Netter's Essential Biochemistry

1008 Pages·2018·28.614 MB·English

by  Peter Ronner

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2015v1.0
NETTER:
It’s How You Know Anatomy.
Netter Collection of Medical
Illustrations, 2nd Edition
The Netter Collection of Medical Illustrations,
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Netter’s Essential Histology
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NET TER’S
ESSENTIAL
BIOCHEMISTRY
PETER RONNER, PhD
Professor of Biochemistry and Molecular Biology
Professor of Pharmaceutical Sciences
Department of Biochemistry and Molecular Biology
Thomas Jefferson University
Philadelphia, Pennsylvania
Illustrations by
Frank H. Netter, MD
Contributing Illustrators
Carlos A.G. Machado, MD
James A. Perkins, MS, MFA
Kip Carter, MS, CMI
Tiffany Davanzo, MA, CMI
1600 John F. Kennedy Blvd.
Ste 1800
Philadelphia, PA 19103-2899
NETTER’S ESSENTIAL BIOCHEMISTRY ISBN: 978-1-929007-63-9
Copyright © 2018 by Elsevier, Inc. All rights reserved.
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Notices
Knowledge and best practice in this field are constantly changing. As new research and experience broaden
our understanding, changes in research methods, professional practices, or medical treatment may become
necessary.
Practitioners and researchers must always rely on their own experience and knowledge in evaluating and
using any information, methods, compounds, or experiments described herein. In using such information
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Library of Congress Cataloging-in-Publication Data
Names: Ronner, Peter, 1951- author. | Netter, Frank H. (Frank Henry),
1906-1991, illustrator. | Machado, Carlos A. G., illustrator. |
Craig, John A., illustrator. | Perkins, James A., illustrator.
Title: Netter’s biochemistry / Peter Ronner ; illustrations by
Frank H. Netter ; contributing illustrators, Carlos A.G. Machado,
John A. Craig, James A. Perkins.
Other titles: Biochemistry
Description: Philadelphia, PA : Elsevier, [2018] | Includes bibliographical
references and index.
Identifiers: LCCN 2016024484 | ISBN 9781929007639 (pbk. : alk. paper)
Subjects: | MESH: Biochemical Phenomena | Biochemistry
Classification: LCC QP514.2 | NLM QU 34 | DDC 572–dc23 LC record available at
https://lccn.loc.gov/2016024484
Executive Content Strategist: Elyse O’Grady
Content Development Specialist: Stacy Eastman
Publishing Services Manager: Patricia Tannian
Senior Project Manager: Carrie Stetz
Design Direction: Julia Dummitt
Printed in China
Last digit is the print number: 9 8 7 6 5 4 3 2 1
To Wanda and Lukas
About the Author
Peter Ronner, PhD, is Professor of Biochemistry and Molecular Biology at the
Sidney Kimmel College of Medicine at Thomas Jefferson University in Phila-
delphia. He holds a secondary appointment as Professor of Pharmaceutical
Sciences in the College of Pharmacy at Thomas Jefferson University. Dr. Ronner
received his PhD in Biochemistry from the Swiss Federal Institute of Technol-
ogy (ETH) in Zurich. His former laboratory research involved studies of pan-
creatic hormone secretion. Dr. Ronner has taught medical students for nearly
30 years and pharmacy students for almost 10 years. He is also a past chair of
the Association of Biochemistry Course Directors (now Association of Bio-
chemistry Educators). At Jefferson, he has received numerous awards for his
teaching, including a Lindback Award and a portrait painting.
vi
Acknowledgments
Many people have helped me write this book, but Dr. John John Thomas, PhD
Thomas from New York University School of Medicine Research Associate Professor (Retired)
deserves a place of honor. He and I worked on this book Department of Biochemistry and Molecular Pharmacology
together until we were forced to pause for a few years. New York University School of Medicine
Years earlier, at the University of Pennsylvania, the late Dr. New York, New York
Annemarie Weber introduced me to teaching biochemistry to
medical students. She was a tremendous role model. At I am very thankful to Emine Ercikan Abali for coauthoring
Thomas Jefferson University, Dr. Darwin Prockop planted in the chapters on Triglycerides and Hypertriglyceridemia and
my mind the idea of writing a biochemistry textbook. Many Cholesterol Metabolism and Hypercholesterolemia.
years later, Paul Kelly (then at Icon Learning Systems),
approached me with the idea of using Dr. Netter’s images for Emine Ercikan Abali, PhD
a biochemistry review book. This appealed to me because Associate Professor of Biochemistry and Molecular Biology
biochemistry is taught as a rather abstract science that stu- Rutgers Robert Wood Johnson Medical School
dents have difficulty linking to actual patients. The Netter Piscataway, New Jersey
images, I hoped, would provide the views of the practicing
physician. Thanks to the support of my chairman, Dr. Jeffrey I am very grateful to the following persons for contributing
Benovic, this book project became part of my scholarly pur- their expertise and reviewing chapters:
suits. I am thankful for the invaluable feedback the many
students of medicine and pharmacy at Jefferson gave me over David Axelrod, MD
the years. Associate Professor of Medicine
I would like to thank the team at Elsevier for their support, Department of Medicine
especially Elyse O’Grady (Senior Content Strategist), Stacy Sidney Kimmel Medical College at Thomas Jefferson
Eastman and Marybeth Thiel (Content Development Special- University
ists), as well as Carrie Stetz (Senior Project Manager/ Philadelphia, Pennsylvania
Specialist).
Finally, I would like to thank my family and friends for Samir K. Ballas, MD
their support while writing this book. Emeritus Professor of Medicine and Pediatrics
This book is dedicated to my wife Wanda and my son Cardeza Foundation for Hematologic Research
Lukas. Wanda has been a key influence on me, because she Department of Medicine, Division of Hematology
has continuously given me her perspective as a practicing Sidney Kimmel Medical College at Thomas Jefferson
physician and medical student educator. Lukas, a chemistry University
major and current medical student, has been my most trusted Philadelphia, Pennsylvania
adviser on questions about young learners, chemistry, and
artwork, and he has reviewed much of my writing. James C. Barton, MD
Medical Director
Southern Iron Disorders Center;
COAUTHORS AND CHAPTER REVIEWERS
Clinical Professor of Medicine
I am deeply indebted to John Thomas for his contribu- Department of Medicine
tions, which involved designing this book and writing drafts University of Alabama at Birmingham
of several chapters: Clinical Tests Based on DNA or RNA; Birmingham, Alabama
Basic Genetics for Biochemistry; Transcription and RNA
Processing; Translation and Posttranslational Protein Pro- Jeffrey L. Benovic, PhD
cessing; Pentose Phosphate Pathway, Oxidative Stress, and Professor
Glucose 6-Phosphate Dehydrogenase Deficiency; Oxidative Department of Biochemistry and Molecular Biology
Phosphorylation and Mitochondrial Diseases; Fatty Acids, Sidney Kimmel Medical College at Thomas Jefferson
Ketone Bodies, and Ketoacidosis; Triglycerides and Hyper- University
triglyceridemia; Cholesterol Metabolism and Hypercholester- Philadelphia, Pennsylvania
olemia; Steroid Hormones and Vitamin D; Eicosanoids; and
Signaling.
vviiii
viii Acknowledgments
Bruno Calabretta, MD, PhD Christopher Haines, MD
Professor Assistant Professor
Department of Cancer Biology Department of Family and Community Medicine
Sidney Kimmel Medical College at Thomas Jefferson Sidney Kimmel Medical College at Thomas Jefferson
University University
Philadelphia, Pennsylvania Philadelphia, Pennsylvania
Gino Cingolani, PhD Steven K. Herrine, MD
Professor Professor
Department of Biochemistry and Molecular Biology Department of Medicine
Sidney Kimmel Medical College at Thomas Jefferson Sidney Kimmel Medical College at Thomas Jefferson
University University
Philadelphia, Pennsylvania Philadelphia, Pennsylvania
Joe Deweese, PhD Jacqueline M. Hibbert, PhD
Associate Professor Associate Professor
Department of Pharmaceutical Sciences Department of Microbiology, Biochemistry and
Lipscomb University, College of Pharmacy and Health Immunology
Sciences Morehouse School of Medicine
Nashville, Tennessee Atlanta, Georgia
Tina Bocker Edmonston, MD Jan B. Hoek, PhD
Associate Professor Professor
Department of Pathology and Laboratory Medicine Department of Pathology, Anatomy, and Cell Biology
Cooper University Health Care, Cooper Medical School at Sidney Kimmel Medical College at Thomas Jefferson
Rowan University University
Camden, New Jersey Philadelphia, Pennsylvania
Steven Ellis, PhD Tanis Hogg, PhD
Professor Associate Professor
Department of Biochemistry Department of Medical Education
University of Louisville Texas Tech University Health Sciences Center El Paso, Paul
Louisville, Kentucky L. Foster School of Medicine
El Paso, Texas
Masumi Eto, PhD
Associate Professor Ya-Ming Hou, PhD
Department of Molecular Physiology and Biophysics Professor
Sidney Kimmel Medical College at Thomas Jefferson Department of Biochemistry and Molecular Biology
University Sidney Kimmel Medical College at Thomas Jefferson
Philadelphia, Pennsylvania University
Philadelphia, Pennsylvania
Andrzej Fertala, PhD
Professor Serge A. Jabbour, DM
Department of Orthopaedic Surgery Professor
Sidney Kimmel Medical College at Thomas Jefferson Department of Medicine
University Sidney Kimmel Medical College at Thomas Jefferson
Philadelphia, Pennsylvania University
Philadelphia, Pennsylvania
Elizabeth Gilje, BS
Medical Student Francis E. Jenney Jr, PhD
Sidney Kimmel Medical College at Thomas Jefferson Associate Professor
University Department of Biomedical Sciences
Philadelphia, Pennsylvania Georgia Campus–PCOM
Suwanee, Georgia