STUDENT C O M P A N I O N
F U N D A M E N T A L S O F
Biochemistry
LIFE AT
THE MOLECULAR
LEVEL
Voet Voet Pratt
F O U R T H E D I T I O N
Akif Uzman Jerry Johnson Joseph Eichberg William Widger
Donald Voet Judith G. Voet Charlotte W. Pratt
,STUDENT COMPANION TO ACCOMPANY
FUNDAMENTALS OF
BIOCHEMISTRY
LIFE AT THE MOLECULAR LEVEL
Fourth Edition
Akif Uzman
University of Houston
Jerry Johnson
University of Houston
Joseph Eichberg
University of Houston
William Widger
University of Houston
Donald Voet
University of Pennsylvania
Judith G. Voet
Swarthmore College
Charlotte W. Pratt
Seattle, Washington
JOHN WILEY & SONS, INC
,Cover Designer Madelyn Lesure
Cover Illustration Norm Christiansen
Cover Photos
(Vitruvian Man): © Odysseus/Alamy and © Dennis Hallinan/Alamy
Molecular structures clockwise from top: based on X-ray structures that were respectively determined
by Richard Dickerson and Horace Drew, Caltech; Gerard Bunick, University of Tennessee; Thomas
Steitz, Yale University; Alfonso Mondragón, Northwestern University; Venki Ramakrishnan, MRC
Laboratory of Molecular Biology, Cambridge, U.K.; Andrew Leslie and John Walker, MRC Laboratory
of Molecular Biology, Cambridge, U.K.
Founded in 1807, John Wiley & Sons, Inc. has been a valued source of knowledge and understanding for
more than 200 years, helping people around the world meet their needs and fulfill their aspirations. Our
company is built on a foundation of principles that include responsibility to the communities we serve and
where we live and work. In 2008, we launched a Corporate Citizenship Initiative, a global effort to
address the environmental, social, economic, and ethical challenges we face in our business. Among the
issues we are addressing are carbon impact, paper specifications and procurement, ethical conduct within
our business and among our vendors, and community and charitable support. For more information,
please visit our website: www.wiley.com/go/citizenship.
Copyright © 2012 John Wiley & Sons, Inc.
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or
transmitted in any form or by any means, electronic, mechanical, photocopying, recording, scanning or
otherwise, except as permitted under Sections 107 or 108 of the 1976 United States Copyright Act,
without either the prior written permission of the Publisher, or authorization through payment of the
appropriate per-copy fee to the Copyright Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA
01923, website www.copyright.com. Requests to the Publisher for permission should be addressed to the
Permissions Department, John Wiley & Sons, Inc. 111 River Street, Hoboken, NJ 07030-5774, (201)
748-6011, fax (201) 748-6008, website http://www.wiley.com/go/permissions.
To order books or for customer service, please call 1-800-CALL WILEY (225-5945).
Student Companion (paperback) ISBN-13 978-1-1182-1827-3
Printed in the United States of America
10 9 8 7 6 5 4 3 2 1
, Contents
1 Introduction to the Chemistry of Life 1
2 Water 9
3 Nucleotides, Nucleic Acids, and Genetic Information 21
4 Amino Acids 29
5 Proteins: Primary Structure 40
6 Proteins: Three-Dimensional Structure 49
7 Protein Function: Myoglobin and Hemoglobin, Muscle
Contraction, and Antibodies 60
8 Carbohydrates 72
9 Lipids and Biological Membranes 80
10 Membrane Transport 90
11 Enzymatic Catalysis 96
12 Enzyme Kinetics, Inhibition, and Control 108
13 Biochemical Signaling 121
14 Introduction to Metabolism 128
15 Glucose Catabolism 137
16 Glycogen Metabolism and Gluconeogenesis 146
17 Citric Acid Cycle 154
18 Electron Transport and Oxidative Phosphorylation 160
F U N D A M E N T A L S O F
Biochemistry
LIFE AT
THE MOLECULAR
LEVEL
Voet Voet Pratt
F O U R T H E D I T I O N
Akif Uzman Jerry Johnson Joseph Eichberg William Widger
Donald Voet Judith G. Voet Charlotte W. Pratt
,STUDENT COMPANION TO ACCOMPANY
FUNDAMENTALS OF
BIOCHEMISTRY
LIFE AT THE MOLECULAR LEVEL
Fourth Edition
Akif Uzman
University of Houston
Jerry Johnson
University of Houston
Joseph Eichberg
University of Houston
William Widger
University of Houston
Donald Voet
University of Pennsylvania
Judith G. Voet
Swarthmore College
Charlotte W. Pratt
Seattle, Washington
JOHN WILEY & SONS, INC
,Cover Designer Madelyn Lesure
Cover Illustration Norm Christiansen
Cover Photos
(Vitruvian Man): © Odysseus/Alamy and © Dennis Hallinan/Alamy
Molecular structures clockwise from top: based on X-ray structures that were respectively determined
by Richard Dickerson and Horace Drew, Caltech; Gerard Bunick, University of Tennessee; Thomas
Steitz, Yale University; Alfonso Mondragón, Northwestern University; Venki Ramakrishnan, MRC
Laboratory of Molecular Biology, Cambridge, U.K.; Andrew Leslie and John Walker, MRC Laboratory
of Molecular Biology, Cambridge, U.K.
Founded in 1807, John Wiley & Sons, Inc. has been a valued source of knowledge and understanding for
more than 200 years, helping people around the world meet their needs and fulfill their aspirations. Our
company is built on a foundation of principles that include responsibility to the communities we serve and
where we live and work. In 2008, we launched a Corporate Citizenship Initiative, a global effort to
address the environmental, social, economic, and ethical challenges we face in our business. Among the
issues we are addressing are carbon impact, paper specifications and procurement, ethical conduct within
our business and among our vendors, and community and charitable support. For more information,
please visit our website: www.wiley.com/go/citizenship.
Copyright © 2012 John Wiley & Sons, Inc.
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or
transmitted in any form or by any means, electronic, mechanical, photocopying, recording, scanning or
otherwise, except as permitted under Sections 107 or 108 of the 1976 United States Copyright Act,
without either the prior written permission of the Publisher, or authorization through payment of the
appropriate per-copy fee to the Copyright Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA
01923, website www.copyright.com. Requests to the Publisher for permission should be addressed to the
Permissions Department, John Wiley & Sons, Inc. 111 River Street, Hoboken, NJ 07030-5774, (201)
748-6011, fax (201) 748-6008, website http://www.wiley.com/go/permissions.
To order books or for customer service, please call 1-800-CALL WILEY (225-5945).
Student Companion (paperback) ISBN-13 978-1-1182-1827-3
Printed in the United States of America
10 9 8 7 6 5 4 3 2 1
, Contents
1 Introduction to the Chemistry of Life 1
2 Water 9
3 Nucleotides, Nucleic Acids, and Genetic Information 21
4 Amino Acids 29
5 Proteins: Primary Structure 40
6 Proteins: Three-Dimensional Structure 49
7 Protein Function: Myoglobin and Hemoglobin, Muscle
Contraction, and Antibodies 60
8 Carbohydrates 72
9 Lipids and Biological Membranes 80
10 Membrane Transport 90
11 Enzymatic Catalysis 96
12 Enzyme Kinetics, Inhibition, and Control 108
13 Biochemical Signaling 121
14 Introduction to Metabolism 128
15 Glucose Catabolism 137
16 Glycogen Metabolism and Gluconeogenesis 146
17 Citric Acid Cycle 154
18 Electron Transport and Oxidative Phosphorylation 160
