STUDENT C O M P A N I O N
F U N D A M
5 6
E N T A L S O F
Biochemistry
LIFE56AT
THE5 6 MOLECULA
R5L6 EVEL
Voet Voet Pratt
F O U R T H E D I T I O N
Akif56Uzman Jerry5 6 Johnson Joseph56Eichberg
William56Widger56Donald56Voet
Judith56G.56Voet Charlotte56W.56Pratt
,This56page56is56intentionally56left56blank
,STUDENT56COMPANION56TO56ACCOMPANY
FUNDAMENTALS OF
BIOCHEMISTRY
LIFE AT THE MOLECULAR LEVEL
Fourth Edition
Akif56Uzman
University56of56Houston
Jerry56Johnson
University56of56Houston
Joseph56Eichberg
University56of56Houston
William56Widger
University56of56Houston
Donald56Voet
University56of56Pennsylvania
Judith56G.56Voet
Swarthmore56College
Charlotte56W.56Pratt
Seattle,56Washington
JOHN56WILEY56&56SONS,56INC
, Cover56Designer56Madelyn56Lesure5
6Cover56Illustration56Norm56Christia
nsen56Cover56Photos
(Vitruvian56Man):56©56Odysseus/Alamy56and56©56Dennis56Hallinan/Alamy
Molecular56structures56clockwise56from56top:56based56on56X-
ray56structures56that56were56respectively56determined56by56Richard56Dickerson56and56Horace56Drew,5
6Caltech;56Gerard56Bunick,56University56of56Tennessee;56Thomas56Steitz,56Yale56University;56Alfons
o56Mondragón,56Northwestern56University;56Venki56Ramakrishnan,56MRC56Laboratory56of56Molecula
r56Biology,56Cambridge,56U.K.;56Andrew56Leslie56and56John56Walker,56MRC56Laboratory56of56Molecu
lar56Biology,56Cambridge,56U.K.
Founded56in561807,56John56Wiley56&56Sons,56Inc.56has56been56a56valued56source56of56knowledge56and5
6understanding56for56more56than5620056years,56helping56people56around56the56world56meet56their56needs
56and56fulfill56their56aspirations.56Our56company56is56built56on56a56foundation56of56principles56that56incl
ude56responsibility56to56the56communities56we56serve56and56where56we56live56and56work.56In562008,56we
56launched56a56Corporate56Citizenship56Initiative,56a56global56effort56to56address56the56environmental,56s
ocial,56economic,56and56ethical56challenges56we56face56in56our56business.56Among56the56issues56we56are
56addressing56are56carbon56impact,56paper56specifications56and56procurement,56ethical56conduct56within5
6our56business56and56among56our56vendors,56and56community56and56charitable56support.56For56more56in
formation,56please56visit56our56website:56www.wiley.com/go/citizenship.
Copyright56©562012 John56Wiley56&56Sons,56Inc.
All56rights56reserved.56No56part56of56this56publication56may56be56reproduced,56stored56in56a56retrieval5
6system56or56transmitted56in56any56form56or56by56any56means,56electronic,56mechanical,56photocopying
,56recording,56scanning56or56otherwise,56except56as56permitted56under56Sections5610756or5610856of56the
56197656United56States56Copyright56Act,56without56either56the56prior56written56permission56of56the56Pu
blisher,56or56authorization56through56payment56of56the56appropriate56per-
copy56fee56to56the56Copyright56Clearance56Center,56Inc.,5622256Rosewood56Drive,56Danvers,56MA5601
923,56website56www.copyright.com.56Requests56to56the56Publisher56for56permission56should56be56addr
essed56to56the56Permissions56Department,56John56Wiley56&56Sons,56Inc.5611156River56Street,56Hoboken,
56NJ5607030-5774,56(201)56748-6011,56fax56(201)56748-
6008,56website56http://www.wiley.com/go/permissions.
To56order56books56or56for56customer56service,56please56call561-800-CALL56WILEY56(225-5945).
Student56Companion56(paperback) ISBN-135 6 978-1-1182-1827-3
Printed56in56the56United56States56of56
America56105 6 95685675 6 65655645 6 35
62561
F U N D A M
5 6
E N T A L S O F
Biochemistry
LIFE56AT
THE5 6 MOLECULA
R5L6 EVEL
Voet Voet Pratt
F O U R T H E D I T I O N
Akif56Uzman Jerry5 6 Johnson Joseph56Eichberg
William56Widger56Donald56Voet
Judith56G.56Voet Charlotte56W.56Pratt
,This56page56is56intentionally56left56blank
,STUDENT56COMPANION56TO56ACCOMPANY
FUNDAMENTALS OF
BIOCHEMISTRY
LIFE AT THE MOLECULAR LEVEL
Fourth Edition
Akif56Uzman
University56of56Houston
Jerry56Johnson
University56of56Houston
Joseph56Eichberg
University56of56Houston
William56Widger
University56of56Houston
Donald56Voet
University56of56Pennsylvania
Judith56G.56Voet
Swarthmore56College
Charlotte56W.56Pratt
Seattle,56Washington
JOHN56WILEY56&56SONS,56INC
, Cover56Designer56Madelyn56Lesure5
6Cover56Illustration56Norm56Christia
nsen56Cover56Photos
(Vitruvian56Man):56©56Odysseus/Alamy56and56©56Dennis56Hallinan/Alamy
Molecular56structures56clockwise56from56top:56based56on56X-
ray56structures56that56were56respectively56determined56by56Richard56Dickerson56and56Horace56Drew,5
6Caltech;56Gerard56Bunick,56University56of56Tennessee;56Thomas56Steitz,56Yale56University;56Alfons
o56Mondragón,56Northwestern56University;56Venki56Ramakrishnan,56MRC56Laboratory56of56Molecula
r56Biology,56Cambridge,56U.K.;56Andrew56Leslie56and56John56Walker,56MRC56Laboratory56of56Molecu
lar56Biology,56Cambridge,56U.K.
Founded56in561807,56John56Wiley56&56Sons,56Inc.56has56been56a56valued56source56of56knowledge56and5
6understanding56for56more56than5620056years,56helping56people56around56the56world56meet56their56needs
56and56fulfill56their56aspirations.56Our56company56is56built56on56a56foundation56of56principles56that56incl
ude56responsibility56to56the56communities56we56serve56and56where56we56live56and56work.56In562008,56we
56launched56a56Corporate56Citizenship56Initiative,56a56global56effort56to56address56the56environmental,56s
ocial,56economic,56and56ethical56challenges56we56face56in56our56business.56Among56the56issues56we56are
56addressing56are56carbon56impact,56paper56specifications56and56procurement,56ethical56conduct56within5
6our56business56and56among56our56vendors,56and56community56and56charitable56support.56For56more56in
formation,56please56visit56our56website:56www.wiley.com/go/citizenship.
Copyright56©562012 John56Wiley56&56Sons,56Inc.
All56rights56reserved.56No56part56of56this56publication56may56be56reproduced,56stored56in56a56retrieval5
6system56or56transmitted56in56any56form56or56by56any56means,56electronic,56mechanical,56photocopying
,56recording,56scanning56or56otherwise,56except56as56permitted56under56Sections5610756or5610856of56the
56197656United56States56Copyright56Act,56without56either56the56prior56written56permission56of56the56Pu
blisher,56or56authorization56through56payment56of56the56appropriate56per-
copy56fee56to56the56Copyright56Clearance56Center,56Inc.,5622256Rosewood56Drive,56Danvers,56MA5601
923,56website56www.copyright.com.56Requests56to56the56Publisher56for56permission56should56be56addr
essed56to56the56Permissions56Department,56John56Wiley56&56Sons,56Inc.5611156River56Street,56Hoboken,
56NJ5607030-5774,56(201)56748-6011,56fax56(201)56748-
6008,56website56http://www.wiley.com/go/permissions.
To56order56books56or56for56customer56service,56please56call561-800-CALL56WILEY56(225-5945).
Student56Companion56(paperback) ISBN-135 6 978-1-1182-1827-3
Printed56in56the56United56States56of56
America56105 6 95685675 6 65655645 6 35
62561