Molecular Biology of THE CELL Fifth Edition ALBERTS JOHNSON LEWIS RAFF . (on multicellular systems) in electronic (PDF) form on the accompanying disc. Molecular Biology of the Cell. Fifth Edition. Chapter The Cytoskeleton. Chapter The Cytoskeleton. Copyright © Garland Science Alberts • Johnson. cells and their components are drawn on a logarithmic scale, indicating the range of objects that can be readily resolved by the naked eye and in the light and.
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A companion volume, Molecular Biology of the Cell, Fifth Edition: The in electronic (PDF) form on the accompanying disc, while retaining in the printed volume. Martin Raffreceived his M.D. from McGill University and is at the Medical Research Council Laboratory for Molecular Cell Biology and the Biology Department at. MOLECULAR BIOLOGY OF THE CELL, 5TH EDITION. Editors: Bruce Alberts, PhD; Alexander Johnson, PhD; Julian Lewis, DPhil; Martin Raff MD; Keith Roberts.
DNA bases. Vishu Lingappa University of California. Melvin I. Elliott Margulies National Institutes of Health. Nicholas Harberd John Innes Centre. Robert Horvitz Massachusetts Institute of Technology. Keith Yamamoto University of California.
Julie Theriot Production Design and Development: Bruce Alberts received his Ph. For 12 years, he served as President of the U. National Academy of Sciences — Alexander Johnson received his Ph.
Julian Lewis received his D. Martin Raff received his M. Keith Roberts received his Ph. Peter Walter received his Ph. This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, and sources are indicated. A wide variety of references are listed. Reasonable efforts have been made to publish reliable data and information, but the author and the publisher cannot assume responsibility for the validity of all materials or for the consequences of their use.
All rights reserved. No part of this book covered by the copyright heron may be reproduced or used in any format in any form or by any means—graphic, electronic, or mechanical, including photocopying, recording, taping, or information storage and retrieval systems—without permission of the publisher.
Molecular biology. Alberts, Bruce. M64 Scientists can calculate the age of the Sun and predict when it will cease to shine, but we cannot explain how it is that a human being may live for eighty years but a mouse for only two. We know the complete genome sequences of these and many other species, but we still cannot predict how a cell will behave if we mutate a previously unstudied gene.
Stars may be times bigger, but cells are more complex, more intricately structured, and more astonishing products of the laws of physics and chemistry.
With each edition of this book, we marvel at the new information that cell biologists have gathered in just a few years. But we are even more amazed and daunted at the sophistication of the mechanisms that we encounter. The deeper we probe into the cell, the more we realize how much remains to be understood. In the days of our innocence, working on the first edition, we hailed the identi- fication of a single protein—a signal receptor, say—as a great step forward. Genome sequencing has given us virtu- ally complete molecular parts-lists for many different organisms; genetics and biochemistry have told us a great deal about what those parts are capable of individually and which ones interact with which others; but we have only the most primitive grasp of the dynamics of these biochemical systems, with all their interlocking control loops.
Therefore, although there are great achieve- ments to report, cell biologists face even greater challenges for the future. In this edition, we have included new material on many topics, ranging from epigenetics, histone modifications, small RNAs, and comparative genomics, to genetic noise, cytoskeletal dynamics, cell-cycle control, apoptosis, stem cells, and novel cancer therapies.
As in previous editions, we have tried above all to give readers a conceptual framework for the mass of information that we now have about cells. This means going beyond the recitation of facts. The goal is to learn how to put the facts to use—to reason, to predict, and to control the behavior of living systems. To help readers on the way to an active understanding, we have for the first time incorporated end-of-chapter problems, written by John Wilson and Tim Hunt.
These emphasize a quantitative approach and the art of reasoning from experiments. The Problems Book ISBN , by the same authors, gives com- plete answers to these problems and also contains more than additional problems and solutions. Other ancillaries available for the book include a bank of test questions and lecture outlines, available to qual- ified instructors, and a set of full-color overhead transparencies.
Perhaps the biggest change is in the physical structure of the book. In an effort to make the standard Student Edition somewhat more portable, we are providing Chapters 21—25, covering multicellular systems, in electronic PDF form on the accompanying disc, while retaining in the printed volume Chapters 1—20, covering the core of the usual cell biology curriculum.
But we should emphasize that the final chapters have been revised and updated as thoroughly as the rest of the book and we sincerely hope that they will be read! A Reference Edition ISBN , containing the full set of chapters as printed pages, is also available for those who prefer it. Full details of the conventions adopted in the book are given in the Note to the Reader that follows this Preface. As explained there, we have taken a drastic approach in confronting the different rules for the writing of gene names in dif- ferent species: As always, we are indebted to many people.
Full acknowledgments for sci- entific help are given separately, but we must here single out some exceptionally important contributions: Wallace Marshall and Laura Attardi provided substantial help with Chapters 8 and 20, respectively, as did Maynard Olson for the genomics section of Chapter 4, Xiaodong Wang for Chap- ter 18, and Nicholas Harberd for the plant section of Chapter Denise Schanck directed the whole enterprise and shepherded the wayward authors along the road with wisdom, skill, and kindness.
Nigel Orme put the artwork into its final form and supervised the visual aspects of the book, including the back cover, with his usual flair. Matthew McClements designed the book and its front cover. Emma Jeffcock laid out its pages with extraordinary speed and unflappable effi- ciency, dealing impeccably with innumerable corrections. Eleanor Lawrence and Sherry Granum updated and enlarged the glossary. Jackie Harbor and Sigrid Masson kept us orga- nized. Adam Sendroff kept us aware of our readers and their needs and reactions.
Marjorie Anderson, Bruce Goatly, and Sherry Granum combed the text for obscu- rities, infelicities, and errors. We thank them all, not only for their professional skill and dedication and for efficiency far surpassing our own, but also for their unfailing helpfulness and friendship: Lastly, and with no less gratitude, we thank our spouses, families, friends and colleagues. Without their patient, enduring support, we could not have pro- duced any of the editions of this book.
Cells and Genomes 1 2. Cell Chemistry and Biosynthesis 45 3. DNA, Chromosomes, and Genomes 5. How Cells Read the Genome: From DNA to Protein 7.
Membrane Structure Intracellular Compartments and Protein Sorting Intracellular Vesicular Traffic Energy Conversion: Mitochondria and Chloroplasts Mechanisms of Cell Communication The Cytoskeleton The Cell Cycle Sexual Reproduction: Meiosis, Germ Cells, and Fertilization Development of Multicellular Organisms Pathogens, Infection, and Innate Immunity We would like to thank the following for their suggestions in preparing this edition, as well as those who helped in preparing the first, second, third and fourth editions.
Those who helped on this edition are listed first, those who helped with the first, second, third and fourth editions follow. Chapter 1: Xiaodong Wang [substantial contribution] The Chapter The first three chapters of Part I cover elementary principles and basic biochemistry.
They can serve either as an introduction for those who have not studied biochemistry or as a refresher course for those who have. Merrall-Ross International, Ltd. Permissions Coordinator: Peter Walter Narrated by: Julie Theriot Production Design and Development: Michael Morales. Bruce Alberts received his Ph. For 12 years, he served as President of the U. National Academy of Sciences — Alexander Johnson received his Ph. Julian Lewis received his D. Martin Raff received his M.
Keith Roberts received his Ph. Peter Walter received his Ph. This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, and sources are indicated. A wide variety of references are listed. Reasonable efforts have been made to publish reliable data and information, but the author and the publisher cannot assume responsibility for the validity of all materials or for the consequences of their use.
All rights reserved. No part of this book covered by the copyright heron may be reproduced or used in any format in any form or by any means—graphic, electronic, or mechanical, including photocopying, recording, taping, or information storage and retrieval systems—without permission of the publisher. Molecular biology. Alberts, Bruce. M64 Printed in the United States of America 15 14 13 12 11 10 9 8 7 6 5 4 3 2. Now we appreciate that each protein is generally part of a complex with many others.
This provides hundreds of movies and animations. This means going beyond the recitation of facts. To help readers on the way to an active understanding.
In the days of our innocence. In this edition. Stars may be times bigger. We know the complete genome sequences of these and many other species. As in previous editions. Scientists can calculate the age of the Sun and predict when it will cease to shine. A companion volume. These emphasize a quantitative approach and the art of reasoning from experiments.
Fifth Edition: Through heredity and natural selection. Molecular Biology of the Cell. Genome sequencing has given us virtually complete molecular parts-lists for many different organisms. The deeper we probe into the cell. With each edition of this book. But we are even more amazed and daunted at the sophistication of the mechanisms that we encounter. The goal is to learn how to put the facts to use—to reason.
As always. Jackie Harbor and Sigrid Masson kept us organized. Perhaps the biggest change is in the physical structure of the book. Without their patient. Eleanor Lawrence and Sherry Granum updated and enlarged the glossary.
Matthew McClements designed the book and its front cover. But we should emphasize that the final chapters have been revised and updated as thoroughly as the rest of the book and we sincerely hope that they will be read! Adam Sendroff kept us aware of our readers and their needs and reactions. Wallace Marshall and Laura Attardi provided substantial help with Chapters 8 and Nigel Orme put the artwork into its final form and supervised the visual aspects of the book.
Marjorie Anderson. Xiaodong Wang for Chapter We thank them all. In an effort to make the standard Student Edition somewhat more portable. Full acknowledgments for scientific help are given separately. Other ancillaries available for the book include a bank of test questions and lecture outlines. Full details of the conventions adopted in the book are given in the Note to the Reader that follows this Preface.
Denise Schanck directed the whole enterprise and shepherded the wayward authors along the road with wisdom. Julie Theriot is almost entirely responsible for Chapters 16 Cytoskeleton and 24 Pathogens. Bruce Goatly.
Emma Jeffcock laid out its pages with extraordinary speed and unflappable efficiency. As explained there. Michael Morales managed the transformation of a mass of animations. Cell Adhesion. Amino Acids. Germ Cells. Stem Cells. To Metastasize. Cell Differentiation. But Not All. Used Repeatedly.
Creates All-or-None Outcomes. Ciliated Cells. Pore Formation. Heart Disease. San Francisco. Azim Surani University of Cambridge Chapter 8: Wallace Marshall [major contribution] University of California. Harry Noller University of California. James Haber Brandeis University. Roger Nicoll University of California. Roger Kornberg Stanford University. Douglas Kellogg University of California. Chapel Hill. Donald Voet University of Pennsylvania. Karsten Weis University of California.
Los Angeles. Robert Sauer Massachusetts Institute of Technology. Santa Cruz. Frankfurt am Main. Daniel Finley Harvard University. UK Chapter 4: David Allis The Rockefeller University. Rick Wood University of Pittsburgh Chapter 6: Raul Andino University of California. The Netherlands. Eugene Koonin National Institutes of Health. Eric Green National Institutes of Health.
Christine Guthrie University of California. Aziz Sancar University of North Carolina. Bertil Hille University of Washington. Carol Gross University of California. Richard Ebright Rutgers University. San Francisco Chapter 7: Daniel Schnell University of Massachusetts.
Ulrich Laemmli University of Geneva. Ronald Vale University of California. Steve Harrison Harvard University. Elliott Margulies National Institutes of Health.
Chapter 1: Joan Steitz Yale University. Greg Petsko Brandeis University. Roy Parker University of Arizona. Peter Shaw John Innes Centre. We would like to thank the following for their suggestions in preparing this edition. Susan Wente Vanderbilt University. Louise Johnson University of Oxford. Frank Holstege University Medical Center.
Dresden Chapter Adrian Bird Wellcome Trust Centre. Jonathan Weissman University of California. Geeta Narlikar University of California. Hiten Madhani University of California. David Eisenberg University of California. Jack Szostak Harvard Medical School. Douglas Koshland Carnegie Institution of Washington. Ford Doolittle Dalhousie University. Robert Stroud University of California. Thomas Lindahl Cancer Research. Ueli Schibler University of Geneva.
Nancy Kleckner Harvard University. Gary Felsenfeld National Institutes of Health. Maynard Olson University of Washington Chapter 5: Elizabeth Blackburn University of California. Those who helped on this edition are listed first. Joachim Li University of California. Michael Cox University of Wisconsin.
Steven West Cancer Research. Barbara Panning University of California. Howard Hughes Medical Institute. David Bartel Massachusetts Institute of Technology. Ramanujan Hegde National Institutes of Health. Robert Edwards University of California. Nikolaus Pfanner University of Freiburg. Joseph Gall Carnegie Institution of Washington. Lily Jan University of California. Alexander Varshavsky California Institute of Technology.
Susan Gasser University of Geneva. Quinn Mitrovich University of California. Michael Lynch Indiana University. Amherst Chapter Christopher Mathews Oregon State University. Rodney Rothstein Columbia University. Janet Thornton European Bioinformatics Institute. Patrick Williamson University of Massachusetts.
San Francisco Chapter 9: David Tollervey University of Edinburgh. Arshad Desai University of California. Patricia Calarco University of California. Ralph Steinman The Rockefeller University. Len Stephens The Babraham Institute. Gerald Schatten Pittsburgh Development Center. Douglas Hanahan University of California. Richard Hynes Massachusetts Institute of Technology. Robin Irvine University of Cambridge. Graham Warren Yale University. Renee Reijo University of California.
Bruce Ponder University of Cambridge. San Diego. Martin Humphries University of Manchester. Ira Mellman Yale University. Kansas City. UK Chapter Julie Theriot [major contribution] Stanford University. Anton Berns Netherlands Cancer Institute.
Tony Pawson Mount Sinai Hospital. Hans Clevers Hubrecht Institute. Daniel St. Pat Williamson University of Massachusetts. William Paul National Institutes of.
Morgan Sheng Massachusetts Institute of Technology. David Garrod University of Manchester. Douglas Fearon University of Cambridge. Lewis Lanier University of California. Michael Bishop University of California. Keith Burridge University of North Carolina. Henry Bourne University of California. John Carroll University College London. Neil Hunter University of California. Acknowledgments Medical Center. Bjorn Olsen Harvard Medical School. Holger Gerhardt London Research Institute.
Julie Pitcher University College London. Scott Emr University of California. Stan Falkow Stanford University. Andrew Halestrap University of Bristol. Ian Tomlinson Cancer Research. Richard Locksley University of California. Fiona Watt Cancer Research Institute. Laura Machesky The University of Birmingham. Johnson The Gurdon Institute. Timothy Mitchison Harvard Medical School. John Dick University of Toronto.
Matthew Freeman Laboratory of Molecular Biology. Fred Bunz Johns Hopkins. Diana Myles University of California. Charles Streuli University of Manchester.
Elliott Meyerowitz California Institute of Technology. Michael Yaffe University of California. Laura Attardi [substantial contribution] Stanford University. Harald von Boehmer Harvard Medical School. San Diego Chapter Nicholas Harberd [substantial contribution] John Innes Centre. Ralf Adams London Research Institute. Kenneth Irvine Rutgers University. Larry Goldstein University of California.
Simon Hughes Kings College. Jonathan Pines Gurdon Institute. Roel Nusse Stanford University. Paul Edwards University of Cambridge. Craig Thompson University of Pennsylvania. Julie Ahringer The Gurdon Institute. University of Pennsylvania. Gary Ward University of Vermont Chapter Daniel Portnoy University of California. Stuart Orkin Harvard Medical School. Michael Nussenzweig Rockefeller University. Japan Chapter Lynn Enquist Princeton University.
Charles Streuli The University of Manchester. Caroline Damsky University of California. Richard Harland University of California. Michael Karin University of California. Daniel Goodenough Harvard Medical School. Matthias Falk Lehigh University.
John Couchman Imperial College. Caetano Reis e Sousa Cancer Research. William McGinnis University of California. Kim Nasmyth University of Oxford. Daniel Louvard Institut Curie. Shigekazu Nagata Kyoto University.
James Ferrell Stanford University. Thomas Reh University of Washington. Giampietro Schiavo London Research Institute. Abby Dernburg University of Califonia. Azim Surani The Gurdon Institute. Ken Keegstra Michigan State University. Frankfurt am Main Chapter Michael Gray Dalhousie University. Austin Smith University of Edinburgh. Douglas Green St. Dennis Bray University of Cambridge.
Charles Sherr St. Brigid Hogan Duke University. Jeffrey Gordon Washington University. Ben Glick University of Chicago. Jim Smith The Gurdon Institute. Michael Neuberger University of Cambridge. Michael Glotzer University of Chicago. Rebecca Heald University of California. San Francisco Chapter David Morgan [major contribution] University of California. John Harris University of Otago.
New Zealand. Ruth Ellman Institute of Cancer Research. Leroy Hood Institute for Systems Biology. Carl Branden deceased. Frank Gertler Massachusetts Institute of Technology. Roderick Capaldi University of Oregon. Max Burger University of Basel. Leslie Dale University College London. Graham Hardie University of Dundee. Stuart Ferguson University of Oxford.
John Hopfield Princeton University. Emmanuel Kreidl University of Vienna. Pierre Chambon University of Strasbourg. Martin Brand University of Cambridge. Michael Akam University of Cambridge. David Hanke University of Cambridge. Jonathan Howard University of Washington. Darwin Berg University of California. Ray Evert University of Wisconsin. Comprehensive Cancer Center.
Bernie Gilula deceased. Piet Borst Jan Swammerdam Institute. University of Basel. Stanley Falkow Stanford University. Ira Herskowitz deceased. Michael Grunstein University of California. Larry Fowke University of Saskatchewan. Peter Garland Institute of Cancer Research. Canada First. Beverly Emerson The Salk Institute. Glossary Eleanor Lawrence.
Harvey Florman Tufts University. Brian Gunning Australian National University. Michael Carey University of California. Adelaide Carpenter University of California. Cornelia Bargmann University of California. Charles Cantor Columbia University. Ben Barres Stanford University.
Michael Brown University of Oxford. Russell Doolittle University of California. Lewis Cantley Harvard Medical School. David Haig Harvard University. Reid Gilmore University of Massachusetts. Howard Green Harvard University. Jonathan Ashmore University College London.
Michael Dexter The Wellcome Trust. Leslie Grivell University of Amsterdam.
James Hudspeth The Rockefeller University. Robert Fletterick University of California. Leland Hartwell University of Washington. Charles Gilvarg Princeton University. Sharyn Endow Duke University. Peter Baker deceased. Jeffrey Hall Brandeis University. Adrian Harris Cancer Research. Glenn Herrick University of Utah. Robert Cohen University of California.
Charles Emerson University of Virginia. Hans Bode University of California. Stephen Harrison Harvard University. Robert Horvitz Massachusetts Institute of Technology.
James Crow University of Wisconsin. John Cairns Radcliffe Infirmary. Gerard Evan University of California. Daniel Friend University of California. Alan Grafen University of Oxford. Michael Green University of Massachusetts.
Elaine Fuchs University of Chicago. Ernst Hafen Universitat Zurich. Stony Brook. Nick Rudzik University of Toronto.
Ari Helenius Yale University. Alan Boyde University College London. Judah Folkman Harvard Medical School. Clay Armstrong University of Pennsylvania. Dea Shahinas University of Toronto. John Hall University of Southampton. David Housman Massachusetts Institute of Technology. Nancy Craig Johns Hopkins University.
Julian Downward Cancer Research.
Klaus Rajewsky Harvard Medical School. Tim Hunt Health. Philip Cohen University of Dundee. Michael Ashburner University of Cambridge. John Gerhart University of California. Richard Gardner University of Oxford.
Tariq Enver Institute of Cancer Research. Zach Hall University of California. Jim Dunwell John Innes Institute. Walter Gehring Biozentrum. Martha Arnaud University of California. Tayna Awabdy University of California. Christopher Dobson University of Cambridge. Anthony DeFranco University of California. Roger Cooke University of California.
Alan Hinnebusch National Institutes of Health. Nicholas Harberd John Innes Centre. Frank Grosveld Erasmus Universiteit. University of Amsterdam. Michael Banda University of California. Christine Field Harvard Medical School. John Heath University of Birmingham. David Epel Stanford University. Michael Glotzer University of Vienna. Zacheus Cande University of California. Gary Firestone University of California. Joseph Gall Yale University. Jim Goodrich University of Colorado.
Benny Geiger Weizmann Institute of Science. Merton Bernfield Harvard Medical School. Mario Capecchi University of Utah. Michael Berridge The Babraham Institute. Enrico Coen John Innes Institute. David Baldwin Stanford University. Gerald Fischbach Columbia University. Barbara Meyer University of California. Mark E. Alex Levitzki Hebrew University. Tom Maniatis Harvard University. Mark Marsh Institute of Cancer Research. Richard Losick Harvard University. James Priess University of Washington.
William Otto Cancer Research. Roger Keynes University of Cambridge. Michelle Moritz University of California. Martin Rechsteiner University of Utah. David Lane University of Dundee. Mark Mooseker Yale University. Michael Schramm Hebrew University. Laura Machesky University of Birmingham. Gary Ruvkun Massachusetts General Hospital. Avrion Mitchison University College London.
Vishu Lingappa University of California. Paul National Institutes of Health. Phillips Robbins Massachusetts Institute of Technology. David Nicholls University of Dundee. Paul Nurse Cancer Research. Dale Purves Duke University. Jane Langdale University of Oxford. Nelson University of Illinois. Alan Munro University of Cambridge. William W. Richard Myers Stanford University. Tom Kornberg University of California. John Kuriyan University of California.
Mitchison University College London. Colin Manoil Harvard Medical School. George Ratcliffe University of Oxford. Joshua Sanes Harvard University. Laurence Hurst University of Bath.
Elliot Meyerowitz California Institute of Technology. Gordon Peters Cancer Research. Stephanie Mel University of California. Charles Janeway deceased. Daniel Koshland University of California. Elaine Robson University of Reading.
Frank McNally University of California. Norman Iscove Ontario Cancer Institute. Richard Scheller Stanford University. Gail Martin University of California. Montrose Moses Duke University. Gottfried Schatz Biozentrum.
Philip Ingham University of Sheffield. George Palade deceased. Ray Keller University of California. Jeremy Hyams University College London. Tom Jessell Columbia University. Salt Lake City. Elio Raviola Harvard Medical School. Robert Perry Institute of Cancer Research. Richard Klausner National Institutes of Health.
Chris Miller Brandeis University. Marc Kirschner Harvard University. Joel Rosenbaum Yale University. Janet Rossant Mount Sinai Hospital. Alan Sachs University of California.
Efraim Racker Cornell University. Trevor Lamb University of Cambridge. Anne Mudge University College London. Dale Oxender University of Michigan. David Sabatini New York University. Jodi Nunnari University of California. Keith Mostov University of California. Michael Levine University of California. Louis Reichardt University of California.
Peter Sarnow Stanford University. Tom Pollard Yale University. Ron Kaback University of California. Lefkowitz Duke University. Klaus Rajewsky University of Cologne. Howard Schachman University of California. Robert Roeder The Rockefeller University. Parson University of Washington.
Conly Rieder Wadsworth Center. Peter Mombaerts The Rockefeller University. Darwin Prockop Tulane University. Warren Levinson University of California. Kelly Komachi University of California. Marilyn Kozak University of Pittsburgh. Acknowledgments Cancer Research. William E. Juan Korenbrot University of California. Mark Krasnow Stanford University. Shirley Lowe University of California. David Ish-Horowicz Cancer Research. Richard McCarty Cornell University. David Phillips The Rockefeller University.
Mike Klymkowsky University of Colorado. James Schwartz Columbia. John Owen University of Birmingham. Stuart Kornfeld Washington University. Andy Johnston John Innes Institute. Robert Kingston Massachusetts General Hospital.
Jay Lash University of Pennsylvania. Walter Neupert University of Munich. Giampietro Schiavo Cancer Research. Regis Kelly University of California. Freiderick Meins Freiderich Miescher Institut. Clive Lloyd John Innes Institute. Tim Mitchison Harvard Medical School. Robert J. Randy Schekman University of California.
Jesse Roth National Institutes of Health. David Morgan University of California. Suzanne Noble University of California. Jordan Queen Elizabeth College. Dan Portnoy University of California. Tomas Lindahl Cancer Research. Cynthia Kenyon University of California. Fred Richards Yale University. Hartmut Land Cancer Research. Robert Mishell University of Birmingham. Judith Kimble University of Wisconsin. Brian McCarthy University of California.