce envc|ve(:
inIe(cccce: icnicc:
|icccce: pc( pcnIe: ce hic(ccnic
inIe(cccce: Vcn ce( Wcc|:.

Centro oct|vo:
Lccc| ce fixcccc {ccmLinc ccm S pc( |icccce: f(ccc:)
CenI(c ccIc||Iicc {ccIuc :cL(e c S |evcncc-c c :cf(e( c (eccccc cu|micc)

Fc(c umc |iccccc eficienIe nece::c(ic cue:
1. Hcjc cce:cc enI(e c: mc|cu|c: E e S
2. /: mc|cu|c: S e:Iejcm c(ienIccc: cc((ecIcmenIe pc(c c cenI(c ccIivc
3. Se c(iem fc(cc: ce ccn:I(cncimenIc nc ccmp|exc ES

Mode|os
Mcce|c Chcve/Fechccu(c: F(ev um enccixe pe(feiIc cc :uL:I(cIc nc |ccc| ce
|iccccc. cue :e(ic (|cicc ccmc umc fechccu(c. Cu :ejc. c :uL:I(cIc ceve(ic pc::ui(
umc ceIe(minccc fc(mc ce mccc c pcce(-:e enccixc( nc enzimc.

Mcce|c cc /ju:Ie lncuzicc: Secuncc Kc:|cnc c: ccceic: ce cmincccicc: cc cenI(c
ccIivc :cc f|ex|vei: e :u:cepI|vei: ce (ec((cnjc:. pe(miIincc umc c(ienIcccc fcvc(cve|
ccm c mc|cu|c ce :uL:I(cIc.
93

Espec|||c|dode enz|mt|co: c f|exiLi|iccce e:I(uIu(c| cc: cmincccicc: exp|icc c
cife(encc ce cimen:ce: enI(e c enzimc e c :uL:I(cIc e c e:pecificiccce enzimcIicc.
/ exi:Incic ce enzimc: cue ccIucm em |icccce: cc me:mc Iipc. |ccc|izccc: em
e:I(uIu(c: cu|micc: cife(enIe:. evicencic cue c e:pecificiccce enzimcIicc ncc e:Ic
|imiIccc c pc(Ie cc mc|cu|c ce :uL:I(cIc cue ccIuc nc cenI(c ccIivc. mc: e:Ience-
:e c Iccc c mc|cu|c.
Espec|||c|dode obso|uto: c enzimc ncc cceiIc umc ceIe(minccc mc|cu|c ce
:uL:I(cIc.
Espec|||c|dode re|ot|vo: c enzimc I(cn:fc(mc um ccnjunIc ce :uL:I(cIc: cc me:mc
Iipc.
Estereoespec|||c|dode: c: enzimc: cpenc: ccIc|i:cm um cc: encnIicme(c: cu
cic:Ie(ecme(c:. excepIc nc cc:c cc: i:cme(c:e:. cue inIe(ccnve(Iem i:cme(c:.

Isoenz|mos:
E|ecI(cfc(eIiccmenIe cife(enIe: {pi: cife(enIe:)
CcIc|izcm c me:mc (eccccc
Ccmpc(IimenIc: {e Iecicc:) cife(enIe:. ccm cife(enIe: ccncicce: ce pH.
Iempe(cIu(c.

Mecon|smos de cot||se enz|mt|co
Cot||se gero| c|do-bose: um p(cIcc I(cn:fe(icc nc e:Iccc ce I(cn:iccc. pc( um cc
ccm ccceic: |cIe(ci: icnizcvei:.
cicc H
+
cu e
-

8c:e H
+
cu e
-

Feccccc ccIc|i:ccc pc( um ccicc {H/)
1. H + X <-> XH
+
+ /
-

2. XH
+
<-> Y + H
+

Ioro do $ubstroto: cepence cc Ic(ccc cc :uL:I(cIc incuzicc pe|c |iccccc cc
me:mc ccm c cenI(c ccIivc cc enzimc. c|ccnccncc c e:Iccc ce I(cn:iccc e
e:Iimu|cncc c :uc ccnve(:cc em p(ccuIc.

Cot||se Covo|ente
Fe:u|Ic cc cIccue nuc|ecfi|icc cu e|ecI(cfi|icc ce um (ccicc| cc cenI(c ccIc||Iicc :cL(e
c :uL:I(cIc. |iccncc-c ccvc|enIemenIe c enzimc e incuzincc c :uc I(cn:fc(mcccc em
p(ccuIc. Envc|ve ccm f(ecuncic c pc(Iicipcccc ce ccenzimc:.
/ p(e:encc cc enzimc pcce incuzi( c cpc(ecimenIc ce cc(cc: pc:iIivc: cu
neccIivc: nc :upe(f|cie cc :uL:I(cIc. fcci|iIcncc c cIccue ce ccenIe: nuc|ecf||icc:
cu e|ecI(cf||icc:. De um mccc ce(c| c: enzimc: ccIucm ccmc ccenIe:
nuc|ecf||icc:.
keoco nuc|eo|||co: envc|ve c :uL:IiIuiccc ce um c(upc pc( cuI(c.

/ enzimc pcce fc(nece( um p(cIcc cc :uL:I(cIc. fccu|Icncc c I(cn:fe(ncic ce cuI(c
p(cvenienIe ce ccenzimc.

94

E|e|tos de D|m|nu|o do Entrop|o: /: enzimc: cjuccm nc pc:icicncmenIc e nc
cefiniccc cc e:IecuicmeI(ic cc((ecIc cc (eccccc. fcci|iIcncc c: meccni:mc:
cnIe(ic(e:.

Iec(ic: ce ccIc|i:e
Fenomenos de super|c|e: c enzimc ccn:ice(ccc ccmc um |ccc| ce enccnI(c ccm c
:uL:I(cIc fcci|iIccc c fc(mcccc cc ccmp|exc {ES).
Frox|m|dode e or|entoo: c mc|cu|c ce cL:I(ccIc ceve(c e:Ic( p(cximc e Lem
c(ienIccc em (e|cccc cc: c(upc: funcicnci: cc cenI(c ccIivc cc enzimc
Aporec|mento de constrong|mento ||berdode do mo|cu|o de substroto. cue
fc(cccc c pe(mcnece( em |ccc| e:pec|ficc. e:IcLe|ecencc |icccce: ccm c: c(upc:
funcicnci: cc: cmincccicc: ce :upc(Ie e ccIc||Iicc:
Formoo de comp|exos |ntermed|r|os pc::uicc(e: ce cc|c(e: (ecuzicc: ce ene(cic
ce ccIivcccc. C vc|c( ce ene(cic ce ccIivcccc ce umc (eccccc enzimcIicc infe(ic(
cc cue :e cLIm nc (eccccc ncc enzimcIicc cnc|ccc.

CINIICA ENIIMIICA
/ cinIicc ce umc enzimc e:Iucccc cvc|icncc-:e c cucnIiccce ce p(ccuIc
fc(mccc cu c cucnIiccce ce :uL:I(cIc ccn:umicc pc( uniccce ce Iempc ce (eccccc
{ve|cciccce ce (eccccc).
E + S {ES] E+F

C ccmp|exc enzimc/:uL:I(cIc {ES) Iem umc ene(cic ce ccIivcccc |icei(cmenIe menc(
cue c cc :uL:I(cIc i:c|ccc. e c :uc fc(mcccc |evc cc cpc(ecimenIc cc e:Iccc ce
I(cn:iccc I:. / fc(mcccc ce F' c pc(Ii( ce ES c eIcpc |imiIcnIe cc ve|cciccce cc
(eccccc. / ve|cciccce ce umc (eccccc enzimcIicc cepence cc: ccncenI(ccce: cc
Enzimc e cc SuL:I(cIc.
Cucncc I(cIcmc: cc cinIicc ce (ecccce: enzimcIicc: hc cue Ie( em cIenccc:
1) Ve|cciccce inicic| cc (eccccc:
2) lnf|uncic cc ccncenI(cccc cc enzimc nc cinIicc:
3) lnf|uncic cc ccncenI(cccc cc :uL:I(cIc nc cinIicc:
4) lnf|uncic ce FccIc(e: ExIe(nc:.

1. Ve|cciccce inicic| cc (eccccc
/ ve|cciccce cc (eccccc enzimcIicc ncc :e
mcnIm ccn:IcnIe cc |cncc cc Iempc.
cec(e:cencc p(cc(e::ivcmenIe. /::im. nece::c(ic
incicc( c cue Iempc :e (efe(e umc ce(Ic ve|cciccce.
Ccn:ice(c-:e c ve|cciccce inicic| Vc. nume(iccmenIe
icuc| cc cec|ive cc (ecIc IcncenIe cc c(cficc ce
p(ccuIc fc(mccc em funccc cc Iempc. /
ve|cciccce inicic| :c cepence cc:
CcncenI(cccc cc enzimc
CcncenI(cccc cc :uL:I(cIc
/finiccce cc enzimc pe|c :uL:I(cIc

2.lnf|uncic cc {E] nc cinIicc
/nIe: ce umc (eccccc enzimcIicc cIinci( c
ecui||L(ic c ve|cciccce cc me:mc cumenIc pc(
cc(:cimc cc enzimc
/ ccncenI(cccc cc enzimc. ncc cfecIc c
ccn:IcnIe ce ecui||L(ic.

3. lnf|uncic cc {S] nc cinIicc

95

Fc(c Lcixc: ccncenI(ccce: ce :uL:I(cIc -
ve|oc|dode u concentroo {(eccccc ce 1
c(cem em (e|cccc cc :uL:I(cIc)
Fc(c c|Ic: ccncenI(ccce: - ve|cciccce Ience
pc(c um vc|c( c::impIcIicc - ve|oc|dode
mx|mo* (Vmx) {(eccccc ce c(cem ze(c).

* /: mc|cu|c: ce enzimc e:Icc :cIu(ccc: pe|c:
mc|cu|c: ce :uL:I(cIc. i:Ic . exi:Ie um exce::c
ce :uL:I(cIc (e|cIivcmenIe c enzimc - ccncicce:
:cIu(cnIe:.

3.c. Fcucccc ce /|cnce||:-/enten |!!3}

Michce|i: e MenIen - mcce|c ce (eccccc enzimcIicc pc(c cpenc: um :uL:I(cIc

K1 - ccn:IcnIe ce ve|cciccce ce fc(mcccc ce ES
K-1 - ccn:IcnIe ce ve|cciccce ce cec(cccccc ce ES
K2 - ccn:IcnIe ce ve|cciccce ce ceccmpc:iccc ce ES em p(ccuIc:

K-1 >> K2

Equoo de M|choe||s-Menten: (ep(e:enIc c efeiIc cc ccncenI(cccc ce :uL:I(cIc
:cL(e c ve|cciccce ce (eccccc enzimcIicc.

Km (constonte de o||n|dode ou de M|choe||s-Menten): ccncenI(cccc ce :uL:I(cIc em
cue c ve|cciccce meIcce cc ve|cciccce mcximc.
Km= [$] poro o quo| v= Vmx/ 2

l:Ic :icnificc cue. pc(c e:Ic ccncenI(cccc. c enzimc e:Ic 507 :cIu(ccc.
Exp(ime c cfiniccce cc(ccIe(|:Iicc. mc: inve(:cmenIe: cucnIc mcic( Km menc( :e(c
c cfiniccce cu e:pecificiccce.
Vmx = K2 [E0]

Fostu|odos do equoo de M|choe||s-Menten:
C ccmp|exc ES e:Ic e:Iccicnc(ic: {ES] K
Iccc c enzimc ccnve(Iicc nc ccmp|exc ES {ncc hc enzimc: |iv(e:)
Cucncc Iccc c E e:Ic :cL c fc(mc ce ES. enIcc V0 = Vmx = K2 [E$]

Cucncc [$] << Km : Feccccc ce 1 c(cem {ve|cciccce u {S])

Cucncc [$] >> Km : Feccccc ce c(cem ze(c

Estodo estoc|onr|o (8r|ggs e Ho|done): c ccmp|exc E: ncc e:Ic em ecui||L(ic {num
ceIe(minccc inIe(vc|c ce Iempc cp(ecicve|) ccm E e S. mc: c ve|cciccce ce
ceccmpc:iccc ce ES icnIicc c ve|cciccce ce fc(mcccc.
V = Vmx x [$]
Km + [$]
K1 K2
E + $ E$ E + F
K-1
V = K2[E0] x [$]
Km
V = Vmx
96

C e:Iccc e:Iccicnc(ic ccc((e cucncc c fc(mcccc ce ES icuc| c fc(mcccc ce F.

Cucncc c enzimc e:Ic ccmp|eIcmenIe :cIu(ccc pe|c :uL:I(cIc pcce-:e cefini(:
N de Iroco ou renovoo = Kcot = n ce mc|cu|c: ce S ccnve(Iicc: em p(ccuIc/pc(
cenI(c ccIivc/pc( uniccce ce Iempc {ccn:IcnIe ce ve|cciccce ce 1 c(cem)

bn|dodes de expresso do oct|v|dode Enz|mt|co
Ul = uniccce enzimcIicc: cucnIiccce ce enzimc cue ccIc|izc c I(cn:fc(mcccc ce 1
1mc|e ce SuL:I(cIc pc( 1min. c 25C. nc: ccncicce: cpiImc: ce pH e ccncenI(cccc
ce :uL:I(cIc {1mc|e S/min)
/cIiviccce E:pec|ficc: Ul/mc p(cIe|nc
/cIiviccce mc|ecu|c(: Ul/mc|e enzimc
KcIc|: cucnIiccce cue ccIc|i:c c I(cn:fc(mcccc ce 1 mc|e ce :uL:I(ccIc pc( :ecuncc
{1 KcIc| = 0 x 10
Ul)

3.L. / (ep(e:enIcccc ce Linewecve(-8u(k

/ Ecucccc ce Michce|i:-MenIen pcce :cf(e( (ec((cnjc: ce mccc c Ic(nc( mci: (icc(c:c
c ceIe(mincccc cc: vc|c(e: Ecucccc ce Linewecve(-8u(k

ScL e:Ic fc(mc Iemc: umc ecucccc cc Iipc y = cx + L. cnce |/v e 1/{S] :cc c:
vc(icvei: y e x.
Se nc ecucccc. fize( y = 0. i:Ic . |/v = 0. cLIm c vc|c( -1/Km. Se fize( |/{S] = 0 cu x =
0. cLIm 1/Vmcx.
C vc|c( cc Km |icc ccm (icc(. pci: :imI(icc cc inve(:c cc inIe(:ecccc ccm c
eixc cc: cLci::c:.

3.c. lniLiccc EnzimcIicc
E|ectores: vc(icm c ve|cciccce ce (eccccc: ccmpc:Ic: cue pc::cm c ccIivc( c:
enzimc: cu c iniLi( c :uc ccccc.

In|b|dores enz|mt|cos: efecIc(e: cue :e |iccm c umc enzimc. ciminuincc c :uc
ccIiviccce.
In|b|o enz|mt|co reversve|: c iniLicc( |icc-:e c enzimc ce um mccc ncc ccvc|enIe.
:encc c ccIiviccce Lic|ccicc cc enzimc (ecupe(ccc cpc: c ci::ccicccc:
Iipc::
i. CcmpeIiIivc
ii. Ncc ccmpeIiIivc
iii. lnccmpeIiIivc
In|b|o |rreversve|: hc mccificcccc ccvc|enIe e cefiniIivc nc :|Iic ce |iccccc cu nc
:|Iic ccIc||Iicc cc enzimc.

3.c.|. In|b|o enz|mt|co reversve| compet|t|vo
V = Vmx x [$]
Km + [$]
1 = (Km x 1 )+ 1
v Vmx [$]
V
97

C iniLicc( e:I(uIu(c|menIe :eme|hcnIe cc :uL:I(cIc e |icc-:e (eve(:ive|menIe cc
me:mc :|Iic ce |iccccc cc :uL:I(cIc
C efeiIc (eve(Iicc cumenIcncc-:e c ccncenI(cccc ce :uL:I(cIc.
E:Ie Iipc ce iniLiccc cepence cc: ccncenI(ccce: cc :uL:I(cIc e cc iniLicc(.
C iniLicc( ccmpeIe pe|c cenI(c ccIivc ccm c :uL:I(cIc. ciminuincc pc( i::c c
cfiniccce cc enzimc pe|c :uL:I(cIc {Km oumento). mc: c Vmx montm-se
inc|Ie(ccc.

3.c.||. In|b|o enz|mt|co reversve| no-compet|t|vo

C: iniLicc(e: fixcm-:e (eve(:ive|menIe nc enzimc ncuI(c |ccc| cue ncc c cenI(c
ccIivc. ncc ciminuincc c :uc ccccc pc( cciccc ce um exce::c ce :uL:I(cIc.
/ enzimc pcce |icc(-:e cc me:mc Iempc cc iniLicc( ncc ccmpeIiIivc e cc
:uL:I(cIc.
Nc enIcnIc. ccmc c ccnfc(mcccc cc cenI(c ccIivc ce c|cum mccc cfecIccc
pe|c p(e:encc cc iniLicc(. e::c |iccccc efecIuc-:e ccm menc: fcci|iccce e c
ve|cciccce cc (eccccc ciminui.
Um iniLicc( ncc ccmpeIiIivc ccmpc(Ic-:e ccmc :e (emcve::e c enzimc ccIivc cc
:c|uccc. (e:u|Icncc nc d|m|nu|o do ve|oc|dode mx|mo. encucnIc c o||n|dode
no se o|tero.
E:Ie Iipc ce iniLiccc cepence cpenc: cc ccncenI(cccc cc iniLicc(.

3.c.|||. In|b|o enz|mt|co reversve| |ncompet|t|vo

C iniLicc( |icc-:e cc ccmp|exc ES. e ncc c enzimc nc fc(mc |iv(e.
Ne:Ie cc:c. IcnIc c Km como o Vmx d|m|nuem.

I|po de In|b|o Km Vmx
CcmpeIiIivc /umenIc Ccn:IcnIe
Ncc ccmpeIiIivc Ccn:IcnIe Diminuiccc
lnccmpeIiIivc Diminuiccc Diminuiccc

4. FccIc(e: exIe(nc: cue inf|uencicm c ve|cciccce ce umc (eccccc enzimcIicc

4.c. Iempe(cIu(c

/ ve|cciccce ce Iccc: c: (ecccce: cu|micc: cumenIc ccm c
Iempe(cIu(c {mcic( nme(c ce chccue:). mc: i:Ic :c :e cp|icc
cI ceIe(minccc vc|c( ce Iempe(cIu(c - Iemperoturo pt|mo.
l:Ic . cI :e( cIincicc c Iempe(cIu(c c pc(Ii( cc cuc| c
p(cIe|nc enzimcIicc. pc( ccmecc( c ce:ncIu(c(. v IcmLm
ciminui( c :uc ccIiviccce.
98

4.L. pH

Exi:Ie um pH pt|mo cnce c ci:I(iLuiccc ce cc(cc: e|cI(icc:
cc mc|cu|c cc enzimc e. em e:pecic| cc cenI(c ccIc||Iicc.
icec| pc(c c ccIc|i:e.
/ icnizcccc cc: cmincccicc: cc cenI(c ccIc||Iicc pcce iniLi( c
ccIc|i:e: Lem ccmc c icnizcccc ce cuI(c: cmincccicc: fc(c cc
cenI(c ccIc||Iicc. pcce incuzi( umc c|Ie(cccc ccnfc(mccicnc|.
c|Ie(cncc c::im c cfiniccce enI(e c enzimc e c :uL:I(cIc.
C pH pcce cincc mccificc( c c(cu ce icnizcccc cc :uL:I(cIc.
pH: exI(emc: ce:ncIu(cm c: p(cIe|nc: e ccn:ecuenIemenIe c ccIiviccce
enzimcIicc.
Ccmc cccc enzimc Iem um pH cpIimc ce ccIucccc. exi:Iem enzimc: cue
ccIucm:
Em meic ccicc. ccmc c: enzimc: cc:I(icc:
Em meic neuI(c. ccmc c: cc Lccc
Em meic c|cc|inc ccmc c: cc cuccenc.

Apostila Completa Bioquimica Rita Luz

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Apostila Completa Bioquimica Rita Luz

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Fccu|ccce ce Mecicinc Mcc|c ll.

/ I(cn:ceIc|c:e (ecue( c pc(Iicipcccc cc ccenzimc Iicminc pi(cfc:fcIc {IFF).

keoces do sntese do g||cogn|o Enzimc:

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