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The Cover
The cover image shows the formation of an autophagosome, in which organ-
elles and cytosol are engulfed in cytoplasmic membranes. Original painting
by David S. Goodsell, based on the scientific design of Daniel J. Klionsky.

The Artist
David S. Goodsell is an Associate Professor of Molecular Biology at the
Scripps Research Institute. His illustrated books, The Machinery of Life and
Our Molecular Nature, explore biological molecules and their diverse roles
within living cells, and his new book, Bionanotechnology: Lessons from Nature,
presents the growing connections between biology and nanotechnology.
More information may be found at: http://mgl.scripps.edu/people/goodsell

The Cell: A Molecular Approach, Seventh Edition


Copyright 2016 by Geoffrey M. Cooper. All rights reserved.
This book may not be reproduced in whole or in part without permission.

Address orders and requests for examination copies to: Sinauer Associates
P.O. Box 407, 23 Plumtree Road, Sunderland, MA 01375 U.S.A.
Phone: 413-549-4300
FAX: 413-549-1118
email: orders@sinauer.com
www.sinauer.com

Library of Congress Cataloging-in-Publication Data

Cooper, Geoffrey M., author.


The cell : a molecular approach / Geoffrey M. Cooper, Robert E. Hausman.
-- Seventh edition.
p. ; cm.
Includes bibliographical references.
ISBN 978-1-60535-290-9
I. Hausman, Robert E., 1947-2015, author. II. Title.
[DNLM: 1. Cells. 2. Cell Biology. 3. Cell Physiological Phenomena.
4. Molecular Biology. QU 300]
QH581.2
571.6--dc23
2015032684
Printed in U.S.A
6 5 4 3 2 1
To my friend, colleague, and coauthor,
Robert E. Hausman
(1947–2015)

Geoffrey M. Cooper
Brief Table of Contents
PART I
Fundamentals and Foundations 1
Chapter 1 An Overview of Cells and Cell Research 3
Chapter 2 Molecules and Membranes 47
Chapter 3 Bioenergetics and Metabolism 81
Chapter 4 Fundamentals of Molecular Biology 111
Chapter 5 Genomics, Proteomics, and Systems Biology 157

PART II
The Flow of Genetic Information 185
Chapter 6 Genes and Genomes 187
Chapter 7 Replication, Maintenance, and Rearrangements
of Genomic DNA 217
Chapter 8 RNA Synthesis and Processing 259
Chapter 9 Protein Synthesis, Processing, and Regulation 317

PART III
Cell Structure and Function 365
Chapter 10 The Nucleus 367
Chapter 11 Protein Sorting and Transport 397
Chapter 12 Mitochondria, Chloroplasts, and Peroxisomes 447
Chapter 13 The Cytoskeleton and Cell Movement 479
Chapter 14 The Plasma Membrane 531
Chapter 15 Cell Walls, the Extracellular Matrix, and Cell Interactions 571

PART IV
Cell Regulation 599
Chapter 16 Cell Signaling 601
Chapter 17 The Cell Cycle 651
Chapter 18 Cell Death and Cell Renewal 691
Chapter 19 Cancer 723
Contents
PART I
Fundamentals and Foundations 1
Chapter 1 An Overview of Cells and Cell Research 3
The Origin and Evolution of Cells 4 Tools of Cell Biology 23
The first cell 4 Light microscopy 23
The evolution of metabolism 7 Electron microscopy 30
Present-day prokaryotes 8 Super-resolution light microscopy 32
Eukaryotic cells 9 Subcellular fractionation 34
The origin of eukaryotes 11 Growth of animal cells in culture 37
The development of multicellular organisms 13 Key Experiment
HeLa Cells 39
Cells as Experimental Models 17 Culture of plant cells 39
E. coli 17
Viruses 40
Yeasts 18
Molecular Medicine
Caenorhabditis elegans 19 Viruses and Cancer 41
Drosophila melanogaster 20 SUMMARY AND KEY TERMS 43
Arabidopsis thaliana 20 QUESTIONS 44
Vertebrates 20 REFERENCES AND FURTHER READING 45

Chapter 2 Molecules and Membranes 47


The Molecules of Cells 47 The catalytic activity of enzymes 62
Carbohydrates 48 Mechanisms of enzymatic catalysis 63
Lipids 50 Coenzymes 66
Nucleic acids 54 Regulation of enzyme activity 68
Proteins 56 Cell Membranes 70
Key Experiment Membrane lipids 70
The Folding of Polypeptide Chains 58
Membrane proteins 71
Enzymes as Biological Catalysts 62 Key Experiment
The Structure of Cell Membranes 73
viii Contents

Transport across cell membranes 74 QUESTIONS 78


SUMMARY AND KEY TERMS 77 REFERENCES AND FURTHER READING 78

Chapter 3 Bioenergetics and Metabolism 81


Metabolic Energy and ATP 81 ATP synthesis 98
Free energy and ATP 81 Synthesis of glucose 99
Glycolysis and the Krebs cycle 84 The Biosynthesis of Cell Constituents 101
The derivation of energy from lipids 89 Carbohydrates 101
Electron Transport and Oxidative Lipids 102
Phosphorylation 89 Proteins 103
The electron transport chain 90 Key Experiment
Antimetabolites and Chemotherapy 105
Chemiosmotic coupling 92
Key Experiment Nucleic acids 106
The Chemiosmotic Theory 93 SUMMARY AND KEY TERMS 107
QUESTIONS 108
Photosynthesis 95
REFERENCES AND FURTHER READING 109
Electron transport 96

Chapter 4 Fundamentals of Molecular Biology 111


Heredity, Genes, and DNA 111 Detection of Nucleic Acids and Proteins 135
Genes and chromosomes 111 Amplification of DNA by the polymerase chain
Genes and enzymes 115 reaction 135
Identification of DNA as the genetic material 115 Nucleic acid hybridization 137
The structure of DNA 116 Antibodies as probes for proteins 140
Replication of DNA 118 Gene Function in Eukaryotes 142
Expression of Genetic Information 119 Gene transfer in plants and animals 142
Colinearity of genes and proteins 120 Mutagenesis of cloned DNAs 144
The role of messenger RNA 120 Introducing mutations into cellular genes 146
The genetic code 121 Genome engineering by the CRISPR/Cas system 148
RNA viruses and reverse transcription 124 Interfering with cellular gene expression 148
Key Experiment Key Experiment
The DNA Provirus Hypothesis 125 RNA Interference 151
Recombinant DNA 126 SUMMARY AND KEY TERMS 153
Restriction endonucleases 126 QUESTIONS 154
Generation of recombinant DNA molecules 128 REFERENCES AND FURTHER READING 155
Vectors for recombinant DNA 130
DNA sequencing 132
Expression of cloned genes 134
Contents ix

Chapter 5 Genomics, Proteomics, and Systems Biology 157


Genomes and Transcriptomes 157 Global analysis of protein localization 170
The genomes of bacteria and yeast 157 Protein interactions 171
The genomes of Caenorhabditis elegans, Drosophila Systems Biology 173
melanogaster, and Arabidopsis thaliana 159
Systematic screens of gene function 174
The human genome 160
Regulation of gene expression 175
The genomes of other vertebrates 160
Networks 176
Key Experiment
Synthetic biology 178
The Human Genome 161
Molecular Medicine
Next-generation sequencing and personal genomes 164
Malaria and Synthetic Biology 180
Global analysis of gene expression 166
SUMMARY AND KEY TERMS 181
Proteomics 167 QUESTIONS 182
Identification of cell proteins 167 REFERENCES AND FURTHER READING 183

PART II
The Flow of Genetic Information 185
Chapter 6 Genes and Genomes 187
The Structure of Eukaryotic Genes 187 Repetitive sequences 197
Introns and exons 188 Gene duplication and pseudogenes 201
Key Experiment Chromosomes and Chromatin 203
The Discovery of Introns 190
Chromatin 204
Roles of introns 192 Centromeres 208
Noncoding Sequences 194 Telomeres 213
Key Experiment SUMMARY AND KEY TERMS 214
The ENCODE Project 195 QUESTIONS 215
Noncoding RNAs 196 REFERENCES AND FURTHER READING 215

Chapter 7 Replication, Maintenance, and Rearrangements


of Genomic DNA 217
DNA Replication 217 Telomeres and telomerase: Maintaining the ends of
chromosomes 230
DNA polymerases 217
The replication fork 218 DNA Repair 232
The fidelity of replication 225 Direct reversal of DNA damage 233
Origins and the initiation of replication 226 Excision repair 235
x Contents

Translesion DNA synthesis 240 Antibody genes 245


Molecular Medicine Key Experiment
Colon Cancer and DNA Repair 241 Rearrangement of Immunoglobulin Genes 246
Repair of double-strand breaks 241 Gene amplification 252
SUMMARY AND KEY TERMS 254
DNA Rearrangements 243
QUESTIONS 255
Yeast mating types 244
REFERENCES AND FURTHER READING 256

Chapter 8 RNA Synthesis and Processing 259


Transcription in Bacteria 259 Regulation of elongation 283
RNA polymerase and transcription 260 Chromatin and Epigenetics 284
Repressors and negative control of transcription 263 Histone modifications 284
Positive control of transcription 264 Chromatin remodeling factors 289
Eukaryotic RNA Polymerases and General Histones and epigenetic inheritance 290
Transcription Factors 265 DNA methylation 292
Eukaryotic RNA polymerases 265 Noncoding RNAs 293
General transcription factors and initiation of RNA Processing and Turnover 295
transcription by RNA polymerase II 266
Processing of ribosomal and transfer RNAs 295
Transcription by RNA polymerases I and III 269
Processing of mRNA in eukaryotes 297
Regulation of Transcription in Eukaryotes 271 Splicing mechanisms 300
cis-acting regulatory sequences: promoters and Key Experiment
enhancers 271 The Discovery of snRNPs 303
Transcription factor binding sites 274 Alternative splicing 306
Transcriptional regulatory proteins 276 RNA editing 307
Key Experiment RNA degradation 309
Isolation of a Eukaryotic Transcription Factor 279 SUMMARY AND KEY TERMS 310
Structure and function of transcriptional activators 280 QUESTIONS 313
Eukaryotic repressors 282 REFERENCES AND FURTHER READING 313

Chapter 9 Protein Synthesis, Processing, and Regulation 317


Translation of mRNA 317 Protein misfolding diseases 341
Transfer RNAs 317 Molecular Medicine
The ribosome 319 Alzheimer’s Disease 342
The organization of mRNAs and the initiation of Enzymes that catalyze protein folding 343
translation 324 Protein cleavage 344
The process of translation 326 Glycosylation 346
Regulation of translation 332 Attachment of lipids 347

Protein Folding and Processing 338 Regulation of Protein Function 350


Chaperones and protein folding 338 Regulation by small molecules 350
Contents xi

Protein phosphorylation and other modifications 351 The ubiquitin-proteasome pathway 357
Key Experiment Lysosomal proteolysis 358
The Discovery of Tyrosine Kinases 353 SUMMARY AND KEY TERMS 360
Protein–protein interactions 356 QUESTIONS 361
Protein Degradation 357 REFERENCES AND FURTHER READING 362

PART III
Cell Structure and Function 365
Chapter 10 The Nucleus 367
The Nuclear Envelope and Traffic between the Chromatin localization and transcriptional activity 382
Nucleus and the Cytoplasm 367 Replication and transcription factories 385
Structure of the nuclear envelope 367 Nuclear Bodies 386
Molecular Medicine The nucleolus and rRNA 387
Nuclear Lamina Diseases 371 Polycomb bodies: Centers of transcriptional
The nuclear pore complex 372 repression 391
Selective transport of proteins to and from the Cajal bodies and speckles: Processing and storage of
nucleus 373 snRNPs 391
Key Experiment SUMMARY AND KEY TERMS 392
Identification of Nuclear Localization Signals 374 QUESTIONS 394
Transport of RNAs 378 REFERENCES AND FURTHER READING 394
Regulation of nuclear protein import 380
The Organization of Chromosomes 381
Chromosome territories 381

Chapter 11 Protein Sorting and Transport


The Endoplasmic Reticulum, Golgi Apparatus, and Lysosomes 397
The Endoplasmic Reticulum 397 Organization of the Golgi 422
The endoplasmic reticulum and protein secretion 398 Protein glycosylation within the Golgi 423
Targeting proteins to the endoplasmic reticulum 400 Lipid and polysaccharide metabolism in the Golgi 426
Key Experiment Protein sorting and export from the Golgi
The Signal Hypothesis 402 apparatus 427
Insertion of proteins into the ER membrane 405 The Mechanism of Vesicular Transport 430
Protein folding and processing in the ER 411 Experimental approaches to understanding vesicular
Quality control in the ER 414 transport 430
The smooth ER and lipid synthesis 418 Cargo selection, coat proteins, and vesicle budding 431
Export of proteins and lipids from the ER 420 Vesicle fusion 435
The Golgi Apparatus 421 Lysosomes 436
xii Contents

Lysosomal acid hydrolases 436 Phagocytosis and autophagy 440


Molecular Medicine SUMMARY AND KEY TERMS 441
Gaucher Disease 437 QUESTIONS 443
Endocytosis and lysosome formation 438 REFERENCES AND FURTHER READING 444

Chapter 12 Mitochondria, Chloroplasts, and Peroxisomes 447


Mitochondria 447 The chloroplast genome 461
Organization and function of mitochondria 448 Import and sorting of chloroplast proteins 461
The genetic system of mitochondria 450 Other plastids 465
Molecular Medicine Peroxisomes 467
Diseases of Mitochondria: Leber’s Hereditary Functions of peroxisomes 468
Optic Neuropathy 452
Peroxisome assembly 470
Protein import and mitochondrial assembly 453
Molecular Medicine
Mitochondrial lipids 457
Peroxisome Biogenesis Disorders 472
Transport of metabolites across the inner
SUMMARY AND KEY TERMS 475
membrane 457
QUESTIONS 476
Chloroplasts and Other Plastids 458 REFERENCES AND FURTHER READING 476
The structure and function of chloroplasts 459

Chapter 13 The Cytoskeleton and Cell Movement 479


Structure and Organization of Actin Microtubule Motors and Movement 508
Filaments 479 Microtubule motor proteins 508
Assembly and disassembly of actin filaments 479 Key Experiment
Organization of actin filaments 484 The Isolation of Kinesin 509
Association of actin filaments with the plasma Cargo transport and intracellular organization 512
membrane 486 Cilia and flagella 514
Microvilli 489 Reorganization of microtubules during mitosis 517
Cell surface protrusions and cell movement 490 Chromosome movement 518
Myosin Motors 493 Intermediate Filaments 520
Muscle contraction 493 Intermediate filament proteins 520
Contractile assemblies of actin and myosin Assembly of intermediate filaments 521
in nonmuscle cells 498 Intracellular organization of intermediate filaments 522
Unconventional myosins 500 Functions of intermediate filaments: Keratins and
diseases of the skin 524
Microtubules 501
Key Experiment
Structure and dynamic organization of
Expression of Mutant Keratin Causes Abnormal
microtubules 501
Skin Development 525
Assembly of microtubules 505
SUMMARY AND KEY TERMS 526
Organization of microtubules within cells 506
QUESTIONS 528
REFERENCES AND FURTHER READING 529
Contents xiii

Chapter 14 The Plasma Membrane 531


Structure of the Plasma Membrane 531 Active transport driven by ion gradients 558
The lipid bilayer 531 Endocytosis 560
Plasma membrane proteins 534 Phagocytosis 560
Plasma membrane domains 540 Clathrin-mediated endocytosis 561
Transport of Small Molecules 544 Key Experiment
The LDL Receptor 564
Facilitated diffusion and carrier proteins 544
Ion channels 546 Clathrin-independent endocytosis 567
SUMMARY AND KEY TERMS 567
Active transport driven by ATP hydrolysis 553
QUESTIONS 568
Molecular Medicine
Cystic Fibrosis 557 REFERENCES AND FURTHER READING 569

Chapter 15 Cell Walls, the Extracellular Matrix, and Cell Interactions 571
Cell Walls 571 Cell–Cell Interactions 587
Bacterial cell walls 571 Adhesion junctions 587
Eukaryotic cell walls 573 Tight junctions 590
Gap junctions 591
The Extracellular Matrix and Cell–Matrix
Interactions 577 Molecular Medicine
Gap Junction Diseases 593
Matrix structural proteins 578
Plasmodesmata 594
Matrix polysaccharides 581
SUMMARY AND KEY TERMS 595
Adhesion proteins 582
QUESTIONS 596
Cell–matrix interactions 583
REFERENCES AND FURTHER READING 597
Key Experiment
The Characterization of Integrin 584

PART IV
Cell Regulation 599
Chapter 16 Cell Signaling 601
Signaling Molecules and Their Receptors 601 Eicosanoids 608
Modes of cell–cell signaling 602 Plant hormones 610
Steroid hormones and the nuclear receptor G Proteins and Cyclic AMP Signaling 611
superfamily 603
G proteins and G protein-coupled receptors 612
Nitric oxide and carbon monoxide 605
Key Experiment
Neurotransmitters 606 G Protein-Coupled Receptors and Odor
Peptide hormones and growth factors 607 Detection 613
xiv Contents

The cAMP pathway: Second messengers and protein Phospholipase C and Ca2+ 635
phosphorylation 615
Cyclic GMP 618
Receptors Coupled to Transcription
Factors 637
Tyrosine Kinases and Signaling by MAP Kinase, The TGF-b/Smad pathway 637
PI 3-Kinase, and Phospholipase C/Calcium NF-kB signaling 638
Pathways 619 The Hedgehog, Wnt, and Notch pathways 638
Receptor tyrosine kinases 619
Nonreceptor tyrosine kinases 622
Signaling Dynamics and Networks 641
Feedback loops and signaling dynamics 641
Molecular Medicine
Cancer: Signal Transduction and the Networks and crosstalk 642
ras Oncogenes 625 SUMMARY AND KEY TERMS 644
MAP kinase pathways 626 QUESTIONS 646
The PI 3-kinase/Akt and mTOR pathways 630 REFERENCES AND FURTHER READING 647

Chapter 17 The Cell Cycle 651


The Eukaryotic Cell Cycle 651 DNA damage checkpoints 669
Phases of the cell cycle 652 The Events of M Phase 670
Regulation of the cell cycle by cell growth and Stages of mitosis 670
extracellular signals 653
Entry into mitosis 673
Cell cycle checkpoints 656
The spindle assembly checkpoint and progression to
Regulators of Cell Cycle Progression 657 anaphase 677
Protein kinases and cell cycle regulation 657 Cytokinesis 678
Key Experiment Meiosis and Fertilization 679
The Discovery of MPF 658
The process of meiosis 679
Key Experiment Regulation of oocyte meiosis 682
The Identification of Cyclin 661
Fertilization 685
Families of cyclins and cyclin-dependent kinases 663
SUMMARY AND KEY TERMS 686
Growth factors and the regulation of G1 Cdk’s 665
QUESTIONS 688
S phase and regulation of DNA replication 668
REFERENCES AND FURTHER READING 688

Chapter 18 Cell Death and Cell Renewal 691


Programmed Cell Death 691 Signaling pathways that regulate apoptosis 698
The events of apoptosis 692 Alternative pathways of programmed cell death 702
Key Experiment Stem Cells and the Maintenance of Adult
Identification of Genes Required for Programmed
Tissues 703
Cell Death 694
Proliferation of differentiated cells 703
Caspases: The executioners of apoptosis 695
Stem cells 705
Central regulators of apoptosis: The Bcl-2 family 696
Contents xv

Medical applications of adult stem cells 711 Somatic cell nuclear transfer 715
Induced pluripotent stem cells 717
Pluripotent Stem Cells, Cellular
Transdifferentiation of somatic cells 718
Reprogramming, and Regenerative
SUMMARY AND KEY TERMS 719
Medicine 712
QUESTIONS 720
Embryonic stem cells 712
REFERENCES AND FURTHER READING 721
Key Experiment
Culture of Embryonic Stem Cells 713

Chapter 19 Cancer 723


The Development and Causes of Cancer 723 Functions of oncogene products 745
Types of cancer 723 Tumor Suppressor Genes 749
The development of cancer 725 Identification of tumor suppressor genes 750
Causes of cancer 727 Functions of tumor suppressor gene products 753
Properties of cancer cells 728 Cancer genomics 757
Transformation of cells in culture 732
Molecular Approaches to Cancer
Tumor Viruses 733 Treatment 758
Hepatitis B and C viruses 733 Prevention and early detection 759
Small DNA tumor viruses 734 Treatment 760
Herpesviruses 735 Molecular Medicine
Retroviruses 736 Imatinib: Cancer Treatment Targeted against
the bcr/abl Oncogene 763
Oncogenes 737
SUMMARY AND KEY TERMS 765
Retroviral oncogenes 737
QUESTIONS 767
Proto-oncogenes 739
REFERENCES AND FURTHER READING 767
Key Experiment
The Discovery of Proto-Oncogenes 740
Oncogenes in human cancer 741

Answers to Questions 771 Illustration Credits 803


Glossary 783 Index 805
Preface
Learning cell biology can be a daunting task because the field is so vast and
rapidly moving, characterized by a continual explosion of new information.
The challenge is how to master the fundamental concepts without becoming
bogged down in details. Students need to understand the principles of cell
biology and be able to appreciate new advances, rather than just memoriz-
ing “the facts” as we see them today. At the same time, the material must be
presented in sufficient depth to thoughtfully engage students and provide a
sound basis for further studies. The Cell provides a balance of concepts and
details designed to meet these needs of today’s students and their teachers.
The Seventh Edition of The Cell continues the central goal of helping stu-
dents understand the principles and concepts of contemporary cell biology
while gaining an appreciation of the excitement and importance of research
in this rapidly moving field. Our understanding of cell and molecular biology
has progressed in many ways over the last three years, and these important
advances have been incorporated into the current edition. Some of the most
striking advances have continued to come from progress in genomics and
proteomics. A new chapter in the current edition—Genomics, Proteomics,
and Systems Biology—focuses on these rapidly advancing areas and includes
discussion of the growing field of synthetic biology. Other notable advances
covered in the current edition include super-resolution microscopy, genome
editing by the CRISPR/Cas system, epigenetics, the organization of chro-
mosome domains in the nucleus, and an updated discussion of prions and
Alzheimer’s disease.
These and other advances have been incorporated into the Seventh
Edition of The Cell not only to provide current information but, even more
importantly, to give students a sense of the excitement and challenges of
research in our field. At the same time, I have sought to minimize details and
focus on concepts in order to ensure that The Cell remains an accessible and
readable text for undergraduates who are taking their first course in cell and
molecular biology. Distinguishing features of The Cell include the Molecular
Medicine and Key Experiment essays, which highlight clinical applications
and describe seminal research papers, respectively. Together with additional
experiments discussed throughout the text, these essays give the students
a sense of how progress in our field is made and a feel for how hypotheses
are framed and results interpreted. As in previous editions, each chapter
also includes several short “FYIs” designed to highlight areas of interest or
medical relevance, as well as a set of questions at the end of each chapter
with answers at the back of the book. In keeping with The Cell’s focus on
experimental analysis, many of these questions ask the student to think of
experiments or interpret results, as well as providing a review of the material.
A new feature of this edition is the addition of Data Analysis Problems to the
Instructor’s Resource Library. These problems present data and figures from
xviii Preface

original research papers and engage students in the analysis of experimental


methods and results. They also provide excellent material for discussions
and opportunities for student participation in active learning.
My most important goal in writing has been to convey the excitement
and challenges of research in contemporary cell and molecular biology. The
opportunities in our field are greater than ever, and I hope The Cell stimulates
today’s students to contribute to the research on which future texts will be based.

Acknowledgments
I am particularly grateful to Alexandra Adams for carefully reviewing the
entire text of the Seventh Edition of The Cell. The book has also benefited from
the comments and suggestions of reviewers, colleagues, and instructors who
used the previous edition. I am pleased to thank the following reviewers for
their thoughtful comments and advice:

Sage Arbor* Marian University


Rebecca Balish Miami University
Esther Biswas-Fiss Thomas Jefferson University
Edward Bonder Rutgers University
Lucinda Carnell Central Washington University
Xiao-Wen Cheng Miami University
Heather Cook* Wagner College
Gary Coombs* Waldorf College
Paul Davis University of Nebraska at Omaha
Jennifer Freytag* The Sage Colleges
Michael Garcia University of Missouri
David Gardner Marian University College of Osteopathic Medicine
Brittany Gasper Florida Southern College
Mark Grimes University of Montana
Karen Guzman Campbell University
Neil Haave* University of Alberta
Joyce Hardy* Chadron State College
Amber Heck Rocky Vista University College of
Osteopathic Medicine
Jose Hernandez Midwestern University
Peter Hoffman Notre Dame of Maryland University
Lisa Kadlec* Wilkes University
Ondra Kielbasa Alvernia University
Janet Kirkley Knox College
Jon Lowrance Lipscomb University
Kaushiki Menon Mt. San Antonio College
Vida Mingo* Columbia College
Craig Moyer Western Washington University
Perpetua Muganda North Carolina A&T State University
Laura Rhoads* State University of New York at Potsdam
Robert Roberson Arizona State University
German Rosas-Acosta The University of Texas at El Paso
D. Scott Samuels University of Montana
Ken Savage The University of British Columbia
Alexander Schreiber St. Lawrence University
Deborah Schulman Lake Erie College
Preface xix

José Serrano-Moreno Western Washington University


Vishal Shah Dowling College
Mohammad Rafique Uddin LeMoyne–Owen College
Shannon Vandaveer University of Kansas
Thomas Wolkow University of Colorado, Colorado Springs
Kathleen Wood* University of Mary Hardin-Baylor
Robin Young The University of British Columbia
*Reviewed chapter(s) and Data Analysis Problems.

It is also a pleasure to thank our publisher for continuing support. Chelsea


Holabird did a superb editorial job. Jen Basil-Whitaker created a beautiful
design and page layouts. Andy Sinauer, Dean Scudder, and Christopher
Small were once again a real pleasure to work with.
Geoffrey M. Cooper
September, 2015
Organization and Features of
The Cell, Seventh Edition
The Cell has been designed to be an approachable and teachable text that
can be covered in a single semester while allowing students to master the
material in the entire book. It is assumed that most students will have had
introductory biology and general chemistry courses, but will not have had
previous courses in organic chemistry, biochemistry, or molecular biology.
Several aspects of the organization and features of the book will help students
to approach and understand its subject matter.

Organization
The Cell is divided into four parts, each of which is self-contained, so that
the order and emphasis of topics can be easily varied according to the needs
of individual courses.
Part I provides background chapters on the evolution of cells, methods for
studying cells, the chemistry of cells, the fundamentals of modern molecular
biology, and the fields of genomics and systems biology. For those students
who have a strong background from either a comprehensive introductory
biology course or a previous course in molecular biology, various parts of
these chapters can be skipped or used for review.
Part II focuses on the molecular biology of cells and contains chapters
dealing with genome organization and sequences; DNA replication, repair,
and recombination; transcription and RNA processing; and the synthesis,
processing, and regulation of proteins. The order of chapters follows the flow
of genetic information (DNA RNA protein) and provides a concise but
up-to-date overview of these topics.
Part III contains the core block of chapters on cell structure and function,
including chapters on the nucleus, cytoplasmic organelles, the cytoskeleton,
the plasma membrane, and the extracellular matrix. This part of the book
starts with coverage of the nucleus, which puts the molecular biology of Part
II within the context of the eukaryotic cell, and then works outward through
cytoplasmic organelles and the cytoskeleton to the plasma membrane and
the exterior of the cell. These chapters are relatively self-contained, however,
and could be used in a different order should that be more appropriate for
a particular course.
Finally, Part IV focuses on the exciting and fast-moving area of cell
regulation, including coverage of topics such as cell signaling, the cell cycle,
programmed cell death, and stem cells. This part of the book concludes with
a chapter on cancer, which synthesizes the consequences of defects in basic
cell regulatory mechanisms.
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up to your Harding reputations, as far as I can see—Babbie the
Butterfly, Madeline the Bohemian, Betty a Benevolent Adventurer.”
“And the moral of that is,” put in Babbie quickly, “what you are at
home, that you will be abroad.”
“Unless you drop all your individuality and become a Tourist, with
a capital T,” added Roberta.
“Or change your spots and turn from a man-hater into a fiancée,”
suggested Bob.
“That’s not changing your spots,” declared Mary wisely. “It’s just
making up your mind, isn’t it, Babe?”
“How in the world did you know that, Mary Brooks?” demanded
Babe in such awe-struck tones that her friends shrieked with
laughter, and Dr. Hinsdale came out from his study to ask about the
joke.
The girls had intended to leave early the next afternoon, but
when Georgia Ames appeared, hovering in the Belden House hall,
before dinner was over, and announced that she was giving a
gargoyle party for them that evening, why of course there was
nothing to do but insist that the gargoyle party should be a “small
and early,” and rush to the station to countermand orders for
carriages, and find out about making connections with sleepers at
the junction.
“For we’re not so young as we were once,” said Roberta, hugging
Betty. “We don’t have to be met at Harding by the registrar, and we
may travel at night if we like, as long as two go one way and three
the other.”
The gargoyle party was as mysterious as Mary Brooks’s historic
hair-raising had been. Mary almost wept when Georgia asked her,
and she was obliged to decline because of a previous dinner
engagement—not to mention the dignity of her position. She solaced
herself by making an elaborate costume for Eugenia Ford, a pretty
little freshman who, when Georgia asked her to the party, thanked
her gravely and explained that if gargoyles had anything to do with
gargles she wouldn’t come, because she never could manage to do
it—her throat must be queer. Most of the other guests professed
hapless ignorance of what a gargoyle might be, but Georgia referred
them easily to Bob’s cherished imp, which she had borrowed for the
occasion, together with some post-cards of other grotesque figures.
“Just run in any time this afternoon, and look them over,” she
urged, “and come in costume to-night, if you can. If not, it doesn’t
matter. Mrs. Hinsdale is going to offer a prize for the best one,
though.”
So the chosen few cast English Lit. papers and a possible—nay,
probable—written review in Psych. to the winds, journeyed down-
town to buy masks and draperies, and preëmpted all the desirable
perches in Georgia’s room, marking them with big “Engaged” signs,
which came loose when the wind blew in next time the door was
opened, and gave the room a disconcerting air of having been
snowed under, when Georgia got back to it just before tea.
“But we had to do it,” Eugenia Ford explained, as she helped
Georgia put things to rights for the evening, “because the whole
point of a gargoyle is that it stands somewhere. Lucile Merrifield said
so. And the way you put on your costume makes a difference about
where you are to sit. No, the other way around.”
“Conversely, you mean, my child,” amended Georgia, pleasantly,
putting Mary’s five-pound box of Huyler’s on the chiffonier.
“But that’s got to be cleared off,” objected Eugenia. “That’s Miss
Bob Parker’s place. We all wanted it, but she got it tagged first.
Belden House Annie promised her a step-ladder to climb up by, but
she said a chair would do.”
Georgia sighed and dumped the ornaments of the dresser top,
cover and all, into her upper drawer. “A gargoyle party is a thing that
grows on your hands,” she said sadly. “Let’s go and eat. If there’s
anything else to clear off, we’ll do it later.”
When the gargoyle party opened it was certain that, whether or
not it had grown on Georgia’s hands, it was every bit her room could
hold. Betty and Babbie, who had been too busy enjoying Harding to
bother about costumes, were the only guests who were not wearing
some sort of fantastic disguise. Bob had bought a box of paints and
made her own mask, modeling it and her drapery of brown denim
after the imp that the “B. A.’s Abroad” had given her. Eugenia Ford
was a gryphon,—or at least Mary Brooks said so,—with the most
beautiful pair of wings that had ever appeared at a Harding party.
Polly Eastman was the elephant that sits on the tower of Notre
Dame. Georgia had planned to be the other half of the elephant, in
accordance with Harding usage in the matter of elephants and other
four-footed creatures. But at the last minute she discovered that the
Notre Dame elephant wasn’t four-footed.
“Gargoyles never are,” said Lucile wisely—it was she who had
pointed out the mistake. “But never mind, Georgia. You can be one
of my two heads. I was going to be a two-headed beast if I could.
Only Vesta White changed her mind afterward and wanted to be an
eagle.”
There were other gargoyles, as impossible to classify as the real
ones, and they squatted in rows on Georgia’s bed and her big
window-box, popped up mysteriously from behind her desk, or
lounged in strange attitudes in her easy chairs. Bob Parker actually
did get up on the chiffonier, off the edge of which she hung in such
realistic gargoyle style that the judges, Babbie and Betty,
unhesitatingly awarded her the prize.
“Not a bit fair,” objected young Eugenia, flapping her beautiful
gryphon’s wings disconsolately. “We should all have looked a lot
grander on chiffoniers.”
“But you weren’t all clever enough to grab the one there was,” put
in Georgia pacifically.
“Having a gargoyle of your own makes you notice the attitudes
more,” declared Bob proudly. “Never mind, Miss Ford. The prize is
candy, and we’ll pass it around while we wait for Georgia’s
refreshments to materialize.”
“You haven’t forgotten your Harding manners, Bob,” said Betty
severely.
“No, you don’t any of you act a bit like alums,” declared a tall
junior, taking off her mask to breathe.
“You lovely thing!” cried Bob, scrambling down from the chiffonier
to give the appreciative junior first choice of the prize candy.
And then the gargoyles had a dance and a parade, and delicious
“eats,” on which Georgia had rashly spent all that was left of her
month’s allowance. And after that, when the five 19—’s were having
the very best time of all, just sitting around talking and realizing what
a dear, dear place Harding was, it was time to pull Bob out of her
beloved costume and rush for trains.
Later in the evening the five classmates sat in the station at the
junction, Babe and Betty waiting to go west, Bob, Babbie and
Roberta bound for New York.
Babbie looked critically at Babe and Betty. “I shall tell mother that
it worked,” she said. “You went to bed at three, and got up at seven
this morning to go canoeing. You’ve eaten four meals to-day and as
many ices. You’ve been horseback and trolley-riding. You’ve made
dozens of calls. It’s now ten p. m., and you’re fresh as the daisies in
Oban. How’s that for the Harding cure?”
“Don’t you feel exactly as if it was some June?” demanded Bob.
“Not last June, but a regular June, you know, and we were all just
going home for the summer.”
“Exactly,” agreed everybody, and then a sleepy silence settled
upon the group.
“What were those things we had in the ‘Rise of the Drama’
course?” asked Betty Wales suddenly. “Not intervals, but something
like that.”
“You mean Interludes, don’t you?” asked Roberta. “They came
right after the Moralities.”
Betty nodded. “That’s what this summer has been—an Interlude.”
“With Babe for the fascinating heroine,” put in Babbie.
“Yes,” agreed Betty hastily. “And when I get home to-morrow the
real business of life is going to begin.”
“Act I, Scene I, Life of Betty Wales, B. A.,” said Roberta. “Doesn’t
that sound serious? But it won’t be. You’ll play tennis with Nan, and
go to dances with your brother and other people’s brothers, and
amuse that darling little sister of yours, and be nice to everybody
who needs it, just as you always have, except that you won’t be
home on a snippy little vacation.”
“Oh, I hope so,” said Betty, laughing at Roberta’s choice of
details. “But then I want to do something that counts, too.”
“You’re always doing things that count,” Babe declared, giving
her a loving little squeeze.
“That was just fun,” Betty reminded her for the hundredth time at
least.
“But if fun counts, it counts,” declared Roberta. “Just ask
Madeline Ayres if it doesn’t. If you can make fun out of hard work,
then, according to Madeline, you really know how to live.”
“But we’re not the working contingent,” objected Babbie. “K. and
Rachel and Helen are the workers.”
“They are!” breathed Bob indignantly. “Just try taking care of
certain fresh-air youngsters for two weeks.”
“Or typewriting most particular briefs for your most particular
father, who always wants things in a terrific hurry,” added Roberta.
Betty considered. “I’ve helped in little ways of course, but I never
did any one big thing. I’m going to now, though.”
“Here’s to a winter of hard work!” cried Babe. “I shall have to sew,
and I hate it.”
“But you must make fun out of it all the same,” Betty told her, with
the flash of gay courage in her eyes that had won over Mr. Morton. “I
shall, no matter what happens, and whatever we do, think of the fun
we’ll have talking it over when we all get together again. Oh, is that
our train, Babe?” And with her curls flying and her eyes dancing with
eagerness Betty Wales turned merrily from her happy summer’s
Interlude to “the real business of life.”
THE END
Transcriber’s Notes:
Minor corrections (addition or deletion) of double quote marks have been made on pages
188, 196, 230 and 317, to conform to accepted usage.
Splended, on page 153, has been changed to splendid.
Cooperation, on page 218, has been changed to coöperation, to conform to other
occurrences in this e-book.
On page 270, Louxembourg has been changed to Luxembourg.
All other hyphenation and variant and archaic spellings have been retained as typeset.
Illustrations have been moved to avoid interrupting paragraphs.
*** END OF THE PROJECT GUTENBERG EBOOK BETTY WALES,
B. A.: A STORY FOR GIRLS ***

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