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 ESSENTIAL
CELL BIOLOGY

FIFTH EDITION
 FI F T H
ED I T I O N

ESSENTIAL

CELL BIOLOGY
Bruce Alberts
UNIVERSIT Y OF CALIFORNIA, SAN FRANCISCO

Karen Hopkin
SCIENCE WRITER

Alexander Johnson
UNIVERSIT Y OF CALIFORNIA, SAN FRANCISCO

David Morgan
UNIVERSIT Y OF CALIFORNIA, SAN FRANCISCO

Martin Raff
UNIVERSIT Y COLLEGE LONDON (EMERITUS)

Keith Roberts
UNIVERSIT Y OF EAST ANGLIA (EMERITUS)

Peter Walter
UNIVERSIT Y OF CALIFORNIA, SAN FRANCISCO

n 
W . W . N O R T O N & C O M PA N Y
NE W YORK • LONDON
W. W. Norton & Company has been independent since its founding in 1923, when William
Warder Norton and Mary D. Herter Norton first published lectures delivered at the Peo-
ple’s Institute, the adult education division of New York City’s Cooper Union. The firm soon
expanded its program beyond the Institute, publishing books by celebrated academics
from America and abroad. By midcentury, the two major pillars of Norton’s publishing
program—trade books and college texts—were firmly established. In the 1950s, the Norton
family transferred control of the company to its employees, and today—with a staff of four
hundred and a comparable number of trade, college, and professional titles published
each year—W. W. Norton & Company stands as the largest and oldest publishing house
owned wholly by its employees.

Copyright © 2019 by Bruce Alberts, Dennis Bray, Karen Hopkin, Alexander Johnson, the Estate of Julian
Lewis, David Morgan, Martin Raff, Nicole Marie Odile Roberts, and Peter Walter

All rights reserved

Printed in Canada

Editors: Betsy Twitchell and Michael Morales

Associate Editor: Katie Callahan
Editorial Consultant: Denise Schanck
Senior Associate Managing Editor, College: Carla L. Talmadge
Editorial Assistants: Taylere Peterson and Danny Vargo
Director of Production, College: Jane Searle
Managing Editor, College: Marian Johnson
Managing Editor, College Digital Media: Kim Yi
Media Editor: Kate Brayton
Associate Media Editor: Gina Forsythe
Media Project Editor: Jesse Newkirk
Media Editorial Assistant: Katie Daloia
Ebook Production Manager: Michael Hicks
Content Development Specialist: Todd Pearson
Marketing Manager, Biology: Stacy Loyal
Director of College Permissions: Megan Schindel
Permissions Clearer: Sheri Gilbert
Composition: Emma Jeffcock of EJ Publishing Services
Illustrations: Nigel Orme
Design Director: Hope Miller Goodell
Designer: Matthew McClements, Blink Studio, Ltd.
Indexer: Bill Johncocks
Manufacturing: Transcontinental Interglobe—Beauceville, Quebec

Permission to use copyrighted material is included alongside the appropriate content.

Library of Congress Cataloging-in-Publication Data

Names: Alberts, Bruce, author.

Title: Essential cell biology / Bruce Alberts, Karen Hopkin, Alexander
Johnson, David Morgan, Martin Raff, Keith Roberts, Peter Walter.
Description: Fifth edition. | New York : W.W. Norton & Company, [2019] |
Includes index.
Identifiers: LCCN 2018036121 | ISBN 9780393679533 (hardcover)
Subjects: LCSH: Cytology. | Molecular biology. | Biochemistry.
Classification: LCC QH581.2 .E78 2019 | DDC 571.6—dc23 LC record available at
https://lccn.loc.gov/2018036121

W. W. Norton & Company, Inc., 500 Fifth Avenue, New York, NY 10110
wwnorton.com
W. W. Norton & Company Ltd., 15 Carlisle Street, London W1D 3BS

1 2 3 4 5 6 7 8 9 0
PREFACE

Nobel Prize–winning physicist Richard Feynman once noted that nature

has a far, far better imagination than our own. Few things in the universe
illustrate this observation better than the cell. A tiny sac of molecules
capable of self-replication, this marvelous structure constitutes the fun-
damental building block of life. We are made of cells. Cells provide all
the nutrients we consume. And the continuous activity of cells makes
our planet habitable. To understand ourselves—and the world of which
we are a part—we need to know something of the life of cells. Armed
with such knowledge, we—as citizens and stewards of the global
community—will be better equipped to make well-informed decisions
about increasingly sophisticated issues, from climate change and food
security to biomedical technologies and emerging epidemics.
In Essential Cell Biology we introduce readers to the fundamentals of
cell biology. The Fifth Edition introduces powerful new techniques that
allow us to examine cells and their components with unprecedented
precision—such as super-resolution fluorescence microsocopy and
cryoelectron microscopy—as well as the latest methods for DNA
sequencing and gene editing. We discuss new thinking about how cells
organize and encourage the chemical reactions that make life possible,
and we review recent insights into human origins and genetics.
With each edition of Essential Cell Biology, its authors re-experience the
joy of learning something new and surprising about cells. We are also
reminded of how much we still don’t know. Many of the most fascinat-
ing questions in cell biology remain unanswered. How did cells arise on
the early Earth, multiplying and diversifying through billions of years of
evolution to fill every possible niche—from steaming vents on the ocean
floor to frozen mountaintops—and, in doing so, transform our planet’s
entire environment? How is it possible for billions of cells to seamlessly
cooperate and form large, multicellular organisms like ourselves? These
are among the many challenges that remain for the next generation of
cell biologists, some of whom will begin a wonderful, lifelong journey
with this textbook.
Readers interested in learning how scientific inquisitiveness can fuel break-
throughs in our understanding of cell biology will enjoy the stories of dis-
covery presented in each chapter’s “How We Know” feature. Packed with
experimental data and design, these narratives illustrate how biologists
tackle important questions and how experimental results shape future
ideas. In this edition, a new “How We Know” recounts the discoveries that
first revealed how cells transform the energy locked in food molecules into
the forms used to power the metabolic reactions on which life depends.
As in previous editions, the questions in the margins and at the end of
each chapter not only test comprehension but also encourage careful
thought and the application of newly acquired information to a broader
biological context. Some of these questions have more than one valid
v
vi Preface

answer and others invite speculation. Answers to all of the questions

are included at the back of the book, and many provide additional
information or an alternative perspective on material presented in the
main text.
More than 160 video clips, animations, atomic structures, and high-
resolution micrographs complement the book and are available online.
The movies are correlated with each chapter and callouts are highlighted
in color. This supplemental material, created to clarify complex and critical
concepts, highlights the intrinsic beauty of living cells.

For those who wish to probe even more deeply, Molecular Biology of
the Cell, now in its sixth edition, offers a detailed account of the life of
the cell. In addition, Molecular Biology of the Cell, Sixth Edition: A Prob-
lems Approach, by John Wilson and Tim Hunt, provides a gold mine of
thought-provoking questions at all levels of difficulty. We have drawn
upon this tour-de-force of experimental reasoning for some of the ques-
tions in Essential Cell Biology, and we are very grateful to its authors.

Every chapter of Essential Cell Biology is the product of a communal effort:

both text and figures were revised and refined as drafts circulated from
one author to another—many times over and back again! The numer-
ous other individuals who have helped bring this project to fruition are
credited in the Acknowledgments that follow. Despite our best efforts, it
is inevitable that errors will have crept into the book, and we encourage
eagle-eyed readers who find mistakes to let us know, so that we can
correct them in the next printing.

Acknowledgments
The authors acknowledge the many contributions of professors and
students from around the world in the creation of this Fifth Edition. In
particular, we received detailed reviews from the following instructors
who had used the fourth edition, and we would like to thank them for
their important contributions to our revision:
Delbert Abi Abdallah, Thiel College, Pennsylvania
Ann Aguanno, Marymount Manhattan College
David W. Barnes, Georgia Gwinnett College
Manfred Beilharz, The University of Western Australia
Christopher Brandl, Western University, Ontario
Marion Brodhagen, Western Washington University
David Casso, San Francisco State University
Shazia S. Chaudhry, The University of Manchester, United Kingdom
Ron Dubreuil, The University of Illinois at Chicago
Heidi Engelhardt, University of Waterloo, Canada
Sarah Ennis, University of Southampton, United Kingdom
David Featherstone, The University of Illinois at Chicago
Yen Kang France, Georgia College
Barbara Frank, Idaho State University
Daniel E. Frigo, University of Houston
Marcos Garcia-Ojeda, University of California, Merced
David L. Gard, The University of Utah
Adam Gromley, Lincoln Memorial University, Tennessee
Elly Holthuizen, University Medical Center Utrecht, The Netherlands
Harold Hoops, The State University of New York, Geneseo
Bruce Jensen, University of Jamestown, North Dakota
Andor Kiss, Miami University, Ohio
Annette Koenders, Edith Cowan University, Australia
Arthur W. Lambert, Whitehead Institute for Biomedical Research
Denis Larochelle, Clark University, Massachusetts
David Leaf, Western Washington University
Esther Leise, The University of North Carolina at Greensboro
Bernhard Lieb, University of Mainz, Germany
 Preface vii

Julie Lively, Louisiana State University

Caroline Mackintosh, University of Saint Mary, Kansas
John Mason, The University of Edinburgh, Scotland
Craig Milgrim, Grossmont College, California
Arkadeep Mitra, City College, Kolkata, India
Niels Erik Møllegaard, University of Copenhagen
Javier Naval, University of Zaragoza, Spain
Marianna Patrauchan, Oklahoma State University
Amanda Polson-Zeigler, University of South Carolina
George Risinger, Oklahoma City Community College
Laura Romberg, Oberlin College, Ohio
Sandra Schulze, Western Washington University
Isaac Skromne, University of Richmond, Virginia
Anna Slusarz, Stephens College, Missouri
Richard Smith, University of Tennessee Health Science Center
Alison Snape, King’s College London
Shannon Stevenson, University of Minnesota Duluth
Marla Tipping, Providence College, Rhode Island
Jim Tokuhisa, Virginia Polytechnic Institute and State University
Guillaume van Eys, Maastricht University, The Netherlands
Barbara Vertel, Rosalind Franklin University of Medicine and Science, Illinois
Jennifer Waby, University of Bradford, United Kingdom
Dianne Watters, Griffith University, Australia
Allison Wiedemeier, University of Louisiana at Monroe
Elizabeth Wurdak, St. John’s University, Minnesota
Kwok-Ming Yao, The University of Hong Kong
Foong May Yeong, National University of Singapore
We are also grateful to those readers who alerted us to errors that they
found in the previous edition.
Working on this book has been a pleasure, in part due to the many people
who contributed to its creation. Nigel Orme again worked closely with
author Keith Roberts to generate the entire illustration program with his
usual skill and care. He also produced all of the artwork for both cover
and chapter openers as a respectful digital tribute to the “squeeze-bottle”
paintings of the American artist Alden Mason (1919–2013). As in previ-
ous editions, Emma Jeffcock did a brilliant job in laying out the whole
book and meticulously incorporated our endless corrections. We owe a
special debt to Michael Morales, our editor at Garland Science, who co-
ordinated the whole enterprise. He oversaw the initial reviewing, worked
closely with the authors on their chapters, took great care of us at numer-
ous writing meetings, and kept us organized and on schedule. He also
orchestrated the wealth of online materials, including all video clips
and animations. Our copyeditor, Jo Clayton, ensured that the text was
stylistically consistent and error-free. At Garland, we also thank Jasmine
Ribeaux, Georgina Lucas, and Adam Sendroff.
For welcoming our book to W. W. Norton and bringing this edition to
print, we thank our editor Betsy Twitchell, as well as Roby Harrington,
Drake McFeely, Julia Reidhead, and Ann Shin for their support. Taylere
Peterson and Danny Vargo deserve thanks for their assistance as
the book moved from Garland to Norton and through production.
We are grateful to media editor Kate Brayton and content develop-
ment specialist Todd Pearson, associate editors Gina Forsythe and
Katie Callahan, and media editorial assistant Katie Daloia whose
coordination of electronic media development has resulted in an un-
matched suite of resources for cell biology students and instructors
alike. We are grateful for marketing manager Stacy Loyal’s tireless
enthusiasm and advocacy for our book. Megan Schindel, Ted Szczepanski,
and Stacey Stambaugh are all owed thanks for navigating the permis-
sions for this edition. And Jane Searle’s able management of produc-
tion, Carla Talmadge’s incredible attention to detail, and their shared
knack for troubleshooting made the book you hold in your hands
a reality.
viii Preface

Denise Schanck deserves extra special thanks for providing continuity

as she helped shepherd this edition from Garland to Norton. As always,
she attended all of our writing retreats and displayed great wisdom in
orchestrating everything she touched.
Last but not least, we are grateful, yet again, to our colleagues and our
families for their unflagging tolerance and support. We give our thanks
to everyone in this long list.

Resources for Instructors

and Students
INSTRUCTOR RESOURCES
wwnorton.com/instructors

Smartwork5
Smartwork5 is an easy-to-use online assessment tool that helps stu-
dents become better problem solvers through a variety of interactive
question types and extensive answer-specific feedback. All Smartwork5
questions are written specifically for the book, are tagged to Bloom’s
levels and learning objectives, and many include art and animations.
Get started quickly with our premade assignments or take advantage
of Smartwork5’s flexibility by customizing questions and adding your
own content. Integration with your campus LMS saves you time by al-
lowing Smartwork5 grades to report right to your LMS gradebook, while
individual and class-wide performance reports help you see students’
progress.

Interactive Instructor’s Guide

An all-in-one resource for instructors who want to integrate active
learning into their course. Searchable by chapter, phrase, topic, or
learning objective, the Interactive Instructor’s Guide compiles the many
valuable teaching resources available with Essential Cell Biology. This
website includes activities, discussion questions, animations and videos,
lecture outlines, learning objectives, primary literature suggestions,
medical topics guide, and more.

Coursepacks
Easily add high-quality Norton digital media to your online, hybrid, or
lecture course. Norton Coursepacks work within your existing learning
management system. Content is customizable and includes chapter-
based, multiple-choice reading quizzes, text-based learning objectives,
access to the full suite of animations, flashcards, and a glossary.

Test Bank
Written by Linda Huang, University of Massachusetts Boston, and Cheryl
D. Vaughan, Harvard University Division of Continuing Education,
the revised and expanded Test Bank for Essential Cell Biology includes
65–80 questions per chapter. Questions are available in multiple-choice,
matching, fill-in-the-blank, and short-answer formats, with many using
art from the textbook. All questions are tagged to Bloom’s taxonomy
level, learning objective, book section, and difficulty level, allowing in-
structors to easily create meaningful exams. The Test Bank is available in
ExamView and as downloadable PDFs from wwnorton.com/instructors.
 Preface ix

Animations and Videos

Streaming links give access to more than 130 videos and animations,
bringing the concepts of cell biology to life. The movies are correlated
with each chapter and callouts are highlighted in color.

Figure-integrated Lecture Outlines

All of the figures are integrated in PowerPoint, along with the section
and concept headings from the text, to give instructors a head start
creating lectures for their course.

Image Files
Every figure and photograph in the book is available for download in
PowerPoint and JPG formats from wwnorton.com/instructors.

STUDENT RESOURCES
digital.wwnorton.com/ecb5

Animations and Videos

Streaming links give access to more than 130 videos and animations,
bringing the concepts of cell biology to life. Animations can also be
accessed via the ebook and in select Smartwork5 questions. The movies
are correlated with each chapter and callouts are highlighted in color.

Student Site
Resources for self-study are available on the student site, including
multiple-choice quizzes, cell explorer slides, challenge and concept
questions, flashcards, and a glossary.
 ABOUT THE AUTHORS

BRUCE ALBERTS received his PhD from Harvard University and is a

professor in the Department of Biochemistry and Biophysics at the
University of California, San Francisco. He was the editor in chief of
Science from 2008 to 2013 and served as president of the U.S. National
Academy of Sciences from 1993 to 2005.
KAREN HOPKIN received her PhD from the Albert Einstein College of
Medicine and is a science writer. Her work has appeared in various
scientific publications, including Science, Proceedings of the National
Academy of Sciences, and The Scientist, and she is a regular contributor to
Scientific American’s daily podcast, “60-Second Science.”
ALEXANDER JOHNSON received his PhD from Harvard University and
is a professor in the Department of Microbiology and Immunology at the
University of California, San Francisco.
DAVID MORGAN received his PhD from the University of California, San
Francisco, where he is a professor in the Department of Physiology and
vice dean for research in the School of Medicine.
MARTIN RAFF received his MD from McGill University and is emeritus
professor of biology at the Medical Research Council Laboratory for
Molecular Cell Biology at University College London.
KEITH ROBERTS received his PhD from the University of Cambridge and
was deputy director of the John Innes Centre. He is emeritus professor at
the University of East Anglia.
PETER WALTER received his PhD from The Rockefeller University in New
York and is a professor in the Department of Biochemistry and Biophysics
at the University of California, San Francisco, and an investigator of the
Howard Hughes Medical Institute.

x
 Preface xi

LIST OF CHAPTERS and

SPECIAL FEATURES

CHAPTE R 1 Cells: The Fundamental Units of Life 1

PANEL 1–1 Microscopy 12
TABLE 1–1 Historical Landmarks in Determining Cell Structure 24
PANEL 1–2 Cell Architecture 25
How We Know: Life’s Common Mechanisms 30
TABLE 1–2 Some Model Organisms and Their Genomes 35

CHAPTE R 2 Chemical Components of Cells 39

TABLE 2–1 Length and Strength of Some Chemical Bonds 48
TABLE 2–2 The Chemical Composition of a Bacterial Cell 52
How We Know: The Discovery of Macromolecules 60
PANEL 2–1 Chemical Bonds and Groups 66
PANEL 2–2 The Chemical Properties of Water 68
PANEL 2–3 The Principal Types of Weak Noncovalent Bonds 70
PANEL 2– 4 An Outline of Some of the Types of Sugars 72
PANEL 2–5 Fatty Acids and Other Lipids 74
PANEL 2– 6 The 20 Amino Acids Found in Proteins 76
PANEL 2–7 A Survey of the Nucleotides 78

CHAPTE R 3 Energy, Catalysis, and Biosynthesis 81

PANEL 3–1 Free Energy and Biological Reactions 94
TABLE 3–1 Relationship Between the Standard Free-Energy Change, G°, and the Equilibrium Constant 96
How We Know: “High-Energy” Phosphate Bonds Power Cell Processes 102
TABLE 3–2 Some Activated Carriers Widely Used in Metabolism 109

CHAPTE R 4 Protein Structure and Function 117

PANEL 4 –1 A Few Examples of Some General Protein Functions 118
PANEL 4 –2 Making and Using Antibodies 140
TABLE 4 –1 Some Common Functional Classes of Enzymes 142
How We Know: Measuring Enzyme Performance 144
TABLE 4 –2 Historical Landmarks in Our Understanding of Proteins 160
PANEL 4 –3 Cell Breakage and Initial Fractionation of Cell Extracts 164
PANEL 4 – 4 Protein Separation by Chromatography 166
PANEL 4 –5 Protein Separation by Electrophoresis 167
PANEL 4 – 6 Protein Structure Determination 168

CHAPTE R 5 DNA and Chromosomes 173

How We Know: Genes Are Made of DNA 193

xi
xii List of Chapters and Special Features

CHAPTE R 6 DNA Replication and Repair 199

How We Know: The Nature of Replication 202
TABLE 6 –1 Proteins Involved in DNA Replication 213
TABLE 6 –2 Error Rates 218

CHAPTE R 7 From DNA to Protein: How Cells Read the Genome 227
TABLE 7–1 Types of RNA Produced in Cells 232
TABLE 7–2 The Three RNA Polymerases in Eukaryotic Cells 235
How We Know: Cracking the Genetic Code 246
TABLE 7–3 Antibiotics That Inhibit Bacterial Protein or RNA Synthesis 256
TABLE 7– 4 Biochemical Reactions That Can Be Catalyzed by Ribozymes 261

CHAPTE R 8 Control of Gene Expression 267

How We Know: Gene Regulation—The Story of Eve 280

CHAPTE R 9 How Genes and Genomes Evolve 297

TABLE 9–1 Viruses That Cause Human Disease 318
TABLE 9–2 Some Vital Statistics for the Human Genome 322
How We Know: Counting Genes 324

CHAPTE R 10 Analyzing the Structure and Function of Genes 333

How We Know: Sequencing the Human Genome 348

CHAPTE R 11 Membrane Structure 365

TABLE 11–1 Some Examples of Plasma Membrane Proteins and Their Functions 375
How We Know: Measuring Membrane Flow 384

CHAPTE R 12 Transport Across Cell Membranes 389

TABLE 12–1 A Comparison of Ion Concentrations Inside and Outside a Typical Mammalian Cell 391
TABLE 12–2 Some Examples of Transmembrane Pumps 403
How We Know: Squid Reveal Secrets of Membrane Excitability 412
TABLE 12–3 Some Examples of Ion Channels 419

CHAPTE R 13 How Cells Obtain Energy from Food 427

TABLE 13–1 Some Types of Enzymes Involved in Glycolysis 431
PANEL 13–1 Details of the 10 Steps of Glycolysis 436
PANEL 13–2 The Complete Citric Acid Cycle 442
How We Know: Unraveling the Citric Acid Cycle 444

CHAPTE R 14 Energy Generation in Mitochondria and Chloroplasts 455

TABLE 14 –1 Product Yields from Glucose Oxidation 469
PANEL 14 –1 Redox Potentials 472
How We Know: How Chemiosmotic Coupling Drives ATP Synthesis 476

CHAPTE R 15 Intracellular Compartments and Protein Transport 495

TABLE 15 –1 The Main Functions of Membrane-enclosed Organelles of a Eukaryotic Cell 497
TABLE 15 –2 The Relative Volumes and Numbers of the Major Membrane-enclosed Organelles
in a Liver Cell (Hepatocyte) 498
 List of Chapters and Special Features xiii

TABLE 15 –3 Some Typical Signal Sequences 502

TABLE 15 – 4 Some Types of Coated Vesicles 513
How We Know: Tracking Protein and Vesicle Transport 520

CHAPTE R 16 Cell Signaling 533

TABLE 16 –1 Some Examples of Signal Molecules 536
TABLE 16 –2 Some Foreign Substances That Act on Cell-Surface Receptors 544
TABLE 16 –3 Some Cell Responses Mediated by Cyclic AMP 550
TABLE 16 – 4 Some Cell Responses Mediated by Phospholipase C Activation 552
How We Know: Untangling Cell Signaling Pathways 563

CHAPTE R 17 Cytoskeleton 573

TABLE 17–1 Drugs That Affect Microtubules 584
How We Know: Pursuing Microtubule-associated Motor Proteins 588
TABLE 17–2 Drugs That Affect Filaments 594

CHAPTE R 18 The Cell-Division Cycle 609

TABLE 18–1 Some Eukaryotic Cell-Cycle Durations 611
How We Know: Discovery of Cyclins and Cdks 615
TABLE 18–2 The Major Cyclins and Cdks of Vertebrates 617
PANEL 18–1 The Principal Stages of M Phase in an Animal Cell 628

CHAPTE R 19 Sexual Reproduction and Genetics 651

PANEL 19–1 Some Essentials of Classical Genetics 675
How We Know: Using SNPs to Get a Handle on Human Disease 684

CHAPTE R 20 Cell Communities: Tissues, Stem Cells, and Cancer 691

TABLE 20 –1 A Variety of Factors Can Contribute to Genetic Instability 721
TABLE 20 –2 Examples of Cancer-critical Genes 728
How We Know: Making Sense of the Genes That Are Critical for Cancer 730
 Preface xv

CONTENTS

Preface v
About the Authors x

CHAPTER 1

Cells: The Fundamental Units of Life 1

UNITY AND DIVERSITY OF CELLS 2
Cells Vary Enormously in Appearance and Function 2
Living Cells All Have a Similar Basic Chemistry 3
Living Cells Are Self-Replicating Collections of Catalysts 4
All Living Cells Have Apparently Evolved from the Same Ancestral Cell 5
Genes Provide Instructions for the Form, Function, and Behavior of Cells and Organisms 6
CELLS UNDER THE MICROSCOPE 6
The Invention of the Light Microscope Led to the Discovery of Cells 7
Light Microscopes Reveal Some of a Cell’s Components 8
The Fine Structure of a Cell Is Revealed by Electron Microscopy 9
THE PROKARYOTIC CELL 11
Prokaryotes Are the Most Diverse and Numerous Cells on Earth 14
The World of Prokaryotes Is Divided into Two Domains: Bacteria and Archaea 15
THE EUKARYOTIC CELL 16
The Nucleus Is the Information Store of the Cell 16
Mitochondria Generate Usable Energy from Food Molecules 17
Chloroplasts Capture Energy from Sunlight 18
Internal Membranes Create Intracellular Compartments with Different Functions 19
The Cytosol Is a Concentrated Aqueous Gel of Large and Small Molecules 21
The Cytoskeleton Is Responsible for Directed Cell Movements 22
The Cytosol Is Far from Static 23
Eukaryotic Cells May Have Originated as Predators 24
MODEL ORGANISMS 27
Molecular Biologists Have Focused on E. coli 27
Brewer’s Yeast Is a Simple Eukaryote 28
Arabidopsis Has Been Chosen as a Model Plant 28
Model Animals Include Flies, Worms, Fish, and Mice 29
Biologists Also Directly Study Humans and Their Cells 32
Comparing Genome Sequences Reveals Life’s Common Heritage 33
Genomes Contain More Than Just Genes 35
ESSENTIAL CONCEPTS 36
QUESTIONS 37
xv
xvi Contents

CHAPTER 2

Chemical Components of Cells 39

CHEMICAL BONDS 40
Cells Are Made of Relatively Few Types of Atoms 40
The Outermost Electrons Determine How Atoms Interact 41
Covalent Bonds Form by the Sharing of Electrons 43
Some Covalent Bonds Involve More Than One Electron Pair 44
Electrons in Covalent Bonds Are Often Shared Unequally 45
Covalent Bonds Are Strong Enough to Survive the Conditions Inside Cells 45
Ionic Bonds Form by the Gain and Loss of Electrons 46
Hydrogen Bonds Are Important Noncovalent Bonds for Many Biological Molecules 47
Four Types of Weak Interactions Help Bring Molecules Together in Cells 47
Some Polar Molecules Form Acids and Bases in Water 49
SMALL MOLECULES IN CELLS 50
A Cell Is Formed from Carbon Compounds 50
Cells Contain Four Major Families of Small Organic Molecules 51
Sugars Are both Energy Sources and Subunits of Polysaccharides 52
Fatty Acid Chains Are Components of Cell Membranes 54
Amino Acids Are the Subunits of Proteins 56
Nucleotides Are the Subunits of DNA and RNA 56
MACROMOLECULES IN CELLS 58
Each Macromolecule Contains a Specific Sequence of Subunits 59
Noncovalent Bonds Specify the Precise Shape of a Macromolecule 62
Noncovalent Bonds Allow a Macromolecule to Bind Other Selected Molecules 62
ESSENTIAL CONCEPTS 64
QUESTIONS 65

CHAPTER 3

Energy, Catalysis, and Biosynthesis 81

THE USE OF ENERGY BY CELLS 82
Biological Order Is Made Possible by the Release of Heat Energy from Cells 83
Cells Can Convert Energy from One Form to Another 84
Photosynthetic Organisms Use Sunlight to Synthesize Organic Molecules 85
Cells Obtain Energy by the Oxidation of Organic Molecules 86
Oxidation and Reduction Involve Electron Transfers 87
FREE ENERGY AND CATALYSIS 88
Chemical Reactions Proceed in the Direction That Causes a Loss of Free Energy 89
Enzymes Reduce the Energy Needed to Initiate Spontaneous Reactions 89
The Free-Energy Change for a Reaction Determines Whether It Can Occur 90
G Changes as a Reaction Proceeds Toward Equilibrium 92
The Standard Free-Energy Change, G°, Makes It Possible to Compare the Energetics of
Different Reactions 92
The Equilibrium Constant Is Directly Proportional to G° 96
In Complex Reactions, the Equilibrium Constant Includes the Concentrations of
All Reactants and Products 96
 Contents xvii

The Equilibrium Constant Also Indicates the Strength of Noncovalent Binding Interactions 97
For Sequential Reactions, the Changes in Free Energy Are Additive 98
Enzyme-catalyzed Reactions Depend on Rapid Molecular Collisions 99
Noncovalent Interactions Allow Enzymes to Bind Specific Molecules 100
ACTIVATED CARRIERS AND BIOSYNTHESIS 101
The Formation of an Activated Carrier Is Coupled to an Energetically Favorable Reaction 101
ATP Is the Most Widely Used Activated Carrier 104
Energy Stored in ATP Is Often Harnessed to Join Two Molecules Together 106
NADH and NADPH Are Both Activated Carriers of Electrons 106
NADPH and NADH Have Different Roles in Cells 108
Cells Make Use of Many Other Activated Carriers 108
The Synthesis of Biological Polymers Requires an Energy Input 110
ESSENTIAL CONCEPTS 113
QUESTIONS 114

CHAPTER 4

Protein Structure and Function 117

THE SHAPE AND STRUCTURE OF PROTEINS 119
The Shape of a Protein Is Specified by Its Amino Acid Sequence 119
Proteins Fold into a Conformation of Lowest Energy 122
Proteins Come in a Wide Variety of Complicated Shapes 124
The a Helix and the b Sheet Are Common Folding Patterns 126
Helices Form Readily in Biological Structures 127
b Sheets Form Rigid Structures at the Core of Many Proteins 129
Misfolded Proteins Can Form Amyloid Structures That Cause Disease 129
Proteins Have Several Levels of Organization 129
Proteins Also Contain Unstructured Regions 130
Few of the Many Possible Polypeptide Chains Will Be Useful 131
Proteins Can Be Classified into Families 132
Large Protein Molecules Often Contain More than One Polypeptide Chain 132
Proteins Can Assemble into Filaments, Sheets, or Spheres 134
Some Types of Proteins Have Elongated Fibrous Shapes 134
Extracellular Proteins Are Often Stabilized by Covalent Cross-Linkages 135
HOW PROTEINS WORK 137
All Proteins Bind to Other Molecules 137
Humans Produce Billions of Different Antibodies, Each with a Different Binding Site 138
Enzymes Are Powerful and Highly Specific Catalysts 139
Enzymes Greatly Accelerate the Speed of Chemical Reactions   142
Lysozyme Illustrates How an Enzyme Works 143
Many Drugs Inhibit Enzymes 147
Tightly Bound Small Molecules Add Extra Functions to Proteins 148
HOW PROTEINS ARE CONTROLLED 149
The Catalytic Activities of Enzymes Are Often Regulated by Other Molecules 150
Allosteric Enzymes Have Two or More Binding Sites That Influence One Another 151
Phosphorylation Can Control Protein Activity by Causing a Conformational Change 152
Covalent Modifications Also Control the Location and Interaction of Proteins 153
Regulatory GTP-Binding Proteins Are Switched On and Off by the Gain and Loss of a Phosphate Group 154
xviii Contents

ATP Hydrolysis Allows Motor Proteins to Produce Directed Movements in Cells 154
Proteins Often Form Large Complexes That Function as Machines 155
Many Interacting Proteins Are Brought Together by Scaffolds 156
Weak Interactions Between Macromolecules Can Produce Large Biochemical
Subcompartments in Cells 157
HOW PROTEINS ARE STUDIED 158
Proteins Can Be Purified from Cells or Tissues 158
Determining a Protein’s Structure Begins with Determining Its Amino Acid Sequence 159
Genetic Engineering Techniques Permit the Large-Scale Production, Design, and Analysis of
Almost Any Protein 161
The Relatedness of Proteins Aids the Prediction of Protein Structure and Function 162
ESSENTIAL CONCEPTS 162
QUESTIONS 170

CHAPTER 5

DNA and Chromosomes 173

THE STRUCTURE OF DNA 174
A DNA Molecule Consists of Two Complementary Chains of Nucleotides 175
The Structure of DNA Provides a Mechanism for Heredity 176
THE STRUCTURE OF EUKARYOTIC CHROMOSOMES 178
Eukaryotic DNA Is Packaged into Multiple Chromosomes 179
Chromosomes Organize and Carry Genetic Information 180
Specialized DNA Sequences Are Required for DNA Replication
and Chromosome Segregation 181
Interphase Chromosomes Are Not Randomly Distributed Within the Nucleus 182
The DNA in Chromosomes Is Always Highly Condensed 183
Nucleosomes Are the Basic Units of Eukaryotic Chromosome Structure 184
Chromosome Packing Occurs on Multiple Levels 186
THE REGULATION OF CHROMOSOME STRUCTURE 188
Changes in Nucleosome Structure Allow Access to DNA 188
Interphase Chromosomes Contain both Highly Condensed
and More Extended Forms of Chromatin 189
ESSENTIAL CONCEPTS 192
QUESTIONS 196
 Contents xix

CHAPTER 6

DNA Replication and Repair 199

DNA REPLICATION 200
Base-Pairing Enables DNA Replication 200
DNA Synthesis Begins at Replication Origins 201
Two Replication Forks Form at Each Replication Origin 201
DNA Polymerase Synthesizes DNA Using a Parental Strand as a Template 205
The Replication Fork Is Asymmetrical 206
DNA Polymerase Is Self-correcting 207
Short Lengths of RNA Act as Primers for DNA Synthesis 208
Proteins at a Replication Fork Cooperate to Form a Replication Machine 210
Telomerase Replicates the Ends of Eukaryotic Chromosomes 213
Telomere Length Varies by Cell Type and with Age 214
DNA REPAIR 215
DNA Damage Occurs Continually in Cells 215
Cells Possess a Variety of Mechanisms for Repairing DNA 217
A DNA Mismatch Repair System Removes Replication Errors That Escape Proofreading 218
Double-Strand DNA Breaks Require a Different Strategy for Repair 219
Homologous Recombination Can Flawlessly Repair DNA Double-Strand Breaks 220
Failure to Repair DNA Damage Can Have Severe Consequences for a Cell or Organism 222
A Record of the Fidelity of DNA Replication and Repair Is Preserved in Genome Sequences 223
ESSENTIAL CONCEPTS 224
QUESTIONS 225

CHAPTER 7

From DNA to Protein: How Cells Read

the Genome 227
FROM DNA TO RNA 228
Portions of DNA Sequence Are Transcribed into RNA 229
Transcription Produces RNA That Is Complementary to One Strand of DNA 230
Cells Produce Various Types of RNA 232
Signals in the DNA Tell RNA Polymerase Where to Start and Stop Transcription 233
Initiation of Eukaryotic Gene Transcription Is a Complex Process 235
Eukaryotic RNA Polymerase Requires General Transcription Factors 235
Eukaryotic mRNAs Are Processed in the Nucleus 237
In Eukaryotes, Protein-Coding Genes Are Interrupted
by Noncoding Sequences Called Introns 239
Introns Are Removed from Pre-mRNAs by RNA Splicing 239
RNA Synthesis and Processing Takes Place in “Factories” Within the Nucleus 242
Mature Eukaryotic mRNAs Are Exported from the Nucleus 242
mRNA Molecules Are Eventually Degraded in the Cytosol 242
FROM RNA TO PROTEIN 243
An mRNA Sequence Is Decoded in Sets of Three Nucleotides 244
tRNA Molecules Match Amino Acids to Codons in mRNA 245
xx Contents

Specific Enzymes Couple tRNAs to the Correct Amino Acid 249

The mRNA Message Is Decoded on Ribosomes 249
The Ribosome Is a Ribozyme 252
Specific Codons in an mRNA Signal the Ribosome Where to Start and to Stop Protein
Synthesis 253
Proteins Are Produced on Polyribosomes 255
Inhibitors of Prokaryotic Protein Synthesis Are Used as Antibiotics 255
Controlled Protein Breakdown Helps Regulate the Amount of Each Protein in a Cell 256
There Are Many Steps Between DNA and Protein 257
RNA AND THE ORIGINS OF LIFE 259
Life Requires Autocatalysis 259
RNA Can Store Information and Catalyze Chemical Reactions 260
RNA Is Thought to Predate DNA in Evolution 261
ESSENTIAL CONCEPTS 262
QUESTIONS 264

CHAPTER 8

Control of Gene Expression 267

AN OVERVIEW OF GENE EXPRESSION 268
The Different Cell Types of a Multicellular Organism Contain the Same DNA 268
Different Cell Types Produce Different Sets of Proteins 269
A Cell Can Change the Expression of Its Genes in Response to External Signals 270
Gene Expression Can Be Regulated at Various Steps from DNA to RNA to Protein 270
HOW TRANSCRIPTION IS REGULATED 271
Transcription Regulators Bind to Regulatory DNA Sequences 271
Transcription Switches Allow Cells to Respond to Changes in Their Environment 273
Repressors Turn Genes Off and Activators Turn Them On 274
The Lac Operon Is Controlled by an Activator and a Repressor 275
Eukaryotic Transcription Regulators Control Gene Expression from a Distance 276
Eukaryotic Transcription Regulators Help Initiate Transcription
by Recruiting Chromatin-Modifying Proteins 276
The Arrangement of Chromosomes into Looped Domains Keeps Enhancers in Check 278
GENERATING SPECIALIZED CELL TYPES 278
Eukaryotic Genes Are Controlled by Combinations of Transcription Regulators 279
The Expression of Different Genes Can Be Coordinated by a Single Protein 279
Combinatorial Control Can Also Generate Different Cell Types 282
The Formation of an Entire Organ Can Be Triggered by a Single Transcription Regulator 284
Transcription Regulators Can Be Used to Experimentally Direct the Formation of Specific Cell
Types in Culture 285
Differentiated Cells Maintain Their Identity 286
 Contents xxi

POST-TRANSCRIPTIONAL CONTROLS 287

mRNAs Contain Sequences That Control Their Translation 288
Regulatory RNAs Control the Expression of Thousands of Genes 288
MicroRNAs Direct the Destruction of Target mRNAs 289
Small Interfering RNAs Protect Cells From Infections 290
Thousands of Long Noncoding RNAs May Also Regulate Mammalian Gene Activity 291
ESSENTIAL CONCEPTS 292
QUESTIONS 293

CHAPTER 9

How Genes and Genomes Evolve 297

GENERATING GENETIC VARIATION 298
In Sexually Reproducing Organisms, Only Changes to the Germ Line
Are Passed On to Progeny 299
Point Mutations Are Caused by Failures of the Normal Mechanisms
for Copying and Repairing DNA 300
Mutations Can Also Change the Regulation of a Gene 302
DNA Duplications Give Rise to Families of Related Genes 302
Duplication and Divergence Produced the Globin Gene Family 304
Whole-Genome Duplications Have Shaped the Evolutionary History of Many Species 306
Novel Genes Can Be Created by Exon Shuffling 306
The Evolution of Genomes Has Been Profoundly Influenced by Mobile Genetic Elements 307
Genes Can Be Exchanged Between Organisms by Horizontal Gene Transfer 308
RECONSTRUCTING LIFE’S FAMILY TREE 309
Genetic Changes That Provide a Selective Advantage Are Likely to Be Preserved 309
Closely Related Organisms Have Genomes That Are Similar
in Organization as Well as Sequence 310
Functionally Important Genome Regions Show Up as Islands of Conserved DNA Sequence 310
Genome Comparisons Show That Vertebrate Genomes Gain and Lose DNA Rapidly 313
Sequence Conservation Allows Us to Trace Even the Most Distant Evolutionary Relationships 313
MOBILE GENETIC ELEMENTS AND VIRUSES 315
Mobile Genetic Elements Encode the Components They Need for Movement 315
The Human Genome Contains Two Major Families of Transposable Sequences 316
Viruses Can Move Between Cells and Organisms 317
Retroviruses Reverse the Normal Flow of Genetic Information 318
EXAMINING THE HUMAN GENOME 320
The Nucleotide Sequences of Human Genomes Show How Our Genes Are Arranged 321
Differences in Gene Regulation May Help Explain How Animals with Similar Genomes Can Be So Different 323
The Genome of Extinct Neanderthals Reveals Much about What Makes Us Human 326
Genome Variation Contributes to Our Individuality—But How? 327
ESSENTIAL CONCEPTS 328
QUESTIONS 329
xxii Contents

CHAPTER 10

Analyzing the Structure and Function of

Genes 333
ISOLATING AND CLONING DNA MOLECULES 334
Restriction Enzymes Cut DNA Molecules at Specific Sites 335
Gel Electrophoresis Separates DNA Fragments of Different Sizes 335
DNA Cloning Begins with the Production of Recombinant DNA 337
Recombinant DNA Can Be Copied Inside Bacterial Cells 337
An Entire Genome Can Be Represented in a DNA Library 339
Hybridization Provides a Sensitive Way to Detect Specific Nucleotide Sequences 340
DNA CLONING BY PCR 341
PCR Uses DNA Polymerase and Specific DNA Primers to Amplify
DNA Sequences in a Test Tube 342
PCR Can Be Used for Diagnostic and Forensic Applications 343
SEQUENCING DNA 346
Dideoxy Sequencing Depends on the Analysis of DNA Chains
Terminated at Every Position 346
Next-Generation Sequencing Techniques Make Genome Sequencing Faster and
Cheaper 347
Comparative Genome Analyses Can Identify Genes and Predict Their Function 350
EXPLORING GENE FUNCTION 350
Analysis of mRNAs Provides a Snapshot of Gene Expression 351
In Situ Hybridization Can Reveal When and Where a Gene Is Expressed 352
Reporter Genes Allow Specific Proteins to Be Tracked in Living Cells 352
The Study of Mutants Can Help Reveal the Function of a Gene 354
RNA Interference (RNAi) Inhibits the Activity of Specific Genes 354
A Known Gene Can Be Deleted or Replaced with an Altered Version 355
Genes Can Be Edited with Great Precision Using the Bacterial CRISPR System 358
Mutant Organisms Provide Useful Models of Human Disease 359
Transgenic Plants Are Important for both Cell Biology and Agriculture 359
Even Rare Proteins Can Be Made in Large Amounts Using Cloned DNA 361
ESSENTIAL CONCEPTS 362
QUESTIONS 363
 Contents xxiii

CHAPTER 11

Membrane Structure 365

THE LIPID BILAYER 367
Membrane Lipids Form Bilayers in Water 367
The Lipid Bilayer Is a Flexible Two-dimensional Fluid 370
The Fluidity of a Lipid Bilayer Depends on Its Composition 371
Membrane Assembly Begins in the ER 373
Certain Phospholipids Are Confined to One Side of the Membrane 373
MEMBRANE PROTEINS 375
Membrane Proteins Associate with the Lipid Bilayer in Different Ways 376
A Polypeptide Chain Usually Crosses the Lipid Bilayer as an a Helix 377
Membrane Proteins Can Be Solubilized in Detergents 378
We Know the Complete Structure of Relatively Few Membrane Proteins 379
The Plasma Membrane Is Reinforced by the Underlying Cell Cortex 380
A Cell Can Restrict the Movement of Its Membrane Proteins 381
The Cell Surface Is Coated with Carbohydrate 382
ESSENTIAL CONCEPTS 386
QUESTIONS 387

CHAPTER 12

Transport Across Cell Membranes 389

PRINCIPLES OF TRANSMEMBRANE TRANSPORT 390
Lipid Bilayers Are Impermeable to Ions and Most Uncharged Polar Molecules 390
The Ion Concentrations Inside a Cell Are Very Different from Those Outside 391
Differences in the Concentration of Inorganic Ions Across a Cell Membrane
Create a Membrane Potential 391
Cells Contain Two Classes of Membrane Transport Proteins: Transporters
and Channels 392
Solutes Cross Membranes by Either Passive or Active Transport 392
Both the Concentration Gradient and Membrane Potential Influence the
Passive Transport of Charged Solutes 393
Water Moves Across Cell Membranes Down Its Concentration Gradient—a
Process Called Osmosis 394
TRANSPORTERS AND THEIR FUNCTIONS 395
Passive Transporters Move a Solute Along Its Electrochemical Gradient 396
Pumps Actively Transport a Solute Against Its Electrochemical Gradient 396
The Na+ Pump in Animal Cells Uses Energy Supplied by ATP to Expel Na+ and Bring in K+ 397
The Na+ Pump Generates a Steep Concentration Gradient of Na+ Across the Plasma Membrane 398
Ca2+ Pumps Keep the Cytosolic Ca2+ Concentration Low 399
Gradient-driven Pumps Exploit Solute Gradients to Mediate Active Transport 399
The Electrochemical Na+ Gradient Drives the Transport of Glucose Across the Plasma Membrane of Animal Cells 400
Electrochemical H Gradients Drive the Transport of Solutes in Plants, Fungi, and Bacteria
+
402
ION CHANNELS AND THE MEMBRANE POTENTIAL 403
Ion Channels Are Ion-selective and Gated 404
Membrane Potential Is Governed by the Permeability of a Membrane to Specific Ions 405
xxiv Contents

Ion Channels Randomly Snap Between Open and Closed States 407
Different Types of Stimuli Influence the Opening and Closing of Ion Channels 408
Voltage-gated Ion Channels Respond to the Membrane Potential 409
ION CHANNELS AND NERVE CELL SIGNALING 410
Action Potentials Allow Rapid Long-Distance Communication Along Axons 411
Action Potentials Are Mediated by Voltage-gated Cation Channels 411
Voltage-gated Ca2+ Channels in Nerve Terminals Convert an Electrical Signal into a Chemical
Signal 416
Transmitter-gated Ion Channels in the Postsynaptic Membrane Convert the Chemical Signal
Back into an Electrical Signal 417
Neurotransmitters Can Be Excitatory or Inhibitory 418
Most Psychoactive Drugs Affect Synaptic Signaling by Binding to Neurotransmitter
Receptors 419
The Complexity of Synaptic Signaling Enables Us to Think, Act, Learn, and Remember 420
Light-gated Ion Channels Can Be Used to Transiently Activate or Inactivate Neurons in Living
Animals 421
ESSENTIAL CONCEPTS 422
QUESTIONS 424

CHAPTER 13

How Cells Obtain Energy from Food 427

THE BREAKDOWN AND UTILIZATION OF SUGARS AND FATS 428
Food Molecules Are Broken Down in Three Stages 428
Glycolysis Extracts Energy from the Splitting of Sugar 430
Glycolysis Produces both ATP and NADH 431
Fermentations Can Produce ATP in the Absence of Oxygen 433
Glycolytic Enzymes Couple Oxidation to Energy Storage in Activated Carriers 434
Several Types of Organic Molecules Are Converted to Acetyl CoA
in the Mitochondrial Matrix 438
The Citric Acid Cycle Generates NADH by Oxidizing Acetyl Groups to CO2 438
Many Biosynthetic Pathways Begin with Glycolysis or the Citric Acid Cycle 441
Electron Transport Drives the Synthesis of the Majority of the ATP in Most Cells 446
REGULATION OF METABOLISM 447
Catabolic and Anabolic Reactions Are Organized and Regulated 447
Feedback Regulation Allows Cells to Switch from Glucose Breakdown to
Glucose Synthesis 447
Cells Store Food Molecules in Special Reservoirs to Prepare for Periods of Need 449
ESSENTIAL CONCEPTS 451
QUESTIONS 452
 Contents xxv

CHAPTER 14

Energy Generation in Mitochondria

and Chloroplasts 455
Cells Obtain Most of Their Energy by a Membrane-based Mechanism 456
Chemiosmotic Coupling Is an Ancient Process, Preserved in Present-Day Cells 457
MITOCHONDRIA AND OXIDATIVE PHOSPHORYLATION 459
Mitochondria Are Dynamic in Structure, Location, and Number 459
A Mitochondrion Contains an Outer Membrane, an Inner Membrane,
and Two Internal Compartments 460
The Citric Acid Cycle Generates High-Energy Electrons Required for ATP Production 461
The Movement of Electrons Is Coupled to the Pumping of Protons 462
Electrons Pass Through Three Large Enzyme Complexes in the Inner
Mitochondrial Membrane 464
Proton Pumping Produces a Steep Electrochemical Proton Gradient
Across the Inner Mitochondrial Membrane 464
ATP Synthase Uses the Energy Stored in the Electrochemical Proton
Gradient to Produce ATP 465
The Electrochemical Proton Gradient Also Drives Transport Across
the Inner Mitochondrial Membrane 466
The Rapid Conversion of ADP to ATP in Mitochondria Maintains
a High ATP/ADP Ratio in Cells 467
Cell Respiration Is Amazingly Efficient 468
MOLECULAR MECHANISMS OF ELECTRON TRANSPORT AND PROTON PUMPING 469
Protons Are Readily Moved by the Transfer of Electrons 469
The Redox Potential Is a Measure of Electron Affinities 470
Electron Transfers Release Large Amounts of Energy 471
Metals Tightly Bound to Proteins Form Versatile Electron Carriers 471
Cytochrome c Oxidase Catalyzes the Reduction of Molecular Oxygen 474
CHLOROPLASTS AND PHOTOSYNTHESIS 478
Chloroplasts Resemble Mitochondria but Have an Extra Compartment—the Thylakoid 478
Photosynthesis Generates—and Then Consumes—ATP and NADPH 479
Chlorophyll Molecules Absorb the Energy of Sunlight 480
Excited Chlorophyll Molecules Funnel Energy into a Reaction Center 481
A Pair of Photosystems Cooperate to Generate both ATP and NADPH 482
Oxygen Is Generated by a Water-Splitting Complex Associated with Photosystem II 483
The Special Pair in Photosystem I Receives its Electrons from Photosystem II 484
Carbon Fixation Uses ATP and NADPH to Convert CO2 into Sugars 484
Sugars Generated by Carbon Fixation Can Be Stored as Starch or Consumed to Produce ATP 487
THE EVOLUTION OF ENERGY-GENERATING SYSTEMS 488
Oxidative Phosphorylation Evolved in Stages 488
Photosynthetic Bacteria Made Even Fewer Demands on Their Environment 489
The Lifestyle of Methanococcus Suggests That Chemiosmotic Coupling Is an Ancient Process 490
ESSENTIAL CONCEPTS 491
QUESTIONS 492
xxvi Contents

CHAPTER 15

Intracellular Compartments and Protein

Transport 495
MEMBRANE-ENCLOSED ORGANELLES 496
Eukaryotic Cells Contain a Basic Set of Membrane-enclosed Organelles 496
Membrane-enclosed Organelles Evolved in Different Ways 499
PROTEIN SORTING 500
Proteins Are Transported into Organelles by Three Mechanisms 500
Signal Sequences Direct Proteins to the Correct Compartment 502
Proteins Enter the Nucleus Through Nuclear Pores 503
Proteins Unfold to Enter Mitochondria and Chloroplasts 505
Proteins Enter Peroxisomes from both the Cytosol and the Endoplasmic Reticulum 506
Proteins Enter the Endoplasmic Reticulum While Being Synthesized 507
Soluble Proteins Made on the ER Are Released into the ER Lumen 508
Start and Stop Signals Determine the Arrangement of a Transmembrane Protein
in the Lipid Bilayer 509
VESICULAR TRANSPORT 511
Transport Vesicles Carry Soluble Proteins and Membrane Between Compartments 511
Vesicle Budding Is Driven by the Assembly of a Protein Coat 512
Vesicle Docking Depends on Tethers and SNAREs 514
SECRETORY PATHWAYS 515
Most Proteins Are Covalently Modified in the ER 516
Exit from the ER Is Controlled to Ensure Protein Quality 517
The Size of the ER Is Controlled by the Demand for Protein Folding 518
Proteins Are Further Modified and Sorted in the Golgi Apparatus 518
Secretory Proteins Are Released from the Cell by Exocytosis 519
ENDOCYTIC PATHWAYS 523
Specialized Phagocytic Cells Ingest Large Particles 523
Fluid and Macromolecules Are Taken Up by Pinocytosis 524
Receptor-mediated Endocytosis Provides a Specific Route into Animal Cells 525
Endocytosed Macromolecules Are Sorted in Endosomes 526
Lysosomes Are the Principal Sites of Intracellular Digestion 527
ESSENTIAL CONCEPTS 528
QUESTIONS 530
 Contents xxvii

CHAPTER 16

Cell Signaling 533

GENERAL PRINCIPLES OF CELL SIGNALING 534
Signals Can Act over a Long or Short Range 534
A Limited Set of Extracellular Signals Can Produce a Huge Variety of Cell Behaviors 537
A Cell’s Response to a Signal Can Be Fast or Slow 538
Cell-Surface Receptors Relay Extracellular Signals via Intracellular Signaling Pathways 539
Some Intracellular Signaling Proteins Act as Molecular Switches 541
Cell-Surface Receptors Fall into Three Main Classes 543
Ion-Channel-Coupled Receptors Convert Chemical Signals into Electrical Ones 544
G-PROTEIN-COUPLED RECEPTORS 545
Stimulation of GPCRs Activates G-Protein Subunits 545
Some Bacterial Toxins Cause Disease by Altering the Activity of G Proteins 547
Some G Proteins Directly Regulate Ion Channels 548
Many G Proteins Activate Membrane-bound Enzymes That Produce Small
Messenger Molecules 549
The Cyclic AMP Signaling Pathway Can Activate Enzymes and Turn On Genes 549
The Inositol Phospholipid Pathway Triggers a Rise in Intracellular Ca2+ 552
A Ca2+ Signal Triggers Many Biological Processes 553
A GPCR Signaling Pathway Generates a Dissolved Gas That Carries a Signal to Adjacent Cells 554
GPCR-Triggered Intracellular Signaling Cascades Can Achieve Astonishing Speed,
Sensitivity, and Adaptability 555
ENZYME-COUPLED RECEPTORS 557
Activated RTKs Recruit a Complex of Intracellular Signaling Proteins 558
Most RTKs Activate the Monomeric GTPase Ras 559
RTKs Activate PI 3-Kinase to Produce Lipid Docking Sites in the Plasma Membrane 560
Some Receptors Activate a Fast Track to the Nucleus 565
Some Extracellular Signal Molecules Cross the Plasma Membrane and Bind to Intracellular Receptors 565
Plants Make Use of Receptors and Signaling Strategies That Differ from Those Used by Animals 567
Protein Kinase Networks Integrate Information to Control Complex Cell Behaviors 567
ESSENTIAL CONCEPTS 569
QUESTIONS 571
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The Project Gutenberg eBook of Crónicas
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Title: Crónicas imorais

Author: Albino Forjaz de Sampaio

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Language: Portuguese

Original publication: Lisboa: Santos & Vieira, 1918

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*** START OF THE PROJECT GUTENBERG EBOOK CRÓNICAS

IMORAIS ***
 ALBINO FORJAZ DE SAMPAYO

Crónicas
imorais
«O que melhor se ria será
o último a rir-se.»

F. Nietzsche.

3.º MILHAR

Editores—S A N T O S & V I E I R A
EMPREZA LITTERARIA FLUMINENSE
125, Rua dos Retrozeiros, 125
LISBOA
CRÓNICAS IMORAIS

Editores—SANTOS & VIEIRA—LISBOA

Typ. da Empreza Litteraria e Typogr.
(Officinas movidas a electricidade)
R. Elias Garcia, 184 Porto MCMXV
 OBRAS DO MESMO AUTOR:
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ALBINO FORJAZ DE SAMPAYO

Crónicas
imorais
«O que melhor se ria será
o último a rir-se».

F. Nietzsche.

3.º MILHAR

EDITORES—Santos & Vieira

EMPREZA LITTERARIA FLUMINENSE
125, Rua dos Retrozeiros, 125
LISBOA
 AO
DR. BRITO CAMACHO
 Crónicas imorais
OS artigos, crónicas ou antes as impressões que hoje se reúnem
em volume são já do domínio público. Todavia devo declarar que
não foi nele que pensei, quando as escrevi. Os quatro géneros de
criaturas que há no mundo, «criaturas racionais, como os homens;
criaturas sensitivas como os animais; criaturas vegetativas, como as
plantas; criaturas insensíveis, como as pedras», como quere Vieira,
só mediocremente me interessam. Publicam-se hoje porque isso me
apraz, e se são más ou boas, tal não me fará doer a cabeça. Nunca
mais as lerei. São cousas que passaram, notas à margem dum
acontecimento, comentários a uma leitura, impressões de momento,
fumo, cinza, terra, nada. Viveram um dia. Hoje reúne-as um certo
egoismo. Vivi com elas. Acharia a algumas, se as voltasse a ler,
uma ingenuidade primitiva; a outras uma piada estupenda.
Pensadas e escritas à hora em que o chefe da tipografia vem pedir
original, só para mim terão valor. No emtanto marcam cousas
curiosas e são um depoimento pessoal não falho de interêsse.
Umas são indignadas, outras biliosas, algumas pândegas. Não
admira. Filhas do momento que as inspirou, devem tambêm ter o
seu grão de incoerência, de petulância, de atrevimento. Que
importa? Não as renego. Chamei-lhes imorais, porque algumas
tiveram o condão de irritar muita gente boa, que ainda tem moral.
Ah! abjecta gente a que ainda tem moral! Muitas mereceram acres
censuras e outras ainda despertaram louvores. As que despertaram
louvores tive o cuidado de não as juntar aqui. Devem ser más por
fôrça.
Vivendo a vida do jornal, perdendo-se no arquivo do
esquecimento que o jornal é, em pouco tempo estariam esquecidas.
Em livro, não será mais longa a sua vida, mas sempre o entêrro é
melhor engendrado. Imorais disse eu. Imorais sim porque quási
sempre estão em desacôrdo com a moral do meu parceiro. Que
devo confessar-lhes, não me dou nada bem com os que estão de
acôrdo. Quem está de acôrdo não sabe ter razão. Porque a razão
não é dos que a teem, é dos que teem talento para a ter. Devo
tambêm confessar-lhes, que estas crónicas são inofensivas. A
ninguêm fazem mal. Não passam dum bom riso, um grande riso.
Tolice seria tomar tudo isto a sério. O mundo é uma espécie de
revista e quem se mata morre cedo. Se alguma vez, leitor, te
interessares, olha que tudo isto é ilusão. Não vale a pena, em
verdade te digo. Ilusão, ilusão sómente. A comoção, a ironia, o riso,
a tristeza, tudo isto que vês reunido, tudo isto, não é mais do que
cousas que já passaram, recordações, notas breves, leves
apontamentos, terra, pó, nada, cinza.
E não se comovam que se acaba o prólogo.
 Juízo do ano
O ano que começou ontem será um ano igual aos outros. Igual, sem
tirar nem pôr. Haverá nele, como já houve o ano passado, como
houve em todos os anos que passaram e como haverá em todos os
anos que vierem, tolos que enriquecem e tolos que cavam pés de
burro, asnos que se suicidam e asnos que acham isto uma cousa
óptima. Continuará a haver uma ignorância formidável da multidão
dominada e uma patifaria criminosa da minoria dominante;
continuará a haver homens cavalos e homens cavaleiros; sábios
que são burros e burros que são sábios; muita maldade nas
mulheres—a maldade nas mulheres é um pleonasmo!—e muita
estultícia nos homens. Haverá côres para todos os gostos, acepipes
para todos os paladares e partidos para todos os cidadãos. Não
sucederá nada de novo, porque nada é novo debaixo do sol, diz a
voz grave do vélho Eclesiastes, um mágico que sabia muito mais da
patifaria humana do que o snr. Civinini das mulheres que usam
bigode e pêra.
O novo ano será um ano feliz para quem jogar e lhe sair «el
prémio gordo». Para os que joguem e lhes sair branco será um ano
de azar.
Quem trabucar, manduca, a não ser que perca o apetite. Quanto
ao resto, o que êste ano será, é fácil de adivinhar.
Para os tolos, uma felicidade, porque êsses banabóias são felizes
todos os dias; para os que não sejam, nem eu mesmo sei.
Quanto ao ano artístico, será deplorável, porque o que terá menos
é arte, e de artistas nem sombra. Haverá vários quadros assinados
por estes e aqueles fulanos. Se é paisagem, já sabemos o que é:
«Macieira em flor», «Um trecho da Tapada» ou a «Ribeira de
Algés». Se é retrato, um cavalheiro ou uma cavalheira de cara
lambuza e desconfiada, olhos de goraz do vapor, ou então um ar
bonacho de quem diz à gente: «Então que tal? como passou? estou
catita, heim?»
Em arquitectura, continuarão a aparecer projectos... do Palácio da
Justiça, de várias saladas de cal e areia para projecto ou para
habitar, e de «artes várias».
Os nossos escultores continuarão fazendo Senhoras da
Conceição, cruzes para jazigos, e «muchas cosas más» dum
«salero» infinito.
Das finanças: «finanças se chamam as rendas públicas quando
Portugal está a finar-se». É de Camilo o dito. Das finanças dizia eu...
mas em finanças não sou muito forte. Adiante.
O ano literário... Continuará a haver literatura, literatos vulgares
de Linneu, literatos abezelgados e chués, porque, irmãos
caríssimos, «todo o homem tem em si uma porção de inépcia, que
há de sair em prosa ou verso, em palavras ou obras, como o
carnicão dum furúnculo». Homens de génio não haverá, mas em
compensação abundosos se prognostica os homens de génio mau
ou de bom génio, porque cada um é como o pai o fêz. A crítica
continuará a ser como foi sempre: De mostarda, de manteiga, e de
àgua e sal.
E, tenho dito.
Quanto a ti, leitor molesto, eu não tiro o teu horóscopo nem te leio
as ruins tenções de que porventura estejas cheio. Mas é sempre
bom conversar, uma conversa de amigos vélhos e inseparáveis.
Não sei se és rico, se pobre, se alto, se baixo. Se rico, guarda a
bôlsa que não preciso dela; se pobre, tem paciência porque não te
posso valer.
Dito isto, em verdade te digo que tens um ano diante de ti.
Emprega-o bem. Lê a vida do bom homem Ricardo, regula as tuas
digestões, não tenhas excessos e deita-te cedo. A isto se chama em
bom português fazer pela vida.
Faze pois pela vida. Lembra-te que «os mortos caem fácilmente
no olvido», como dizia Bürger na balada de Leonora, traduzida por
êsse tristonho Gerard de Nerval que se enforcou num candieiro da
Rue Vieille-Lanterne, numa manhã gelada de janeiro em que um
corvo, que parecia fugido ao «Never more!» de Poe, lhe crocitava
satânico e lúgubre a sua elegia de tímido, de sonhador e de
incompreendido. ¿E quem hoje, no aniversário da sua morte, se
lembra dêsse pobre Gerard, que trazia sempre os bolsos cheios de
livros, como o Schaunard da Bohème, que traduziu Goethe e visitou
o Oriente? Ah! é bem certo que «os mortos caem fácilmente no
olvido!»
Faze por ter dinheiro. O Dinheiro, alêm de ser tudo o que tu
sabes, é ainda aquilo com que se compram os melões. Se o tiveres
não o emprestes nem o dês. Se precisares não peças, porque
ninguêm te vale. Gritar é inútil tambêm, para que não chames
curiosos à tua desventura.
Prefere «um pássaro na mão a dois voando» e não te fies na
Virgem. Porque se te fias na Virgem e não corres não tarda o fatal e
bem merecido pontapé.
Se és casado não leves amigos a casa. Isto não é para que te
ofendas, é por uma cousa que eu cá sei. Se tens filhos, ao menos
um filho só, que é cousa que tôda a gente tem, seu ou alheio, faz do
teu filho um homem forte. Bom estômago, bons nervos, bons
músculos. Antes o obrigues à frequência do mestre Raku, um sujeito
que ganha a vida a deitar os outros ao chão, do que à de Félix
Pereira, que lhe ensina que meter os dedos no nariz é porcaria. É
preferível ser forte a ser bem-criado. É mesmo preferível ter fôrça a
ter direito, «porque se vai mais longe com a mão cheia de fôrça do
que com um saco cheio de direito», ensina a experiência dos
homens e a sabedoria das nações.
Podes ensinar-lhe muitas mais cousas que tu saibas. Não o
queiras nem artista, nem literato, nem jornalista. Vê se o podes fazer
par do reino, que o pariato é uma cousa que se está dando ou se vai
dar a tôda a gente, exactamente como o hábito de Cristo ou de S.
Tiago. «Dizem que até há barbeiros...» como já o suspeitava o
Baptista dos Maias. Se o fizeres par, porque êle não tenha geiteira
para outra cousa, recomenda-lhe que se cale. «O silêncio é de
ouro» e um tolo calado, conquanto não deixe de ser tolo, passa por
homem sisudo. Porque se êle assim não fôr, tanto pior para êle. Os
que assim não são, nada teem a esperar do ano novo. São e serão
sempre escarnecidos, ridículos e pobres. E o pior mal dum homem é
ser pobre. Ninguêm lhe vale. Ser pobre é...
Não continuo porque um cavalheiro que está vendo o que eu
escrevo, refila, em ar de resposta:
—«Mas há a Caridade. A Caridade, homem!»
—«A Caridade? Ah! sim.»—As misérias do próximo comovem
muito a caridade de cada um. E eu, que tambêm me vou tornando
azêdo como o senhor Silva Pinto, resmunguei e recordei-me. Devia
ser uma caridade como a daquele barão da Falperra que o nosso
Alfredo Mesquita conheceu: Um homem tão caridoso que, depois de
ter ouvido um pobre contar-lhe as suas misérias, a ponto de o fazer
chorar, chamava sempre o criado e, com a voz entrecortada de
soluços, ordenava:—«João, ponha êsse homem no ôlho da rua;
parte-se-me o coração de o ouvir...»
E para os que lhe dissessem que êle lhe não dera nada, retorquia,
que sim, que dera. Dera-lhe... atenção.
E estava certo, como diz ainda o senhor Silva Pinto.
 Artistas
A morte recente dêsse desventurado Augusto Santo, escultor do
Pôrto, veio avocar com amarga intensidade a malfadada sorte que
está guardada a todos que teem a suprema desventura de ter
nascido artistas em Portugal. Augusto Santo foi discípulo de
Falguière, de Rodin e de Soares dos Reis. Ouviu as lições de Taine,
e expôs no Salon. Era quási um desconhecido e morreu na maior
miséria, num catre humilde de hospital.
Ser desconhecido em Portugal é um caso banalíssimo.
Herculano, e êsse Garrett que «num dia de apuro por cem ou
duzentas moedas seria capaz de tôdas as porcarias menos de pôr
num papel a trôco de todo o ouro do mundo uma linha mal escrita»
são, ainda que isto pareça um paradoxo, quási desconhecidos. O
grande público não tem ideia dêles e ainda hoje uma das suas
edições leva uma eternidade para se esgotar.
A indiferença do público pelos artistas é absoluta, e não vai longe
ainda o tempo em que literato era sinónimo de vadio. Camilo no
Pôrto, ao tempo, era sómente um janota que para ali quebrava
esquinas, um tal que não avezava com que mandar cantar um cego.
Quando, pelos romances, ganhava com que forrar de cuidados o
passadio de dois meses, não se imagina o escândalo que aquilo
produzia nos Antónios Josés da Silva e na rua das Congostas. O
Pôrto de então tinha ideias seguras e via as cousas como devia ver.
Literatos neste país?! hum! e torcia o nariz como quem dizia que
aquilo não era prático, nem por aquele caminho se chegava a ter um
rolosito de inscrições, um prédiosito, ou uma velhice sossegada. Era
a vida prática, o balcão é que era o caminho. Por isso, quando
Camilo, já no apogeu de glória, em cartas duma dolorosa
humildade, quási esmolava, para comer, a compra de pratas que os
seus admiradores lhe ofereciam, o Pôrto mui devia rir. Êle bem lhe
dizia! Não era aquele o caminho...
 A Soares dos Réis para lhe fazerem justiça foi preciso que a
morte o tomasse. Em vida foi um obscuro obreiro sem amigos, ou
quási, e teve uma existência bastante precária. Basta dizer que em
tôda a sua vida de trabalho não conseguiu mais do que 4:764$500
réis segundo a autobiografia do artista, que o snr. Joaquim de
Vasconcelos, em 1905, confidenciou ao público por intermédio da
Revista. Para ganhar essa importância confessa Soares dos Réis
que esteve «algumas vezes em relação com a arte industrial».
Mesmo assim quem a achar exagerada deduza-lhe o custo do
material e de auxiliares indispensáveis e verá quanto fica. O Artista
na Infância vendeu-o por 600$000 réis e foi o máximo preço que
levou por uma obra sua, se exceptuarmos, é claro, o monumento ao
Conde de Ferreira.
Em vida ninguêm o auxiliou, ninguêm o encorajou para
prosseguir. Muito ao contrário perseguiam-no, sitiavam-no,
roubavam-no, fechavam-lhe tôdas as portas com intrigas soezes,
com indróminas de sabidos e ronha de marotos. Concurso a que
fôsse, vaga a que concorresse, plano que tentasse viabilizar, certo
era a mediocridade arranjista, por portas travessas, frustrar-lhe tôda
a ambição e todo o Sonho.
Emquanto o homem viveu não puderam os amigos, êsses amigos
do diabo, atormentá-lo mais, nem mais o perseguirem. Depois de
morto não houve lamúria que não chorassem e não houve adjectivo
sonoroso e amelaçado que a criatura não tivesse. A justiça chega
sempre depois da morte, é certo, mas pela injustiça dos vivos.
Com Augusto Santo o caso é o mesmo. Augusto Santo era, antes
de tudo, um impersistente. Não possuía a tenacidade avara, fria e
reservada, a confiança absoluta dêsses brocadores do Ideal,
fadados para o êxito. A menor contrariedade o exasperava. E como
se não fôsse bastante a execução perra, o sonho nebuloso, a vida
material e a falta de tudo, ainda a minar-lhe a existência a
agressividade constante dos outros,—os colegas os primeiros,—não
fôsse o pobre diabo roubar-lhes a glória, e numa avidez de faminto a
guardasse tôda para si. Augusto Santo foi um perseguido desde que
expôs êsse Ismael, a sua única obra a valer, mas ainda assim uma
promessa do que o seu temperamento de artista poderia dar.
 Como vêem, eu não quero dizer que Augusto Santo era um génio
e que a sua morte abre uma clareira formidável entre os Teixeiras e
os Lopes da pedra portuguesa. Não. Augusto Santo não deixou
fauteuil vago. O que eu quero precisar é que era Augusto Santo um
artista a valer, que noutro meio floresceria, um meio que lhe não
fôsse hostil como o seu,—tão hostil que até o deixou morrer de fome
numa enxêrga de hospital. Com esta hostilidade e com esta
ingratidão a arte perdeu, mas ganharam os escultores portugueses.
Podem agora catrapiscar a imortalidade à vontadinha, que ela não
esperará que êles morram para vir, ou o seu cão, sevandijar-lhes em
cima da obra.
Alguns gazeteiros ou gazetíferos, com as palavras de louvor do
cliché, justificam porque Augusto Santo não teve carruagem às
horas, mesa lauta e colchão fôfo. Foi, dizem êles reprimendando os
restos do escultor, porque era um inadaptado, porque desprezou
sempre a arte de engorda, a que dá lucros e considerações, e
preferiu correr atrás dum sonho que o exauria e que o matava, um
sonho de arte irrealisável e irrealisado, arte verdadeira que não tem
preço, embora às vezes se venda, e que não se compra, embora às
vezes se adquira. E com um desdêm olímpico, absoluto, ditatorial,
chamam-lhe... sonhador. Sonhador!... como se isto fôsse o sumo
desprêzo ou a máxima compaixão. Deve ser bem triste morrer,
assim!
Ao menos se essa turba se calasse e governasse vida, vá; mas
vir babujar a sua irresponsabilidade desvergonhada no momento em
que o pobre vencido solavancava a caminho do cemitério, é
intolerável. Não se justifique, que ninguêm lhe tira o ganho: ela tem
a sua utilidade. ¿Quem é que nos havia afinal de fazer os fretes?
Augusto Santo, não reste dúvida, foi um atormentado da forma.
Quando a concepção, megalomania conceptiva era nele, lhe
fabulava maravilhas, logo o barro, parece que conluiado com os
homens, debaixo dos gadanhos convulsos e nervosos tinha formas
brutais, aduncas e agressivas. E a sua frialdade viscosa logo ali
abafava o delírio artístico do pobre impotente e lhe dava crises de
desânimo capazes de vergar um atleta.
 Todo o Pôrto cabareteiro e intelectual conhecia estas torturas do
artista. Sem fôrças para opôr uma resistência, êle consumia-se em
desolações. Cada tentativa malograda deixava-o mais exausto de
fôrças do que uma noite perdida a caminhar. O Sonho vampirizava-o
e exigia-lhe uma impossível produção de calorias. Todavia nada
resta dele, decorrendo como decorreu intramuralhas do seu crânio.
Após a criação dêsse Ismael, que noutro meio mais acarinhante, se
o não celebrizasse, o evidenciaria, Augusto Santo criara, para si
mesmo, responsabilidades. Ficaram por cumprir. A quantos artistas
não sucede outro tanto?! Todos o sabem, Portugal artísticamente é
um país morto. Capaz duma ressurreição? Certamente. Desde que
os poderes públicos olhem com atenção para a Arte, desde que a
percentagem assustadora dos analfabetos diminua e desde que o
público manifeste interêsse por estas cousas. Arte em Portugal? Os
pintores é que sabem o que isso é. Êles bem vêem que não é de
sonhos e de belos quadros que se vive. O que dá são as lições, que
é como quem diz os quadros que fazem e os discípulos assinam. É
uma arte de opereta, de muito riso, com sinfonias de Offenbach e de
que só vai mal a quem a toma a sério. Êsses são os vencidos, os
Soares dos Réis e os Augustos Santos.
Ê pois mais um artista que sucumbiu em luta inglória com o
Destino. Consagrado ou desconhecido, não há dúvida que era um
artista. O Ismael o atesta. E já que ninguêm teve que o auxiliasse e
que no meio da indiferença geral da gente que escreve e do regosijo
da gente que escultura, êle deixou a vida, justo é que eu o admire,
ao seu exemplo de intransigência, à luta que sustentou e à tragédia
da vida que sofreu. E ninguêm se lembrou dele senão a Morte, que
se amerceou de tanto sofrimento. Ó morte remediadora e
sacratíssima, amiga dos pobres, remidora dos desventurados e
consoladora dos aflitos, eu te bemdigo.
 O Jettatore
TEÓFILO Gauthier, o Benvenuto Cellini da prosa francesa, segundo
Camilo, consagrou um dos seus volumes à Jettatura, ou mau
olhado, condão funesto do Jettatore. Nesse volume, obra prima do
lapidário precioso a quem já Baudelaire havia chamado «poeta
impecável, mago e mestre», se romantiza a vida dum jettatore e se
dá conta de todos os malefícios que o seu olhar causou. Paulo
d’Aspremont, francês, é o protagonista do romance, e, sem o saber,
possui o fascino, que, traduzido em vulgar, significa mau olhado. O
seu olhar fatal onde pousa é nefasto. A pata do cavalo de Attila não
teve tanta crueldade. Se se debruça na amurada dum barco, para
olhar a água mansa e tranqùila, logo o mar se encapela e enfuría;
se entra num teatro e fita um actor que ri, logo o espectáculo se
interrompe e a própria Troça fica séria; se olha uma bailarina que
rodopia, um bico de gás incendeia a gaze e a bailarina morre.
Quando fita a sua própria noiva, o pobre jettatore mata-a. Uma
roseta escarlate assoma às faces da condenada inglêsa, um escarro
de sangue vem da goela à bôca, e dali à morte é um ai. Nápoles
inteiro o odeia. «Tôdas as vezes que parava junto duma loja, o dono
parecia assustado, murmurava algumas imprecações a meia voz, e
alongava os dedos como se quisesse apunhalá-lo com o auricular e
o índex; as vendedeiras mais ousadas, acabrunhavam-o de
impropérios e mostravam-lhe o punho.» Por fim cega-se e suicida-
se, arremessando-se ao mar. O seu corpo nunca se encontrou.
Quando êle morreu,—oh meu adorável Gauthier!—quando êle
morreu «a tempestade desencadeou-se então com tôda a fúria: as
ondas assaltaram a praia em filas compactas, como guerreiros
correndo ao assalto, e lançando a cincoenta passos ao ar jactos de
espuma; as nuvens negras alagartaram-se como paredes do
inferno, deixando entrever a ardente fornalha dos relâmpagos; luzes
sulfurosas iluminaram a extensão; o cume do Vesúvio avermelhou-
se e um penacho de sombrio vapor, que o vento impelia, ondulou na
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