Biochemistr
C O N C EP TS AN D C O N N EC T I O N S
SECOND EDITION
Dean R. Appling
THE UNIVERSITY OF TEXAS AT AUSTIN
Spencer J. Anthony-Cahill
WESTERN WASHINGTON UNIVERSITY
Christopher K. Mathews
OREGON STATE UNIVERSITY
330 Hudson Street, New York, NY 10013
A01_APPL1621_02_SE_FM.indd 1
,Editor in Chief: Jeanne Zalesky
Acquisitions Editor: Chris Hess
Director of Development: Jennifer Hart
Marketing Manager: Elizabeth Bell
Development Editor: Matt Walker
Art Development Editor: Jay McElroy
Program Manager: Kristen Flathman
Content Producer: Anastasia Slesareva
Text Permissions Project Manager: Tim Nicholls
Text Permissions Specialist: James Fortney, Lumina Datamatics
Project Management Team Lead: David Zielonka
Production Management: Mary Tindle, Cenveo
Compositor: Cenveo
Design Manager: Marilyn Perry
Interior Designer: Elise Lansdon
Cover Designer: Mark Ong, Side by Side Studios Illustrators:
ImagineeringArt, Inc.
Photo Researcher: Mo Spuhler
Photo Lead: Maya Melenchuk / Eric Shrader
Photo Permissions: Kathleen Zander / Matt Perry
Operations Specialist: Stacey Weinberger
Cover Background Photo
Credit: The Human
Spliceosome
Dr. Berthold Kastner, Max
Planck lnstitute of
Biophysical Chemisty
Molecular graphics
and analyses were
performed with the
UCSF Chimera
package. Chimera is
developed by the
Resource for
Biocomputing,
Visualization, and
Informatics at the
University of
California, San Francisco (supported by NIGMS
P41-GM103311)
Chimpanzee Photo Credit: Fiona Rogers/Getty
A01_APPL1621_02_SE_FM.indd 2
, Images
Copyright © 2019, 2016 Pearson Education, Inc. All Rights Reserved. Printed in the United States of
America. This publication is protected by copyright, and permission should be obtained from the publisher
prior to any prohibited reproduction, storage in a retrieval system, or transmission in any form or by any
means, electronic, mechanical, photocopying, recording, or otherwise. For information regarding
permissions, request forms, and the appropriate contacts within the Pearson Education Global Rights &
Permissions Department, please visit www.pearsoned.com/permissions.
Acknowledgments of third-party content appear on pages C-1–C-3, which constitutes an extension of this
copyright page.
PEARSON, ALWAYS LEARNING, MasteringTM Chemistry, and Learning Catalytics are exclusive
trademarks in the U.S. and/or other countries owned by Pearson Education, Inc. or its affiliates.
Unless otherwise indicated herein, any third-party trademarks that may appear in this work are the
property of their respective owners, and any references to third-party trademarks, logos, or other trade
dress are for demonstrative or descriptive purposes only. Such references are not intended to imply any
sponsorship, endorsement, authorization, or promotion of Pearson’s products by the owners of such
marks, or any relationship between the owner and Pearson Education, Inc. or its affiliates, authors,
licensees, or distributors.
Library of Congress Cataloging-in-Publication Data
Names: Appling, Dean Ramsay, author. | Anthony-Cahill, Spencer J., author. |
Mathews, Christopher K., 1937- author.
Title: Biochemistry : concepts and connections / Dean R. Appling, Spencer J.
Anthony-Cahill, Christopher K. Mathews.
Description: Second edition. | New York : Pearson, [2019] | Includes
bibliographical references and index.
Identifiers: LCCN 2017047599| ISBN 9780134641621 | ISBN 0134641620
Subjects: | MESH: Biochemical Phenomena
Classification: LCC RB112.5 | NLM QU 34 | DDC 612/.015--
dc23 LC record available at https://lccn.loc.gov/2017047599
[ Third-Party Trademark] [TM/®] is a [registered] trademark of [Third Party]. Used under
license. 1 18
ISBN 10: 0-134-64162-0; ISBN 13: 978-0-134-64162-1
www.pearson.com
A01_APPL1621_02_SE_FM.indd 3
, Brief Contents
1 Biochemistry and the Language 16 Lipid Metabolism 512
of Chemistry 2
2 The Chemical Foundation of Life:
Weak 17 Interorgan and Intracellula
Interactions in an Aqueous Environment Metabolism in Vertebrates
18
18 Amino Acid and Nitrogen
3 The Energetics of Life 48
19 Nucleotide Metabolism 61
4 Nucleic Acids 72
20 Mechanisms of Signal Tra
5 Introduction to Proteins: The Primary
Level of Protein Structure 108 6 The Three-
21 Genes, Genomes, and Ch
Dimensional Structure of Proteins 144 22 DNA Replication 686 23
Recombination, and Rear
7 Protein Function and Evolution 190
Transcription and Posttran
8 Enzymes: Biological Catalysts 232 742
9 Carbohydrates: Sugars, 25 Information Decoding: Tra
Saccharides, Posttranslational Protein P
Glycans 278 10 Lipids,
Membranes, and Cellular 26 Regulation of Gene Expre
Transport 304
APPENDIX I: ANSWERS TO SELECTED
11 Chemical Logic of Metabolism 340 A-1
12 Carbohydrate Metabolism: Glycolysis, APPENDIX II: REFERENCES A-20
Gluconeogenesis, Glycogen Metabolism, CREDITS C-1
and the Pentose Phosphate Pathway 374
INDEX I-1
13 The Citric Acid Cycle 420
14 Electron Transport,
Oxidative
Phosphorylation, and Oxygen
Metabolism 450
15 Photosynthesis 486
A01_APPL1621_02_SE_FM.indd 4
C O N C EP TS AN D C O N N EC T I O N S
SECOND EDITION
Dean R. Appling
THE UNIVERSITY OF TEXAS AT AUSTIN
Spencer J. Anthony-Cahill
WESTERN WASHINGTON UNIVERSITY
Christopher K. Mathews
OREGON STATE UNIVERSITY
330 Hudson Street, New York, NY 10013
A01_APPL1621_02_SE_FM.indd 1
,Editor in Chief: Jeanne Zalesky
Acquisitions Editor: Chris Hess
Director of Development: Jennifer Hart
Marketing Manager: Elizabeth Bell
Development Editor: Matt Walker
Art Development Editor: Jay McElroy
Program Manager: Kristen Flathman
Content Producer: Anastasia Slesareva
Text Permissions Project Manager: Tim Nicholls
Text Permissions Specialist: James Fortney, Lumina Datamatics
Project Management Team Lead: David Zielonka
Production Management: Mary Tindle, Cenveo
Compositor: Cenveo
Design Manager: Marilyn Perry
Interior Designer: Elise Lansdon
Cover Designer: Mark Ong, Side by Side Studios Illustrators:
ImagineeringArt, Inc.
Photo Researcher: Mo Spuhler
Photo Lead: Maya Melenchuk / Eric Shrader
Photo Permissions: Kathleen Zander / Matt Perry
Operations Specialist: Stacey Weinberger
Cover Background Photo
Credit: The Human
Spliceosome
Dr. Berthold Kastner, Max
Planck lnstitute of
Biophysical Chemisty
Molecular graphics
and analyses were
performed with the
UCSF Chimera
package. Chimera is
developed by the
Resource for
Biocomputing,
Visualization, and
Informatics at the
University of
California, San Francisco (supported by NIGMS
P41-GM103311)
Chimpanzee Photo Credit: Fiona Rogers/Getty
A01_APPL1621_02_SE_FM.indd 2
, Images
Copyright © 2019, 2016 Pearson Education, Inc. All Rights Reserved. Printed in the United States of
America. This publication is protected by copyright, and permission should be obtained from the publisher
prior to any prohibited reproduction, storage in a retrieval system, or transmission in any form or by any
means, electronic, mechanical, photocopying, recording, or otherwise. For information regarding
permissions, request forms, and the appropriate contacts within the Pearson Education Global Rights &
Permissions Department, please visit www.pearsoned.com/permissions.
Acknowledgments of third-party content appear on pages C-1–C-3, which constitutes an extension of this
copyright page.
PEARSON, ALWAYS LEARNING, MasteringTM Chemistry, and Learning Catalytics are exclusive
trademarks in the U.S. and/or other countries owned by Pearson Education, Inc. or its affiliates.
Unless otherwise indicated herein, any third-party trademarks that may appear in this work are the
property of their respective owners, and any references to third-party trademarks, logos, or other trade
dress are for demonstrative or descriptive purposes only. Such references are not intended to imply any
sponsorship, endorsement, authorization, or promotion of Pearson’s products by the owners of such
marks, or any relationship between the owner and Pearson Education, Inc. or its affiliates, authors,
licensees, or distributors.
Library of Congress Cataloging-in-Publication Data
Names: Appling, Dean Ramsay, author. | Anthony-Cahill, Spencer J., author. |
Mathews, Christopher K., 1937- author.
Title: Biochemistry : concepts and connections / Dean R. Appling, Spencer J.
Anthony-Cahill, Christopher K. Mathews.
Description: Second edition. | New York : Pearson, [2019] | Includes
bibliographical references and index.
Identifiers: LCCN 2017047599| ISBN 9780134641621 | ISBN 0134641620
Subjects: | MESH: Biochemical Phenomena
Classification: LCC RB112.5 | NLM QU 34 | DDC 612/.015--
dc23 LC record available at https://lccn.loc.gov/2017047599
[ Third-Party Trademark] [TM/®] is a [registered] trademark of [Third Party]. Used under
license. 1 18
ISBN 10: 0-134-64162-0; ISBN 13: 978-0-134-64162-1
www.pearson.com
A01_APPL1621_02_SE_FM.indd 3
, Brief Contents
1 Biochemistry and the Language 16 Lipid Metabolism 512
of Chemistry 2
2 The Chemical Foundation of Life:
Weak 17 Interorgan and Intracellula
Interactions in an Aqueous Environment Metabolism in Vertebrates
18
18 Amino Acid and Nitrogen
3 The Energetics of Life 48
19 Nucleotide Metabolism 61
4 Nucleic Acids 72
20 Mechanisms of Signal Tra
5 Introduction to Proteins: The Primary
Level of Protein Structure 108 6 The Three-
21 Genes, Genomes, and Ch
Dimensional Structure of Proteins 144 22 DNA Replication 686 23
Recombination, and Rear
7 Protein Function and Evolution 190
Transcription and Posttran
8 Enzymes: Biological Catalysts 232 742
9 Carbohydrates: Sugars, 25 Information Decoding: Tra
Saccharides, Posttranslational Protein P
Glycans 278 10 Lipids,
Membranes, and Cellular 26 Regulation of Gene Expre
Transport 304
APPENDIX I: ANSWERS TO SELECTED
11 Chemical Logic of Metabolism 340 A-1
12 Carbohydrate Metabolism: Glycolysis, APPENDIX II: REFERENCES A-20
Gluconeogenesis, Glycogen Metabolism, CREDITS C-1
and the Pentose Phosphate Pathway 374
INDEX I-1
13 The Citric Acid Cycle 420
14 Electron Transport,
Oxidative
Phosphorylation, and Oxygen
Metabolism 450
15 Photosynthesis 486
A01_APPL1621_02_SE_FM.indd 4
