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Molecular Biology Of the Cell by Alberts
5th Edition pdf file


In many respects, we understand the structure of the universe better than the
workings of living cells. 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
genomes equenceso f thesea nd many other speciesb, ut we still cannot predict
how a cell will behave if we mutate a previously unstudied gene. Stars may be
l0a3 times bigger, but cells are more complex, more intricately structured, and
more astonishing products of the laws of physics and chemistry. Through heredity
and natural selection, operating from the beginnings of life on Earth to the
presentd ay-that is, for about 20Voo f the ageo f the universe-living cellsh ave
been progressively refining and extending their molecular machinery and
recording the results of their experiments in the genetic instructions they pass
on to their progeny.
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 reafize how much remains to be understood.
In the days of our innocence, working on the first edition, we hailed the identification
of a single protein-a signal receptol say-as a great step forward' Now
we appreciate that each protein is generally part of a complexwith many others,
working together as a system, regulating one another's activities in subtle ways,
and held in specific positions by binding to scaffold proteins that give the chemical
factory a definite spatial structure. Genome sequencing has given us virtually
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 achievements
to report, cell biologistsf ace eveng reaterc hallengesf or the future.
In this edition, we have included new material on many topics, ranging from
epigeneticsh, istonem odificationss, mall RNAs,a nd comparativeg enomicst,o
geneticn oise,c ytoskeletadl lmamics,c ell-cyclec ontrol, apoptosis,s tem 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 incorporatede nd-of-chapterp roblems,w ritten by Iohn Wilson and Tim
Hunt. These emphasize a quantitative approach and the art of reasoning from
experiments. A companion volume, Molecular Biology of the CelI, Fifth Edition:
TheP roblemsB ook0 SBN9 78-0-8153-4110-9b)y, t he samea uthors,g ivesc omplete
answerst o thesep roblemsa nd alsoc ontainsm ore than 1700a dditional
problems and solutions.
A further major adjunct to the main book is the attached Media DVD-ROM
disc. This provides hundreds of movies and animations, including manythat are
new in this edition, showing cells and cellular processesin action and bringing
the text to life; the disc also now includes all the figures and tables from the main
book,p re-loadedin to PowerPoint@pr esentationsO. thera ncillariesa vailablefo r
the book include a bank of test questions and lecture outlines, available to qualified
instructors,a nd a seto f 200f ull-coloro verheadtr ansparencies.
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 2r-25, covering multicellular systems, in electronic (pDF)
form on the accompanying disc, while retaining in the printed volume chapters
l-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 (ISBN9 7s-0-8153-4r11-6c)o, ntainingt he full seto f chaptersa sp rinred
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 Readert hat follows this PrefaceA. s explainedt here,w e have taken a drastic
approach in confronting the different rules for the writing of gene names in different
species: throughout this book, we use the same style, regardless of
speciesa, nd often in defianceo fthe usuals pecies-specifcico nventions.
As always,w e are indebted to many people. Full acknowledgmentsfo r scientific
help are given separatelyb, ut we must here singleo ut somee xceptionally
important contributions: Iulie Theriot is almost entirely responsible for chapters
16 (cytoskeleton)a nd 24 (PathogensI,n fection, and Innate Immunity), and
David Morgan likewise for chapter 17 (cell cycle). wallace Marshall and Laura
Attardi provided substantialh elp with chapters 8 and 20, respectively,a s did
Maynardo lson for the genomicss ectiono f chapter4 ,X iaodongwangf or chapter
18, and Nicholas Harberd for the plant section of Chapter 15.
we also owe a huge debt to the staff of Garland science and others who
helped convert writers' efforts into a polished final product. 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 supervisedt he visuala spectso f the book,i ncluding the backc over,w ith
his usual flair. Matthew Mcclements designed the book and its front cover.
Emma Jeffcockla id out its pagesw ith extraordinarys peeda nd unflappablee fficiency,
d ealingi mpeccablywith innumerablec orrections.M ichaelM oralesm anaged
the transformation of a mass of animations, video clips, and other materials
into a user-friendly DVD-ROM. Eleanor Lawrence and sherry Granum
updatedand enlargedt he glossaryJ. ackieH arbor and SigridM assonk ept us organized.
Adam Sendroffkept us aware ofour readers and their needs and reactions.
Marjorie Anderson, Bruce Goatly, and sherry Granum combed the text for obscurities,
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 helpftrlnessa nd friendship:t hey havem adei t a pleasuret o work on the
Lastly, and with no less gratitude, we thank our spouses, families, friends
and colleaguesw. ithout their patient,e ndurings upport,w e couldn ot havep roduced
any of the editions of this book.



1 Cells and Genomes
2 Cell Chemistry and Biosynthesis
3 Proteins


4 DNA, Chromosomes and Genomes
5 DNA Replication, Repair, and Recombination
6 How Cells Read the Genome: From DNA to Protein
7 Control of Gene Expression


8 Manipulating Proteins, DNA, and RNA
9 Visualizing Cells


10 Membrane Structure
11 Membrane Transport of Small Molecules and the Electrical Properties of Membranes
12 Intracellular Compartments and Protein Sorting
13 Intracellular Vesicular Traffic
14 Energy Conversion Mitochondria and Chloroplasts
15 Mechanisms of Cell Communication
16 The Cytoskeleton
17 The Cell Cycle
18 Apoptosis


19 Cell lunctions, Cell Adhesion, and the Extracellular Matrix
20 Cancer

Structure of the Book

Although the chapters of this book can be read independently of one another,
they are arranged in a logical sequence of five parts. 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.

Part II deals with the storage, expression and transmission of genetic information.

Part III deals with the principles of the main experimental methods for
investigating cells. It is not necessary to read these two chapters in order to
understand the later chapters, but a reader will find it a useful reference.

Part IV discusses the internal organization of the cell.

Part V follows the behavior of cells in multicellular systems, starting with
cell-cell junctions and extracellular matrix and concluding with tvvo chapters on
the immune system.

A selection of problems, written by Iohn Wilson and Tim Hunt, now appears in
the text at the end of each chapter. The complete solutions to these problems
can be found in Molecular Biology of the CelI, Fifth Edition: The Problems Book.


A concise list of selected references is included at the end of each chapter. These
are arranged in alphabetical order under the main chapter section headings.
These references frequently include the original papers in which important discoveries
were first reported. Chapter 8 includes several tables giving the dates of
crucial developments along with the names of the scientists involved. Elsewhere
in the book the policy has been to avoid naming individual scientists.
Media Codes
Media codes are integrated throughout the text to indicate when relevant videos
and animations are available on the DVD-ROM. The four-letter codes are
enclosed in brackets and highlighted in color, like this . The interface for
the CeII Biology Interactiue media player on the DVD-ROM contains a window
where you enter the 4-letter code. lVhen the code is typed into the interface, the
corresponding media item will load into the media player.
Throughout the book, boldface type has been used to highlight key terms at the
point in a chapter where the main discussion of them occurs. Italic is used to set
off important terms with a lesser degree of emphasis. At the end of the book is
the expanded glossary, covering technical terms that are part of the common
currency of cell biology; it is intended as a first resort for a reader who encounters
an unfamiliar term used without explanation.
Nomenclature for Genes and Proteins
Each species has its own conventions for naming genes; the only common feature
is that they are always set in italics. In some species (such as humans)' gene
names are spelled out all in capital letters; in other species (such as zebrafish),
case and rest in lower case; or (as in Drosophila) with different combinations of
upper and lower case, according to whether the first mutant allele to be discovered
gave a dominant or recessive phenotype. conventions for naming protein
products are equally varied.
This typographical chaos drives everyone crazy. lt is not just tiresome and
absurd; it is also unsustainable. we cannot independently define a fresh convention
for each of the next few million species whose genes we may wish to
study. Moreover, there are many occasions, especially in a book such as this,
where we need to refer to a gene generically, without specifliing the mouse version,
the human version, the chick version, or the hippopotamus version,
because they are all equivalent for the purposes of the discussion. .A/hatc onvention
then should we use?
We have decided in this book to cast aside the conventions for individual
species and follow a uniform rule: we write all gene names, like the names of people
and places, with the first letter in upper case and the rest in lower case, but all
in- italics, thus: Apc, Bazooka, cdc2, Disheuelled, Egll. The corresponding protein,
where it is named after the gene, will be written in the same way, but in roman
rather than italic letters: Apc, Bazooka, cdc2, Dishevelled, Egll. lvhen it is necessary
to specify the organism, this can be done with a prefix to the gene name.
For completeness, we list a few further details of naming rules that we shall
follow In some instances an added letter in the gene name is traditionally used
to distinguish between genes that are related by function or evolution; for those
genes we put that letter in upper case if it is usual to do so (LacZ, RecA, HoxA4).
we use no hyphen to separate added letters or numbers from the rest of the
name. Proteins are more of a problem. Many of them have names in their own
right, assigned to them before the gene was named. such protein names take
many forms, although most of them traditionally begin with a lower-case letter
(actin, hemoglobin, catalase), Iike the names of ordinary substances (cheese,
nylon), unless they are acronyms (such as GFB for Green Fluorescent protein, or
BMP4, for Bone Morphogenetic Protein #4).To force all such protein names into
a uniform style would do too much violence to established usages, and we shall
simply write them in the traditional way (actin, GFB etc.). For thl corresponding
gene names in all these cases, we shall nevertheless follow our standard rule:
Actin, Hemoglobin, catalase, Bmp4, G/p. occasionally in our book we need to
highlight a protein name by setting it in italics for emphasis; the intention will
generally be clear from the context.

Library of Congress Cataloging-in-Publication Data
Molecular biology of the cell / Bruce Alberts ... [et al.].-- 5th ed.
ISBN9 78-0-8153-4r05-(5h ardcover-)- -ISBN9 78-0-8f5 g-4t06_Z( paperback)
L Cytology. 2. Molecular biology. I. Alberts, Bruce.
QHsB1.2.M 642 008
2007005475 CIP
Published by Garland science, Taylor & Francis Group, LLC, an informa business,
270 Madison Avenue, NewYork NY f 0016, USA, and 2 park Square, Milton park,
Abingdon, OXl4 4RN, UK.
Printed in the United States of Am

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