Table Of ContentMolecular Biology of
THE CELL
Sixth Edition
The Problems Book
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to match pagination of print book
Molecular Biology of
THE CELL
Sixth Edition
The Problems Book
John Wilson and Tim Hunt
Garland Science
Vice President: Denise Schanck
Associate Editor: Allie Bochicchio
Production Editor and Layout: EJ Publishing Services
Senior Production Editor: Georgina Lucas
Master Reviewer: Alastair Ewing
Copyeditor: Jo Clayton
Proofreader: Sally Huish
Illustrator: Nigel Orme
Designer: Matthew McClements, Blink Studio, Ltd.
Back cover photograph and design: Nigel Orme
Indexer: Medical Indexing Ltd.
Permissions Coordinator: Sheri Gilbert
© 2015 by John Wilson and Tim Hunt
John Wilson received his PhD from the California Institute of Technology and
pursued his postdoctoral work at Stanford University. He is currently Distinguished
Service Professor of Biochemistry and Molecular Biology at Baylor College of
Medicine in Houston. His research interests include genome instability and gene
therapy. He has taught medical and graduate students for many years, coauthored
books on immunology, molecular biology, and biochemistry, and received
numerous teaching honors, including the Robertson Presidential Award for
excellence in education.
Tim Hunt received his PhD from the University of Cambridge where he taught
biochemistry and cell biology for more than 20 years. In the late 1970s and early
1980s he spent his summers teaching cell and molecular biology at the Marine
Biological Laboratory, Woods Hole, Massachusetts. He left Cambridge in 1990 and
moved to the Cancer Research UK Clare Hall Laboratories, just outside London,
where he worked on the control of the cell cycle until his retirement in 2010.
He shared the 2001 Nobel Prize in Physiology or Medicine with Lee Hartwell
and Paul Nurse.
This book contains information obtained from authentic and highly regarded
sources. Every effort has been made to trace copyright holders and to obtain their
permission for the use of copyright material. 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 herein 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.
ISBN: 978-0-8153-4453-7 (paperback)
Published by Garland Science, Taylor & Francis Group, LLC, an informa business,
711 Third Avenue, New York, NY 10017, US
3 Park Square, Milton Park, Abingdon, OX14 4RN, UK
Printed in the United States of America
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Visit our website at http://www.garlandscience.com
We dedicate this book to the memory of our comrade, Julian Lewis.
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vii
Preface
Welcome to he Problems Book, which aims to help students appreciate the ways
in which an understanding of how cells work, as discussed in Molecular Biology
of the Cell, Sixth Edition, by Alberts et al., can be further explored through experi-
ments and simple calculations. As always, we hope to stimulate our readers to
ask questions as well as to learn and digest the stories that “the big book” tells. In
real life, knowledge and understanding come from research, which entails curios-
ity, puzzlement, doubt, criticism, and debate as well as performing experiments.
Groping one’s way through the fog of uncertainty during a project is a slow and
often discouraging process; eureka moments (even if one is lucky) are few and far
between. Nevertheless, those moments catch the essence of the drama, and we
have tended to focus on them, where we have been able to cast them in the form
of a problem. In this way, for student and teacher alike, we hope to encourage a
questioning attitude to biology. Without curiosity there would be neither science
nor scientists.
We have been making up problems together since 1985, and the revision lead-
ing to this new edition of he Problems Book has taken us more than four years.
here are several new things about this edition. First, the book is now in color. As
well as improving its look as a whole, we think this will improve the clarity and
intelligibility of the igures. Second, we’ve added a new type of question, MCAT
Style, modeled on the kind of problems that are found in most medical school
admissions tests. hese were drafted by Doug Kellogg at the University of Cali-
fornia, Santa Cruz, and we think they make a great addition to the book. Indeed,
we were pleasantly surprised to discover that these questions allowed us to frame
problems in new and interesting ways. Elsewhere, we have done a considerable
amount of pruning, partly to make space for these new problems and partly to
eliminate problems that were showing their age or were no longer relevant to the
parent text.
he organization of he Problems Book remains largely the same. here are
Terms to Learn, Deinitions, and True/False sections in every chapter. Next come
hought Problems, of which some are more challenging than others—they may
be playful, or serious, but all are designed to make the reader think. Following
these is a section called Calculations, designed to deal with quantitative aspects
of cell biology. he calculations in this book are mostly very straightforward, usu-
ally involving no more than the interconversion of units, yet they provide a solid
framework for thinking about the cell. Are cell-surface receptors sparse in the
plasma membrane, or jam-packed? Do molecules difuse across a cell slowly, or
in the blink of an eye? Does chromatin occupy most of the nuclear volume, or
just a tiny fraction? How fast could a tomato plant grow, theoretically? Numeri-
cal analysis of such questions is very important if one is to gain an understand-
ing for the molecular basis of cell biology. he Data Handling section contains
research-based problems. Our original brief was to compose problems based on
experiments so as to allow readers to get a better feel for the way in which biologi-
cal knowledge is obtained. It is tremendously important to keep asking, “How do
we know that? What’s the evidence?” or to wonder how one might go about ind-
ing something out. Often it’s not at all obvious, often the initial breakthrough was
a lucky chance observation, made while investigating some completely diferent
viii PREFACE
business. In fact, it takes most of us years of research experience to grasp the idea
of how one simple fact “can illuminate a distant area, hitherto dark” (Boveri,
1902). Seeing how these tiny shards of evidence give rise to the big picture often
involves considerable imagination as well as a certain discipline, to know how
much weight the evidence will bear. We hope we have sometimes, at least, been
able to capture the essence of how experiments lead to understanding. To do
justice to the authors of the experiments we use in these problems, however, we
strongly recommend recourse to the original papers, whose references we always
provide.
A newly compiled section, Medical Links, contains problems of particular
interest to health science students.
We hope that the organization and classiication of problems will help both
student and teacher to ind what they are looking for. How should this book be
used? We composed it by a process of constant dialogue and discussion, and we
suspect that the most fruitful use of the problems will be to stimulate discussions
in class, or between students. Tackling selected problems as homework will also
surely help. Teachers have told us that they ind ideas for exam questions here,
and all the answers to our questions are now provided in he Problems Book, for
many of these problems are diicult to answer and are not intended to be set as
tests. Rather, we hope that readers will be intrigued (as we were) by the questions
we ask, and after thinking a bit will want to see what the answer is, what form the
discussion takes, and how to get at thinking about this particular kind of a prob-
lem.
he answers to the end-of-chapter problems in Molecular Biology of the Cell,
Sixth Edition can also be found in the back of this book, including the answers to
newly written problems for Chapters 21 through 24.
As always, we want to hear from our readers, for despite our best eforts, we do
not always get things right. Please email John Wilson at [email protected] or Tim
Hunt at [email protected] with your comments or queries, and we’ll do our
best to answer them.
ix
Acknowledgments
It remains true that our rate of production, over the years, averages at about one
chapter per year, but even this glacial progress would not have been possible
without a tremendous amount of help from friends and colleagues whose names
are recorded in previous editions of he Problems Book, which appeared in 1989,
1994, 2002, and 2008. As ever, we are indebted to Alastair Ewing who worked
through all the new problems, discovering embarrassing mistakes and inding
better, clearer, and more graceful ways of putting things. Denise Schanck has been
a tower of strength, as always, and Emma Jefcock a brilliant and startlingly ei-
cient designer and friend. Allie Bochicchio coordinated our activities and kept us
in good order. Mike Morales helped to set up an instant home-away-from-home
during meetings in California, and his cheerful humor helped tremendously.
Adam Sendrof, who took care of marketing and gave us audience feedback, was
unfailingly supportive. We are especially grateful to all the authors of Molecular
Biology of the Cell, who have been extremely helpful in the selection and reine-
ment of problems that appear in the main text. We thank them most warmly for
their suggestions. Once again, Nigel Orme has been a huge help with the illustra-
tions, particularly in the business of adding color. he person who made the most
important contribution to this edition, however, is Doug Kellogg of the Univer-
sity of California, Santa Cruz. Doug is an old friend who allowed himself to be
persuaded to take on the task of composing MCAT-style questions, despite his
heavy responsibilities as a teacher, researcher, and new father. We are very lucky,
however, to have the loving support of our spouses and families, Lynda, Mary,
Celia, and Aggie. hey’ve put up heroically with our regular absences and preoc-
cupations.
