B. Alberts, A. Johnson, J. Lewis и др. - Molecular Biology of The Cell (6th edition)

Molecular Biology ofTHE CELLSixth EditionMolecular Biology ofTHE CELLSixth EditionBruce AlbertsAlexander JohnsonJulian LewisDavid MorganMartin RaffKeith RobertsPeter WalterWith problems byJohn WilsonTim HuntGarland ScienceVice President: Denise SchanckAssociate Editor: Allie BochicchioProduction Editor and Layout: EJ Publishing ServicesSenior Production Editor: Georgina LucasText Editors: Sherry Granum Lewis and Elizabeth ZayatzIllustrator: Nigel OrmeStructures: Tiago BarrosDesigner: Matthew McClements, Blink Studio, Ltd.Copyeditor: Jo ClaytonProofreader: Sally HuishIndexer: Bill JohncocksPermissions Coordinator: Sheri GilbertBack Cover Photograph: Photography, Christophe Carlinet;Design, Nigel OrmeMolecular Biology of the Cell Interactive Media:Artistic and Scientific Direction: Peter WalterNarration: Julie TheriotDirector of Digital Publishing: Michael MoralesEditorial Assistant: Leah ChristiansProduction Editor: Natasha Wolfe© 2015 by Bruce Alberts, Alexander Johnson, Julian Lewis,David Morgan, Martin Raff, Keith Roberts, and Peter Walter.This book contains information obtained from authenticand highly regarded sources.

Every effort has been made totrace copyright holders and to obtain their permission forthe use of copyright material. Reprinted material is quotedwith permission, and sources are indicated. A wide variety ofreferences are listed. Reasonable efforts have been made topublish reliable data and information, but the author and thepublisher cannot assume responsibility for the validity of allmaterials or for the consequences of their use.All rights reserved.

No part of this book covered by the copyrightherein may be reproduced or used in any format in any form orby any means—graphic, electronic, or mechanical, includingphotocopying, recording, taping, or information storage andretrieval systems—without permission of the publisher.About the AuthorsBruce Alberts received his PhD from Harvard Universityand is the Chancellor’s Leadership Chair in Biochemistryand Biophysics for Science and Education, University ofCalifornia, San Francisco.

He was the editor-in-chief of Sciencemagazine from 2008 until 2013, and for twelve years he servedas President of the U.S. National Academy of Sciences (1993–2005). Alexander Johnson received his PhD from HarvardUniversity and is Professor of Microbiology and Immunologyat the University of California, San Francisco. Julian Lewis(1946–2014) received his DPhil from the University of Oxfordand was an Emeritus Scientist at the London Research Instituteof Cancer Research UK.

David Morgan received his PhD fromthe University of California, San Francisco, and is Professor ofthe Department of Physiology there as well as the Director ofthe Biochemistry, Cell Biology, Genetics, and DevelopmentalBiology Graduate Program.

Martin Raff received his MD fromMcGill University and is Emeritus Professor of Biology at theMedical Research Council Laboratory for Molecular Cell Biologyat University College London. Keith Roberts received his PhDfrom the University of Cambridge and was Deputy Director ofthe John Innes Centre, Norwich. He is Emeritus Professor at theUniversity of East Anglia.

Peter Walter received his PhD fromthe Rockefeller University in New York and is Professor of theDepartment of Biochemistry and Biophysics at the Universityof California, San Francisco, and an Investigator at the HowardHughes Medical Institute. John Wilson received his PhDfrom the California Institute of Technology and pursued hispostdoctoral work at Stanford University.

He is DistinguishedService Professor of Biochemistry and Molecular Biology atBaylor College of Medicine in Houston. Tim Hunt receivedhis PhD from the University of Cambridge where he taughtbiochemistry and cell biology for more than 20 years. He workedat Cancer Research UK until his retirement in 2010. He sharedthe 2001 Nobel Prize in Physiology or Medicine with Lee Hartwelland Paul Nurse.Cover design: Cell biology is not only about the structure andfunction of the myriad molecules that comprise a cell, but alsoabout how this complex chemistry is controlled.

Understandingthe cell’s elaborate regulatory feedback networks will requirequantitative approaches.Library of Congress Cataloging-in-Publication DataAlberts, Bruce, author.Molecular biology of the cell / Bruce Alberts, Alexander Johnson, Julian Lewis, David Morgan, MartinRaff, Keith Roberts, Peter Walter ; with problems by John Wilson, Tim Hunt.

-- Sixth edition.p. ; cm.Preceded by Molecular biology of the cell / Bruce Alberts ... [et al.]. 5th ed. c2008.Includes bibliographical references and index.ISBN 978-0-8153-4432-2 (hardcover) -- ISBN 978-0-8153-4464-3 (paperback)I. Title.[DNLM: 1. Cells. 2. Molecular Biology. QU 300]QH581.2572.8--dc232014031818Published by Garland Science, Taylor & Francis Group, LLC, an informa business,711 Third Avenue, New York, NY 10017, US3 Park Square, Milton Park, Abingdon, OX14 4RN, UKPrinted in the United States of America15 14 13 12 11 10 9 8 7 6 5 4 3 2 1Visit our website at http://www.garlandscience.comJulian Hart LewisAugust 12, 1946—April 30, 2014viiPrefaceSince the last edition of this book appeared, more than five million scientific papershave been published.

There has been a parallel increase in the quantity of digitalinformation: new data on genome sequences, protein interactions, molecular structures, and gene expression—all stored in vast databases. The challenge, for both scientists and textbook writers, is to convert this overwhelming amount of informationinto an accessible and up-to-date understanding of how cells work.Help comes from a large increase in the number of review articles that attemptto make raw material easier to digest, although the vast majority of these reviewsare still quite narrowly focused.

Meanwhile, a rapidly growing collection of onlineresources tries to convince us that understanding is only a few mouse-clicks away.In some areas this change in the way we access knowledge has been highly successful—in discovering the latest information about our own medical problems, forexample. But to understand something of the beauty and complexity of how livingcells work, one needs more than just a wiki- this or wiki- that; it is enormously hardto identify the valuable and enduring gems from so much confusing landfill.

Muchmore effective is a carefully wrought narrative that leads logically and progressivelythrough the key ideas, components, and experiments in such a way that readerscan build for themselves a memorable, conceptual framework for cell biology—a framework that will allow them to critically evaluate all of the new science and,more importantly, to understand it. That is what we have tried to do in MolecularBiology of the Cell.In preparing this new edition, we have inevitably had to make some difficultdecisions.

In order to incorporate exciting new discoveries, while at the same timekeeping the book portable, much has had to be excised. We have added new sections, such as those on new RNA functions, advances in stem cell biology, newmethods for studying proteins and genes and for imaging cells, advances in thegenetics and treatment of cancer, and timing, growth control, and morphogenesisin development.The chemistry of cells is extremely complex, and any list of cell parts and theirinteractions—no matter how complete—will leave huge gaps in our understanding.We now realize that to produce convincing explanations of cell behavior will requirequantitative information about cells that is coupled to sophisticated mathematical/computational approaches—some not yet invented. As a consequence, an emerging goal for cell biologists is to shift their studies more toward quantitative description and mathematical deduction.

We highlight this approach and some of its methods in a new section at the end of Chapter 8.Faced with the immensity of what we have learned about cell biology, it mightbe tempting for a student to imagine that there is little left to discover. In fact, themore we find out about cells, the more new questions emerge. To emphasize thatour understanding of cell biology is incomplete, we have highlighted some of themajor gaps in our knowledge by including What We Don’t Know at the end of eachchapter.

These brief lists include only a tiny sample of the critical unanswered questions and challenges for the next generation of scientists. We derive great pleasurefrom the knowledge that some of our readers will provide future answers.The more than 1500 illustrations have been designed to create a parallel narrative, closely interwoven with the text. We have increased their consistency betweenchapters, particularly in the use of color and of common icons; membrane pumpsand channels are a good example.