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,this page left intentionally blank,Molecular Cell Biology
,ABOUT THE AUTHORS
HARVEY LODISH is Professor of Biology and Professor of Biological Engineering at the Massachusetts Institute of Technol-
ogy and a Founding Member of the Whitehead Institute for Biomedical Research. Dr. Lodish is also a member of the National
Academy of Sciences and the American Academy of Arts and Sciences and was President (2004) of the American Society for
Cell Biology. He is well known for his work on cell-membrane physiology, particularly the biosynthesis of many cell-surface pro-
teins, and on the cloning and functional analysis of several cell-surface receptor proteins, such as the erythropoietin and TGF–β
receptors. His laboratory also studies long noncoding RNAs and microRNAs that regulate the development and function of
hematopoietic cells and adipocytes. Dr. Lodish teaches undergraduate and graduate courses in cell biology and biotechnology.
Photo credit: John Soares.
ARNOLD BERK holds the UCLA Presidential Chair in Molecular Cell Biology in the Department of Microbiology, Immunology,
and Molecular Genetics and is a member of the Molecular Biology Institute at the University of California, Los Angeles. Dr. Berk
is also a fellow of the American Academy of Arts and Sciences. He is one of the discoverers of RNA splicing and of mechanisms
for gene control in viruses. His laboratory studies the molecular interactions that regulate transcription initiation in mammalian
cells, focusing in particular on adenovirus regulatory proteins. He teaches an advanced undergraduate course in cell biology of
the nucleus and a graduate course in biochemistry. Photo credit: Penny Jennings/UCLA Department of Chemistry & Biochemistry.
CHRIS A. KAISER is the Amgen Inc. Professor in the Department of Biology at the Massachusetts Institute of Technology.
He is also a former Department Head and former Provost. His laboratory uses genetic and cell biological methods to under-
stand how newly synthesized membrane and secretory proteins are folded and stored in the compartments of the secretory
pathway. Dr. Kaiser is recognized as a top undergraduate educator at MIT, where he has taught genetics to undergraduates for
many years. Photo credit: Chris Kaiser.
MONTY KRIEGER is the Whitehead Professor in the Department of Biology at the Massachusetts Institute of Technology and
a Senior Associate Member of the Broad Institute of MIT and Harvard. Dr. Krieger is also a member of the National Academy of
Sciences. For his innovative teaching of undergraduate biology and human physiology as well as graduate cell biology courses,
he has received numerous awards. His laboratory has made contributions to our understanding of membrane trafficking
through the Golgi apparatus and has cloned and characterized receptor proteins important for pathogen recognition and the
movement of cholesterol into and out of cells, including the HDL receptor. Photo credit: Monty Krieger.
ANTHONY BRETSCHER is Professor of Cell Biology at Cornell University and a member of the Weill Institute for Cell and
Molecular Biology. His laboratory is well known for identifying and characterizing new components of the actin cytoskeleton
and elucidating the biological functions of those components in relation to cell polarity and membrane traffic. For this work,
his laboratory exploits biochemical, genetic, and cell biological approaches in two model systems, vertebrate epithelial cells
and the budding yeast. Dr. Bretscher teaches cell biology to undergraduates at Cornell University. Photo credit: Anthony Bretscher.
HIDDE PLOEGH is Professor of Biology at the Massachusetts Institute of Technology and a member of the Whitehead Institute
for Biomedical Research. One of the world’s leading researchers in immune-system behavior, Dr. Ploegh studies the various
tactics that viruses employ to evade our immune responses and the ways our immune system distinguishes friend from foe.
Dr. Ploegh teaches immunology to undergraduate students at Harvard University and MIT. Photo credit: Hidde Ploegh.
ANGELIKA AMON is Professor of Biology at the Massachusetts Institute of Technology, a member of the Koch Institute for
Integrative Cancer Research, and Investigator at the Howard Hughes Medical Institute. She is also a member of the National
Academy of Sciences. Her laboratory studies the molecular mechanisms that govern chromosome segregation during mitosis
and meiosis and the consequences—aneuploidy—when these mechanisms fail during normal cell proliferation and cancer
development. Dr. Amon teaches undergraduate and graduate courses in cell biology and genetics. Photo credit: Pamela DiFraia/
Koch Institute/MIT.
KELSEY C. MARTIN is Professor of Biological Chemistry and Psychiatry and interim Dean of the David Geffen School of Medi-
cine at the University of California, Los Angeles. She is the former Chair of the Biological Chemistry Department. Her laboratory
studies the ways in which experience changes connections between neurons in the brain to store long-term memories—a
process known as synaptic plasticity. She has made important contributions to elucidating the molecular and cell biological
mechanisms that underlie this process. Dr. Martin teaches basic principles of neuroscience to undergraduates, graduate
students, dental students, and medical students. Photo credit: Phuong Pham.
,Molecular Cell
Biology
EIGHTH EDITION
Harvey Lodish
Arnold Berk
Chris A. Kaiser
Monty Krieger
Anthony Bretscher
Hidde Ploegh
Angelika Amon
Kelsey C. Martin
New York
,Publisher: Katherine Ahr Parker
Acquisitions Editor: Beth Cole
Developmental Editors: Erica Champion, Heather Moffat
Editorial Assistants: Nandini Ahuja, Abigail Fagan
Executive Marketing Manager: Will Moore
Senior Project Editor: Elizabeth Geller
Design Manager: Blake Logan
Text Designer: Patrice Sheridan
Cover Design: Blake Logan
Illustration Coordinator: Janice Donnola
Art Development Editor: H. Adam Steinberg, Art for Science
Permissions Manager: Jennifer MacMillan
Photo Editor: Sheena Goldstein
Photo Researcher: Teri Stratford
Text Permissions: Felicia Ruocco, Hilary Newman
Media and Supplements Editors: Amy Thorne, Kathleen Wisneski
Senior Media Producer: Chris Efstratiou
Senior Production Supervisor: Paul Rohloff
Composition: codeMantra
Printing and Binding: RR Donnelley
Cover Image: Dr. Tomas Kirchhausen and Dr. Lei Lu
ABOUT THE COVER: Imaging of the intracellular organelles of a live
human HeLa cell shows the dramatic morphological changes that accompany
the process of cell division. The membrane of the endoplasmic reticulum (ER)
is labeled green by a fluorescently tagged component of the translocon (GFP-
Sec61β) and chromatin is labeled red by a fluorescently tagged histone (H2B-
mRFP). Front: An interphase cell showing uncondensed chromatin filling the
nucleus, with the ER as a reticulum of cisternae surrounding the nucleus and
interconnected with lace-like tubules at the cell periphery. Back: Prior to cell
division the chromatin condenses to reveal the worm-like structure of individual
chromosomes, the nuclear envelope breaks down, and the ER condenses into
an array of cisternae surrounding the condensed chromosomes. As cell division
proceeds the replicated chromosomes will segregate equally into two daughter
cells, nuclear envelopes will form in the daughter cells, and the ER will return to
its characteristic reticular organization. Cover photo: Dr. Tomas Kirchhausen &
Dr. Lei Lu.
Library of Congress Control Number: 2015957295
ISBN-13: 978-1-4641-8339-3
ISBN-10: 1-4641-8339-2
© 2016, 2013, 2008, 2004 by W. H. Freeman and Company
All rights reserved.
Printed in the United States of America
First printing
W. H. Freeman and Company
One New York Plaza, Suite 4500, New York, NY 10004-1562
www.macmillanhighered.com
,TO OUR STUDENTS AND TO OUR TEACHERS,
from whom we continue to learn,
AND TO OUR FAMILIES,
for their support, encouragement, and love
,this page left intentionally blank
,PREFACE
In writing the eighth edition of Molecular Cell Biology, we long-term memories—a process known as synaptic plasticity.
have incorporated many of the spectacular advances made Dr. Martin received her undergraduate degree in English and
over the past four years in biomedical science, driven in part American Language and Literature at Harvard University.
by new experimental technologies that have revolutionized After serving as a Peace Corps volunteer in the Democratic
many fields. Fast techniques for sequencing DNA, allied Republic of the Congo, she earned an MD and PhD at Yale
with efficient methods to generate and study mutations in University. She teaches basic neurobiology to undergraduate,
model organisms and to map disease-causing mutations in graduate, dental, and medical students.
humans, have illuminated a basic understanding of the func-
tions of many cellular components, including hundreds of
human genes that affect diseases such as diabetes and cancer.
Revised, Cutting-Edge Content
For example, advances in genomics and bioinformat- The eighth edition of Molecular Cell Biology includes new
ics have uncovered thousands of novel long noncoding and improved chapters:
RNAs that regulate gene expression, and have generated r “Molecules, Cells, and Model Organisms” (Chapter 1) is an
insights into and potential therapies for many human dis- improved and expanded introduction to cell biology. It retains
eases. Powerful genome editing technologies have led to an the overviews of evolution, molecules, different forms of life,
unprecedented understanding of gene regulation and func- and model organisms used to study cell biology found in previ-
tion in many types of living organisms. Advances in mass ous editions. In this edition, it also includes a survey of eukary-
spectrometry and cryoelectron microscopy have enabled otic organelles, which was previously found in Chapter 9.
dynamic cell processes to be visualized in spectacular de-
tail, providing deep insight into both the structure and the r “Culturing and Visualizing Cells” (Chapter 4) has been
function of biological molecules, post-translational modifi- moved forward (previously Chapter 9) as the techniques
cations, multiprotein complexes, and organelles. Studies of used to study cells become ever more important. Light-sheet
specific nerve cells in live organisms have been advanced by microscopy, super-resolution microscopy, and two-photon
optogenetic technologies. Advances in stem-cell technology excitation microscopy have been added to bring this chapter
have come from studies of the role of stem cells in plant up to date.
development and of regeneration in planaria.
r All aspects of mitochondrial and chloroplast structure
Exploring the most current developments in the field is
and function have been collected in “Cellular Energetics”
always a priority in writing a new edition, but it is also im-
(Chapter 12). This chapter now begins with the structure
portant to us to communicate the basics of cell biology clear-
of the mitochondrion, including its endosymbiotic origin
ly by stripping away as much extraneous detail as possible to
and organelle genome (previously in Chapter 6). The chap-
focus attention on the fundamental concepts of cell biology.
ter now discusses the role of mitochondria-associated mem-
To this end, in addition to introducing new discoveries and
branes (MAMs) and communication between mitochondria
technologies, we have streamlined and reorganized several
and the rest of the cell.
chapters to clarify processes and concepts for students.
r Cell signaling has been reframed to improve student
accessibility. “Signal Transduction and G Protein–Coupled
New Co-Author, Kelsey C. Martin Receptors” (Chapter 15) begins with an overview of the con-
The new edition of MCB introduces a new member to our cepts of cell signaling and methods for studying it, followed
author team, leading neuroscience researcher and edu- by examples of G protein–coupled receptors performing
cator Kelsey C. Martin of the University of California, multiple roles in different cells. “Signaling Pathways That
Los Angeles. Dr. Martin is Professor of Biological Chemis- Control Gene Expression” (Chapter 16) now focuses on
try and Psychiatry and interim Dean of the David Geffen gene expression, beginning with a new discussion of Smads.
School of Medicine at UCLA. Her laboratory uses Aply- Further examples cover the major signaling pathways that
sia and mouse models to understand the cell and molecu- students will encounter in cellular metabolism, protein deg-
lar biology of long-term memory formation. Her group radation, and cellular differentiation. Of particular interest
has made important contributions to elucidating the mo- is a new section on Wnt and Notch signaling pathways con-
lecular and cell biological mechanisms by which experience trolling stem-cell differentiation in planaria. The chapter
changes connections between neurons in the brain to store ends by describing how signaling pathways are integrated
vii
, (a) Point-scanning confocal Two-photon excitation to form a cellular response in insulin and glucagon control
microscopy microscopy of glucose metabolism.
r Our new co-author, Kelsey C. Martin, has extensively
Electron excited state
revised and updated “Cells of the Nervous System”
(Chapter 22) to include several new developments in the
Excitation
photon 2
field. Optogenetics, a technique that uses channelrhodop-
Excitation Emission (960 nm) Emission sins and light to perturb the membrane potential of a cell,
photon photon photon can be used in live animals to link neural pathways with
(488 nm) (507 nm) (507 nm)
behavior. The formation and pruning of neural pathways
Excitation
photon 1 in the central nervous system is under active investigation,
(960 nm) and a new discussion of signals that govern these processes
focuses on the cell-cell contacts involved. This discussion
Electron ground state leads to an entirely new section on learning and memory,
which explores the signals and molecular mechanisms
underlying synaptic plasticity.
(b)
Objective lens of microscope
Increased Clarity, Improved Pedagogy
As experienced teachers of both undergraduate and gradu-
ate students, we are always striving to improve student un-
derstanding. Being able to visualize a molecule in action
Immobilized can have a profound effect on a student’s grasp of the mo-
mouse lecular processes within a cell. With this in mind, we have
updated many of the molecular models for increased clarity
and added models where they can deepen student under-
standing. From the precise fit required for tRNA charging,
to the conservation of ribosome structure, to the dynamic
strength of tropomyosin and troponin in muscle contraction,
these figures communicate the complex details of molecu-
(c) lar structure that cannot be conveyed in schematic diagrams
alone. In conjunction with these new models, their schematic
icons have been revised to more accurately represent them,
allowing students a smooth transition between the molecu-
lar details of a structure and its function in the cell.
New Discoveries, New Methodologies
r Model organisms Chlamydomonas reinhardtii (for study
of flagella, chloroplast formation, photosynthesis, and
phototaxis) and Plasmodium falciparum (novel organelles
and a complex life cycle) (Ch. 1)
r Intrinsically disordered proteins (Ch. 3)
r Chaperone-guided folding and updated chaperone
structures (Ch. 3)
FIGURE 4-21 Two-photo excitation microscopy allows r Unfolded proteins and the amyloid state and disease
deep penetration for intravital imaging. (a) In conventional (Ch. 3)
point-scanning confocal microscopy, absorption of a single
r Hydrogen/deuterium exchange mass spectrometry
photon results in an electron jumping to the excited state.
(HXMS) (Ch. 3)
In two-photon excitation, two lower-energy photons arrive
almost instantaneously and induce the electron to jump to r Phosphoproteomics (Ch. 3)
the excited state. (b) Two-photon microscopy can be used
to observe cells up to 1 mm deep within a living animal r Two-photon excitation microscopy (Ch. 4)
immobilized on the microscope stage. (c) Neurons in a lobster r Light-sheet microscopy (Ch. 4)
were imaged using two-photon excitation microscopy.
[Part (c) unpublished data from Peter Kloppenburg and Warren R. Zipfel.] r Super-resolution microscopy (Ch. 4)
viii t PREFACE
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