STUDENT C O M P A N I O N
F U N D A M
k
E N T A L S O F
Biochemistry
LIFEkAT
THEk MOLECULARk
LEVEL
Voet Voet Pratt
F O U R T H E D I T I O N
AkifkUzman Jerryk Johnson JosephkEichberg
WilliamkWidgerkDonaldkVoet
JudithkG.kVoet CharlottekW.kPratt
,Thiskpagekiskintentionallykleftkblank
,STUDENTkCOMPANIONkTOkACCOMPANY
FUNDAMENTALSkOF
BIOCHEMISTRY
LIFEkATkTHEkMOLECULARkLEVEL
FourthkEdition
AkifkUzman
UniversitykofkHouston
JerrykJohnson
UniversitykofkHouston
JosephkEichberg
UniversitykofkHouston
WilliamkWidger
UniversitykofkHouston
DonaldkVoet
UniversitykofkPennsylvania
JudithkG.kVoet
SwarthmorekCollege
CharlottekW.kPratt
Seattle,kWashington
JOHNkWILEYk&kSONS,kINC
, CoverkDesignerkMadelynkLesurekC
overkIllustrationkNormkChristiansenk
CoverkPhotos
(VitruviankMan):k©kOdysseus/Alamykandk©kDenniskHallinan/Alamy
Molecularkstructureskclockwisekfromktop:kbasedkonkX-
raykstructureskthatkwerekrespectivelykdeterminedkbykRichardkDickersonkandkHoracekDrew,kCaltech;k
GerardkBunick,kUniversitykofkTennessee;kThomaskSteitz,kYalekUniversity;kAlfonsokMondragón,kNort
hwesternkUniversity;kVenkikRamakrishnan,kMRCkLaboratorykofkMolecularkBiology,kCambridge,kU.K
.;kAndrewkLesliekandkJohnkWalker,kMRCkLaboratorykofkMolecularkBiology,kCambridge,kU.K.
Foundedkink1807,kJohnkWileyk&kSons,kInc.khaskbeenkakvaluedksourcekofkknowledgekandkunderstanding
kforkmorekthank200kyears,khelpingkpeoplekaroundkthekworldkmeetktheirkneedskandkfulfillktheirkaspiratio
ns.kOurkcompanykiskbuiltkonkakfoundationkofkprincipleskthatkincludekresponsibilityktokthekcommunitiesk
wekservekandkwherekweklivekandkwork.kInk2008,kweklaunchedkakCorporatekCitizenshipkInitiative,kakglob
alkeffortktokaddresskthekenvironmental,ksocial,keconomic,kandkethicalkchallengeskwekfacekinkourkbusines
s.kAmongkthekissueskwekarekaddressingkarekcarbonkimpact,kpaperkspecificationskandkprocurement,kethic
alkconductkwithinkourkbusinesskandkamongkourkvendors,kandkcommunitykandkcharitableksupport.kForkm
orekinformation,kpleasekvisitkourkwebsite:kwww.wiley.com/go/citizenship.
Copyrightk©k2012 JohnkWileyk&kSons,kInc.
Allkrightskreserved.kNokpartkofkthiskpublicationkmaykbekreproduced,kstoredkinkakretrievalksystemkorktra
nsmittedkinkanykformkorkbykanykmeans,kelectronic,kmechanical,kphotocopying,krecording,kscanningkork
otherwise,kexceptkaskpermittedkunderkSectionsk107kork108kofkthek1976kUnitedkStateskCopyrightkAct,k
withoutkeitherkthekpriorkwrittenkpermissionkofkthekPublisher,korkauthorizationkthroughkpaymentkofkthek
appropriatekper-
copykfeektokthekCopyrightkClearancekCenter,kInc.,k222kRosewoodkDrive,kDanvers,kMAk01923,kwebsit
ekwww.copyright.com.kRequestsktokthekPublisherkforkpermissionkshouldkbekaddressedktokthekPermissio
nskDepartment,kJohnkWileyk&kSons,kInc.k111kRiverkStreet,kHoboken,kNJk07030-5774,k(201)k748-
6011,kfaxk(201)k748-6008,kwebsitekhttp://www.wiley.com/go/permissions.
Tokorderkbookskorkforkcustomerkservice,kpleasekcallk1-800-CALLkWILEYk(225-5945).
StudentkCompanionk(paperback) ISBN-13k 978-1-1182-1827-3
PrintedkinkthekUnitedkStateskofkAmeri
cak10k 9k8k7k 6k5k4k 3k2k1
F U N D A M
k
E N T A L S O F
Biochemistry
LIFEkAT
THEk MOLECULARk
LEVEL
Voet Voet Pratt
F O U R T H E D I T I O N
AkifkUzman Jerryk Johnson JosephkEichberg
WilliamkWidgerkDonaldkVoet
JudithkG.kVoet CharlottekW.kPratt
,Thiskpagekiskintentionallykleftkblank
,STUDENTkCOMPANIONkTOkACCOMPANY
FUNDAMENTALSkOF
BIOCHEMISTRY
LIFEkATkTHEkMOLECULARkLEVEL
FourthkEdition
AkifkUzman
UniversitykofkHouston
JerrykJohnson
UniversitykofkHouston
JosephkEichberg
UniversitykofkHouston
WilliamkWidger
UniversitykofkHouston
DonaldkVoet
UniversitykofkPennsylvania
JudithkG.kVoet
SwarthmorekCollege
CharlottekW.kPratt
Seattle,kWashington
JOHNkWILEYk&kSONS,kINC
, CoverkDesignerkMadelynkLesurekC
overkIllustrationkNormkChristiansenk
CoverkPhotos
(VitruviankMan):k©kOdysseus/Alamykandk©kDenniskHallinan/Alamy
Molecularkstructureskclockwisekfromktop:kbasedkonkX-
raykstructureskthatkwerekrespectivelykdeterminedkbykRichardkDickersonkandkHoracekDrew,kCaltech;k
GerardkBunick,kUniversitykofkTennessee;kThomaskSteitz,kYalekUniversity;kAlfonsokMondragón,kNort
hwesternkUniversity;kVenkikRamakrishnan,kMRCkLaboratorykofkMolecularkBiology,kCambridge,kU.K
.;kAndrewkLesliekandkJohnkWalker,kMRCkLaboratorykofkMolecularkBiology,kCambridge,kU.K.
Foundedkink1807,kJohnkWileyk&kSons,kInc.khaskbeenkakvaluedksourcekofkknowledgekandkunderstanding
kforkmorekthank200kyears,khelpingkpeoplekaroundkthekworldkmeetktheirkneedskandkfulfillktheirkaspiratio
ns.kOurkcompanykiskbuiltkonkakfoundationkofkprincipleskthatkincludekresponsibilityktokthekcommunitiesk
wekservekandkwherekweklivekandkwork.kInk2008,kweklaunchedkakCorporatekCitizenshipkInitiative,kakglob
alkeffortktokaddresskthekenvironmental,ksocial,keconomic,kandkethicalkchallengeskwekfacekinkourkbusines
s.kAmongkthekissueskwekarekaddressingkarekcarbonkimpact,kpaperkspecificationskandkprocurement,kethic
alkconductkwithinkourkbusinesskandkamongkourkvendors,kandkcommunitykandkcharitableksupport.kForkm
orekinformation,kpleasekvisitkourkwebsite:kwww.wiley.com/go/citizenship.
Copyrightk©k2012 JohnkWileyk&kSons,kInc.
Allkrightskreserved.kNokpartkofkthiskpublicationkmaykbekreproduced,kstoredkinkakretrievalksystemkorktra
nsmittedkinkanykformkorkbykanykmeans,kelectronic,kmechanical,kphotocopying,krecording,kscanningkork
otherwise,kexceptkaskpermittedkunderkSectionsk107kork108kofkthek1976kUnitedkStateskCopyrightkAct,k
withoutkeitherkthekpriorkwrittenkpermissionkofkthekPublisher,korkauthorizationkthroughkpaymentkofkthek
appropriatekper-
copykfeektokthekCopyrightkClearancekCenter,kInc.,k222kRosewoodkDrive,kDanvers,kMAk01923,kwebsit
ekwww.copyright.com.kRequestsktokthekPublisherkforkpermissionkshouldkbekaddressedktokthekPermissio
nskDepartment,kJohnkWileyk&kSons,kInc.k111kRiverkStreet,kHoboken,kNJk07030-5774,k(201)k748-
6011,kfaxk(201)k748-6008,kwebsitekhttp://www.wiley.com/go/permissions.
Tokorderkbookskorkforkcustomerkservice,kpleasekcallk1-800-CALLkWILEYk(225-5945).
StudentkCompanionk(paperback) ISBN-13k 978-1-1182-1827-3
PrintedkinkthekUnitedkStateskofkAmeri
cak10k 9k8k7k 6k5k4k 3k2k1
