STUDENT COMPANION
d d
F
d
U
d d
N
d
D A M
dh d d d d
E
d
N
d d
T A L S
d d d d
O F
Biochemistry
LIFEAT h
THE MOLECULAR d
h d
LEVEL h
Voet Voet Pratt
F O U R T H E D I T I O N
Akif Uzman Jerry Johnson JosephEichberg WilliamWidger
h d dh d h h
DonaldVoet Judith G. Voet CharlotteW.Pratt
dh h h d h d h h
,Thispage isintentionallyleftblank
h h
,STUDENTCOMPANIONTOACCOMPANY
FUNDAMENTALSOF
BIOCHEMISTRY
LIFEATTHEMOLECULARLEVEL
FourthEdition
AkifUzman
UniversityofHouston
JerryJohnson
UniversityofHouston
JosephEichberg
UniversityofHouston
WilliamWidger
UniversityofHouston
DonaldVoet
UniversityofPennsylvania
JudithG.Voet
SwarthmoreCollege
CharlotteW.Pratt
Seattle,Washington
JOHNWILEY&SONS,INC
, CoverDesignerMadelyn Lesure h
CoverIllustrationNormChristiansen
d
CoverPhotos
d
(VitruvianMan):©Odysseus/Alamyand© DennisHallinan/Alamy h
Molecularstructuresclockwisefromtop:basedonX-raystructuresthatwererespectivelydeterminedby
h h d
dRichardDickersonandHoraceDrew,Caltech;GerardBunick,UniversityofTennessee;ThomasSteitz,Yale
h h h d
dUniversity;AlfonsoMondragón,NorthwesternUniversity;VenkiRamakrishnan,MRCLaboratoryof
dMolecularBiology,Cambridge,U.K.;AndrewLeslieandJohnWalker,MRCLaboratoryofMolecular h d
dBiology,Cambridge,U.K.
Foundedin1807,JohnWiley&Sons,Inc.hasbeenavaluedsourceofknowledgeandunderstandingformore
h h h h d d
dthan200years,helpingpeoplearoundtheworldmeettheirneedsandfulfilltheiraspirations.Ourcompanyis
h h h d
dbuiltonafoundationofprinciplesthatincluderesponsibilitytothecommunitiesweserveandwhereweliveand
h d
dwork.In2008,welaunchedaCorporateCitizenshipInitiative,aglobalefforttoaddresstheenvironmental,
h d
dsocial,economic,andethicalchallengeswefaceinourbusiness.Amongtheissuesweareaddressingarecarbon d
dimpact,paperspecificationsandprocurement,ethicalconduct withinourbusinessand amongourvendors,and
h h h h d
dcommunityandcharitablesupport.Formoreinformation,pleasevisitourwebsite:
dwww.wiley.com/go/citizenship.
Copyright©2012 JohnWiley&Sons,Inc. h
Allrightsreserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystemortransmittedin h h d
danyformorbyanymeans,electronic,mechanical,photocopying,recording,scanningorotherwise,exceptas h d
dpermittedunderSections107or108ofthe1976UnitedStatesCopyrightAct,withouteitherthepriorwritten d
dpermissionofthePublisher,orauthorizationthroughpaymentoftheappropriateper-copyfeetotheCopyright h d
dClearanceCenter,Inc.,222Rosewood Drive,Danvers,MA01923,websitewww.copyright.com.Requeststo
h d
dthePublisherforpermissionshouldbeaddressedtothePermissionsDepartment,JohnWiley&Sons,Inc.111 h d
dRiverStreet,Hoboken,NJ07030-5774,(201)748-6011,fax(201)748-6008,website
dhttp://www.wiley.com/go/permissions.
Toorderbooksorforcustomerservice,pleasecall1-800-CALLWILEY(225-5945).
StudentCompanion(paperback)
h ISBN-13 978-1-1182-1827-3
d
Printedin theUnited StatesofAmerica10
h h h
d 987 654 321
d d
d d
F
d
U
d d
N
d
D A M
dh d d d d
E
d
N
d d
T A L S
d d d d
O F
Biochemistry
LIFEAT h
THE MOLECULAR d
h d
LEVEL h
Voet Voet Pratt
F O U R T H E D I T I O N
Akif Uzman Jerry Johnson JosephEichberg WilliamWidger
h d dh d h h
DonaldVoet Judith G. Voet CharlotteW.Pratt
dh h h d h d h h
,Thispage isintentionallyleftblank
h h
,STUDENTCOMPANIONTOACCOMPANY
FUNDAMENTALSOF
BIOCHEMISTRY
LIFEATTHEMOLECULARLEVEL
FourthEdition
AkifUzman
UniversityofHouston
JerryJohnson
UniversityofHouston
JosephEichberg
UniversityofHouston
WilliamWidger
UniversityofHouston
DonaldVoet
UniversityofPennsylvania
JudithG.Voet
SwarthmoreCollege
CharlotteW.Pratt
Seattle,Washington
JOHNWILEY&SONS,INC
, CoverDesignerMadelyn Lesure h
CoverIllustrationNormChristiansen
d
CoverPhotos
d
(VitruvianMan):©Odysseus/Alamyand© DennisHallinan/Alamy h
Molecularstructuresclockwisefromtop:basedonX-raystructuresthatwererespectivelydeterminedby
h h d
dRichardDickersonandHoraceDrew,Caltech;GerardBunick,UniversityofTennessee;ThomasSteitz,Yale
h h h d
dUniversity;AlfonsoMondragón,NorthwesternUniversity;VenkiRamakrishnan,MRCLaboratoryof
dMolecularBiology,Cambridge,U.K.;AndrewLeslieandJohnWalker,MRCLaboratoryofMolecular h d
dBiology,Cambridge,U.K.
Foundedin1807,JohnWiley&Sons,Inc.hasbeenavaluedsourceofknowledgeandunderstandingformore
h h h h d d
dthan200years,helpingpeoplearoundtheworldmeettheirneedsandfulfilltheiraspirations.Ourcompanyis
h h h d
dbuiltonafoundationofprinciplesthatincluderesponsibilitytothecommunitiesweserveandwhereweliveand
h d
dwork.In2008,welaunchedaCorporateCitizenshipInitiative,aglobalefforttoaddresstheenvironmental,
h d
dsocial,economic,andethicalchallengeswefaceinourbusiness.Amongtheissuesweareaddressingarecarbon d
dimpact,paperspecificationsandprocurement,ethicalconduct withinourbusinessand amongourvendors,and
h h h h d
dcommunityandcharitablesupport.Formoreinformation,pleasevisitourwebsite:
dwww.wiley.com/go/citizenship.
Copyright©2012 JohnWiley&Sons,Inc. h
Allrightsreserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystemortransmittedin h h d
danyformorbyanymeans,electronic,mechanical,photocopying,recording,scanningorotherwise,exceptas h d
dpermittedunderSections107or108ofthe1976UnitedStatesCopyrightAct,withouteitherthepriorwritten d
dpermissionofthePublisher,orauthorizationthroughpaymentoftheappropriateper-copyfeetotheCopyright h d
dClearanceCenter,Inc.,222Rosewood Drive,Danvers,MA01923,websitewww.copyright.com.Requeststo
h d
dthePublisherforpermissionshouldbeaddressedtothePermissionsDepartment,JohnWiley&Sons,Inc.111 h d
dRiverStreet,Hoboken,NJ07030-5774,(201)748-6011,fax(201)748-6008,website
dhttp://www.wiley.com/go/permissions.
Toorderbooksorforcustomerservice,pleasecall1-800-CALLWILEY(225-5945).
StudentCompanion(paperback)
h ISBN-13 978-1-1182-1827-3
d
Printedin theUnited StatesofAmerica10
h h h
d 987 654 321
d d
