STUDENT C O M P A N I O N
F U N D A M
m
E N T A L S O F
Biochemistry
LIFE AT m
THE MOLECULAR
m m
LEVEL
Voet Voet Pratt
F O U R T H E D I T I O N
Akif Uzman
m Jerry Johnson Joseph Eichberg
m m
William WidgerDonald Voet
m m m
Judith G. Voet Charlotte W. Pratt
m m m m
,Thismpagemismintentionallymleftmblank
,STUDENT COMPANION TO ACCOMPANY m m m
FUNDAMENTALS OF m
BIOCHEMISTRY
LIFE AT THE MOLECULAR LEVEL
m m m m
Fourth Edition
m
Akif Uzman
m
University of Houston
m m
Jerry Johnson
m
University of Houston
m m
Joseph Eichberg m
University of Houston
m m
William Widger m
University of Houston
m m
Donald Voet m
University of Pennsylvania
m m
Judith G. Voet
m m
Swarthmore College m
Charlotte W. Pratt m m
Seattle, Washington
m
JOHNmWILEYm&mSONS,mINC
, CovermDesignermMadelynmLesuremCo
vermIllustrationmNormmChristiansenmC
overmPhotos
(VitruvianmMan):m©mOdysseus/Alamymandm©mDennismHallinan/Alamy
Molecularmstructuresmclockwisemfrommtop:mbasedmonmX-
raymstructuresmthatmweremrespectivelymdeterminedmbymRichardmDickersonmandmHoracemDrew,mCaltech;mG
erardmBunick,mUniversitymofmTennessee;mThomasmSteitz,mYalemUniversity;mAlfonsomMondragón,mNorth
westernmUniversity;mVenkimRamakrishnan,mMRCmLaboratorymofmMolecularmBiology,mCambridge,mU.K.;
mAndrewmLesliemandmJohnmWalker,mMRCmLaboratorym ofmMolecularmBiology,mCambridge,mU.K.
Foundedminm1807,mJohnmWileym&mSons,mInc.mhasmbeenmamvaluedmsourcemofmknowledgemandmunderstanding
mformmoremthanm200myears,mhelpingmpeoplemaroundmthemworldmmeetmtheirmneedsmandmfulfillmtheirmaspiration
s.mOurmcompanymismbuiltmonmamfoundationmofmprinciplesmthatmincludemresponsibilitymtomthemcommunitiesmw
emservemandmwheremwemlivemandmwork.mInm2008,mwemlaunchedmamCorporatemCitizenshipmInitiative,mamglobal
meffortmtomaddressmthemenvironmental,msocial,meconomic,mandmethicalmchallengesmwemfaceminmourmbusiness.
mAmongmthemissuesmwemaremaddressingmaremcarbonmimpact,mpapermspecificationsmandmprocurement,methical
mconductmwithinmourmbusinessmandmamongmourmvendors,mandmcommunitym andmcharitablem support.mFormmor
eminformation,mpleasemvisitmourmwebsite:mwww.wiley.com/go/citizenship.
Copyrightm©m2012 JohnmWileym&mSons,mInc.
Allmrightsmreserved.mNompartmofmthismpublicationmmaymbemreproduced,mstoredminmamretrievalmsystemmormtra
nsmittedminmanymformmormbymanymmeans,melectronic,mmechanical,mphotocopying,mrecording,mscanningmorm
otherwise,mexceptmasmpermittedmundermSectionsm107morm108mofmthem1976mUnitedmStatesmCopyrightmAct,m
withoutmeithermthempriormwrittenmpermissionmofmthemPublisher,mormauthorizationmthroughmpaymentmofmthem
appropriatemper-
copymfeemtomthemCopyrightmClearancemCenter,mInc.,m222mRosewoodmDrive,mDanvers,mMAm01923,mwebsit
emwww.copyright.com.mRequestsmtomthemPublishermformpermissionmshouldmbemaddressedmtomthemPermissio
nsmDepartment,mJohnmWileym&mSons,mInc.m111mRivermStreet,mHoboken,mNJm07030-5774,m(201)m748-
6011,mfaxm(201)m748-6008,mwebsitemhttp://www.wiley.com/go/permissions.
Tomordermbooksmormformcustomermservice,mpleasemcallm1-800-CALLmWILEYm(225-5945).
StudentmCompanionm(paperback) ISBN-13m 978-1-1182-1827-3
PrintedminmthemUnitedmStatesmofmAmeri
cam10m 9m8m7m 6m5m4m 3m2m1
F U N D A M
m
E N T A L S O F
Biochemistry
LIFE AT m
THE MOLECULAR
m m
LEVEL
Voet Voet Pratt
F O U R T H E D I T I O N
Akif Uzman
m Jerry Johnson Joseph Eichberg
m m
William WidgerDonald Voet
m m m
Judith G. Voet Charlotte W. Pratt
m m m m
,Thismpagemismintentionallymleftmblank
,STUDENT COMPANION TO ACCOMPANY m m m
FUNDAMENTALS OF m
BIOCHEMISTRY
LIFE AT THE MOLECULAR LEVEL
m m m m
Fourth Edition
m
Akif Uzman
m
University of Houston
m m
Jerry Johnson
m
University of Houston
m m
Joseph Eichberg m
University of Houston
m m
William Widger m
University of Houston
m m
Donald Voet m
University of Pennsylvania
m m
Judith G. Voet
m m
Swarthmore College m
Charlotte W. Pratt m m
Seattle, Washington
m
JOHNmWILEYm&mSONS,mINC
, CovermDesignermMadelynmLesuremCo
vermIllustrationmNormmChristiansenmC
overmPhotos
(VitruvianmMan):m©mOdysseus/Alamymandm©mDennismHallinan/Alamy
Molecularmstructuresmclockwisemfrommtop:mbasedmonmX-
raymstructuresmthatmweremrespectivelymdeterminedmbymRichardmDickersonmandmHoracemDrew,mCaltech;mG
erardmBunick,mUniversitymofmTennessee;mThomasmSteitz,mYalemUniversity;mAlfonsomMondragón,mNorth
westernmUniversity;mVenkimRamakrishnan,mMRCmLaboratorymofmMolecularmBiology,mCambridge,mU.K.;
mAndrewmLesliemandmJohnmWalker,mMRCmLaboratorym ofmMolecularmBiology,mCambridge,mU.K.
Foundedminm1807,mJohnmWileym&mSons,mInc.mhasmbeenmamvaluedmsourcemofmknowledgemandmunderstanding
mformmoremthanm200myears,mhelpingmpeoplemaroundmthemworldmmeetmtheirmneedsmandmfulfillmtheirmaspiration
s.mOurmcompanymismbuiltmonmamfoundationmofmprinciplesmthatmincludemresponsibilitymtomthemcommunitiesmw
emservemandmwheremwemlivemandmwork.mInm2008,mwemlaunchedmamCorporatemCitizenshipmInitiative,mamglobal
meffortmtomaddressmthemenvironmental,msocial,meconomic,mandmethicalmchallengesmwemfaceminmourmbusiness.
mAmongmthemissuesmwemaremaddressingmaremcarbonmimpact,mpapermspecificationsmandmprocurement,methical
mconductmwithinmourmbusinessmandmamongmourmvendors,mandmcommunitym andmcharitablem support.mFormmor
eminformation,mpleasemvisitmourmwebsite:mwww.wiley.com/go/citizenship.
Copyrightm©m2012 JohnmWileym&mSons,mInc.
Allmrightsmreserved.mNompartmofmthismpublicationmmaymbemreproduced,mstoredminmamretrievalmsystemmormtra
nsmittedminmanymformmormbymanymmeans,melectronic,mmechanical,mphotocopying,mrecording,mscanningmorm
otherwise,mexceptmasmpermittedmundermSectionsm107morm108mofmthem1976mUnitedmStatesmCopyrightmAct,m
withoutmeithermthempriormwrittenmpermissionmofmthemPublisher,mormauthorizationmthroughmpaymentmofmthem
appropriatemper-
copymfeemtomthemCopyrightmClearancemCenter,mInc.,m222mRosewoodmDrive,mDanvers,mMAm01923,mwebsit
emwww.copyright.com.mRequestsmtomthemPublishermformpermissionmshouldmbemaddressedmtomthemPermissio
nsmDepartment,mJohnmWileym&mSons,mInc.m111mRivermStreet,mHoboken,mNJm07030-5774,m(201)m748-
6011,mfaxm(201)m748-6008,mwebsitemhttp://www.wiley.com/go/permissions.
Tomordermbooksmormformcustomermservice,mpleasemcallm1-800-CALLmWILEYm(225-5945).
StudentmCompanionm(paperback) ISBN-13m 978-1-1182-1827-3
PrintedminmthemUnitedmStatesmofmAmeri
cam10m 9m8m7m 6m5m4m 3m2m1