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GENETICS: ANALYSIS & PRINCIPLES, FOURTH EDITION
Published by McGraw-Hill, a business unit of The McGraw-Hill Companies, Inc., 1221 Avenue of the Americas, New
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This book is printed on acid-free paper.
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ISBN 978–0–07–352528–0
MHID 0–07–352528–6
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Library of Congress Cataloging-in-Publication Data
Brooker, Robert J.
Genetics : analysis & principles / Robert J. Brooker. — 4th ed.
p. cm.
Includes index.
ISBN 978–0–07–352528–0 — ISBN 0–07–352528–6 (hard copy : alk. paper) 1. Genetics. I. Title.
QH430.B766 2012
576.5--dc22
2010015380
www.mhhe.com
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B R I E F C O N T E N T S
::
PART I INTRODUCTION PART IV MOLECULAR PROPERTIES
OF GENES
1 Overview of Genetics 1
12 Gene Transcription and RNA Modification 299
PART II PATTERNS OF INHERITANCE
13 Translation of mRNA 326
2 Mendelian Inheritance 17
14 Gene Regulation in Bacteria
3 Reproduction and Chromosome and Bacteriophages 359
Transmission 44
15 Gene Regulation in Eukaryotes 390
4 Extensions of Mendelian Inheritance 71
16 Gene Mutation and DNA Repair 424
5 Non-Mendelian Inheritance 100
17 Recombination and Transposition
6 Genetic Linkage and Mapping in at the Molecular Level 457
Eukaryotes 126
PART V GENETIC TECHNOLOGIES
7 Genetic Transfer and Mapping in Bacteria
and Bacteriophages 160 18 Recombinant DNA Technology 484
8 Variation in Chromosome Structure 19 Biotechnology 518
and Number 189
20 Genomics I: Analysis of DNA 544
PART III MOLECULAR STRUCTURE AND 21 Genomics II: Functional Genomics, Proteomics,
REPLICATION OF THE GENETIC and Bioinformatics 574
MATERIAL
PART VI G ENETIC ANALYSIS
9 Molecular Structure of DNA and RNA 222
OF INDIVIDUALS AND
10 Chromosome Organization and Molecular POPULATIONS
Structure 247
22 Medical Genetics and Cancer 602
11 DNA Replication 270
23 Developmental Genetics 637
24 Population Genetics 670
25 Quantitative Genetics 700
26 Evolutionary Genetics 730
iii
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T A B L E O F C O N T E N T S
::
Preface vii Experiment 7A Conjugation
4 Experiments Can Map Genes Along
A Visual Guide to Genetics: Analysis & EXTENSIONS OF MENDELIAN the E. coli Chromosome 167
Principles xiv INHERITANCE 71 7.2 Intragenic Mapping in
4.1 Inheritance Patterns Bacteriophages 176
PART I
of Single Genes 71
INTRODUCTION 4.2 Gene Interactions 86
8
1 Experiment 4A Bridges Observed an VARIATION IN CHROMOSOME
8:4:3:1 Ratio Because the Cream-Eye STRUCTURE AND NUMBER 189
OVERVIEW OF GENETICS 1
Gene Can Modify the X-Linked Eosin
1.1 The Relationship Between Genes Allele But Not the Red or 8.1 Variation in Chromosome
White Alleles 89 Structure 189
and Traits 4
1.2 Fields of Genetics 10 Experiment 8A Comparative
5 Genomic Hybridization Is Used to
NON-MENDELIAN Detect Chromosome Deletions and
INHERITANCE 100 Duplications 195
PART II 8.2 Variation in Chromosome
5.1 Maternal Effect 100
PATTERNS OF INHERITANCE 17 Number 203
5.2 Epigenetic Inheritance 103
8.3 Natural and Experimental Ways to
2
Experiment 5A In Adult Female
Produce Variations in Chromosome
Mammals, One X Chromosome Has
MENDELIAN INHERITANCE 17 Number 208
Been Permanently Inactivated 105
2.1 Mendel’s Laws of Inheritance 18 5.3 Extranuclear Inheritance 113
Experiment 2A Mendel Followed the
PART III
Outcome of a Single Character for Two
6
Generations 21 GENETIC LINKAGE AND MAPPING MOLECULAR STRUCTURE AND
Experiment 2B Mendel Also Analyzed IN EUKARYOTES 126 REPLICATION OF THE GENETIC
Crosses Involving Two Different MATERIAL 222
Characters 25 6.1 Linkage and Crossing Over 126
2.2 Probability and Statistics 30 Experiment 6A Creighton and 9
McClintock Showed That Crossing MOLECULAR STRUCTURE OF DNA
Over Produced New Combinations of AND RNA 222
3
Alleles and Resulted in the Exchange
REPRODUCTION 9.1 Identification of DNA as the Genetic
of Segments Between Homologous
AND CHROMOSOME Chromosomes 133 Material 222
TRANSMISSION 44
6.2 Genetic Mapping in Plants Experiment 9A Hershey and Chase
3.1 General Features of and Animals 136 Provided Evidence That DNA Is the
Genetic Material of T2 Phage 225
Chromosomes 44 Experiment 6B Alfred Sturtevant Used
9.2 Nucleic Acid Structure 229
3.2 Cell Division 48 the Frequency of Crossing Over in
Dihybrid Crosses to Produce the First Experiment 9B Chargaff Found That
3.3 Sexual Reproduction 54
Genetic Map 138 DNA Has a Biochemical Composition in
3.4 The Chromosome Theory 6.3 Genetic Mapping in Haploid Which the Amount of A Equals T and
of Inheritance and Sex the Amount of G Equals C 232
Eukaryotes 143
Chromosomes 60
6.4 Mitotic Recombination 149
Experiment 3A Morgan’s Experiments 10
Showed a Connection Between a CHROMOSOME
Genetic Trait and the Inheritance of a ORGANIZATION
7
Sex Chromosome in Drosophila 64 AND MOLECULAR
GENETIC TRANSFER AND
STRUCTURE 247
MAPPING IN BACTERIA AND
BACTERIOPHAGES 160
10.1 Viral Genomes 247
7.1 Genetic Transfer and Mapping 10.2 Bacterial Chromosomes 249
iv
in Bacteria 161
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TABLE OF CONTENTS v
10.3 Eukaryotic Chromosomes 252 14.2 Translational and Posttranslational
PART V
Experiment 10A The Repeating Regulation 375
Nucleosome Structure Is Revealed by 14.3 Riboswitches 377 GENE T IC TECHNOLOGIES 484
Digestion of the Linker Region 257 14.4 Gene Regulation in the Bacteriophage 18
11 Reproductive Cycle 378 RECOMBINANT DNA
TECHNOLOGY 484
DNA REPLICATION 270
15 18.1 Gene Cloning Using Vectors 485
11.1 Structural Overview of DNA GENE REGULATION IN 18.2 Polymerase Chain Reaction 491
Replication 270 EUKARYOTES 390
Experiment 18A Early Attempts at
E xperiment 11A Three Different Models
15.1 Regulatory Transcription Factors 391 Monitoring the Course of PCR Used
Were Proposed That Described the Net
Result of DNA Replication 272 15.2 Chromatin Remodeling, Ethidium Bromide as a Detector 498
11.2 Bacterial DNA Replication 274 Histone Variation, and Histone 18.3 DNA Libraries and Blotting
Modification 397 Methods 499
Experiment 11B DNA Replication Can
Be Studied in Vitro 285 15.3 DNA Methylation 403 18.4 Methods for Analyzing DNA- and RNA-
11.3 Eukaryotic DNA Replication 288 15.4 Insulators 406 Binding Proteins 505
15.5 Regulation of RNA Processing, RNA 18.5 DNA Sequencing and Site-Directed
Stability, and Translation 407 Mutagenesis 507
PART IV Experiment 15A Fire and Mello Show 19
That Double-Stranded RNA Is More
MOLECULAR PROPERTIES
Potent Than Antisense RNA at Silencing BIOTECHNOLOGY 518
OF GENES 299
mRNA 411
19.1 Uses of Microorganisms
12 in Biotechnology 518
GENE TRANSCRIPTION AND
16 19.2 Genetically Modified Animals 522
RNA MODIFICATION 299 GENE MUTATION
AND DNA REPAIR 424 19.3 Reproductive Cloning and Stem
12.1 Overview of Transcription 300 Cells 527
12.2 Transcription in Bacteria 302 16.1 Consequences of Mutation 425 19.4 Genetically Modified Plants 532
12.3 Transcription in Eukaryotes 307 16.2 Occurrence and Causes of 19.5 Human Gene Therapy 536
12.4 RNA Modification 310 Mutation 431 Experiment 19A Adenosine Deaminase
Experiment 16A X-Rays Were the First
Experiment 12A Introns Were Deficiency Was the First Inherited Disease
Environmental Agent Shown to Cause
Experimentally Identified via Treated with Gene Therapy 538
Induced Mutations 439
Microscopy 313
13 16.3 DNA Repair 443 20
GENOMICS I: ANALYSIS
OF DNA 544
TRANSLATION OF mRNA 326
17
13.1 The Genetic Basis for Protein RECOMBINATION AND 20.1 Overview of Chromosome Mapping 545
Synthesis 326 TRANSPOSITION AT THE 20.2 Cytogenetic Mapping Via
MOLECULAR LEVEL 457
Experiment 13A Synthetic RNA Helped Microscopy 545
to Decipher the Genetic Code 332 17.1 Homologous Recombination 457 20.3 Linkage Mapping Via Crosses 547
13.2 Structure and Function of tRNA 340 Experiment 17A The Staining of 20.4 Physical Mapping Via Cloning 553
Experiment 13B tRNA Functions as the Harlequin Chromosomes Can Reveal 20.5 Genome-Sequencing Projects 559
Adaptor Molecule Involved in Codon Recombination Between Sister
Experiment 20A Venter, Smith, and
Recognition 340 Chromatids 458
Colleagues Sequenced the First
13.3 Ribosome Structure and 17.2 Site-Specific Recombination 466 Genome in 1995 559
Assembly 345 17.3 Transposition 468
21
13.4 Stages of Translation 347 E xperiment 17B McClintock Found That GENOMICS II: FUNCTIONAL
Chromosomes of Corn Plants Contain GENOMICS, PROTEOMICS,
14 Loci That Can Move 468 AND BIOINFORMATICS 574
GENE REGULATION
IN BACTERIA AND 21.1 Functional Genomics 575
BACTERIOPHAGES 359
Experiment 21A The Coordinate
14.1 Transcriptional Regulation 360 Regulation of Many Genes Is Revealed
by a DNA Microarray Analysis 577
Experiment 14A The lacI Gene Encodes
21.2 Proteomics 583
a Diffusible Repressor Protein 365
21.3 Bioinformatics 587
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vi TABLE OF CONTENTS
23.3 Vertebrate Development 652 26
PART VI EVOLUTIONARY
23.4 Plant Development 656
GENETICS 730
GENETIC ANALYSIS
23.5 Sex Determination in Animals
OF INDIVIDUALS AND and Plants 659 26.1 Origin of Species 731
POPULATIONS 602 26.2 Phylogenetic Trees 738
24
26.3 Molecular Evolution 744
22
POPULATION GENETICS 670 Experiment 26A Scientists Can
MEDICAL GENETICS
Analyze Ancient DNA to Examine the
AND CANCER 602 24.1 Genes in Populations and the
Relationships Between Living and
22.1 Inheritance Patterns of Genetic Hardy-Weinberg Equation 670 Extinct Flightless Birds 748
Diseases 603 24.2 Factors That Change Allele 26.4 Evo-Devo: Evolutionary Developmental
22.2 Detection of Disease-Causing and Genotype Frequencies in Biology 753
Populations 675
Alleles 609
22.3 Prions 613 Experiment 24A The Grants Have Appendix A
Observed Natural Selection in
22.4 Genetic Basis of Cancer 614 Galápagos Finches 686 Experimental Techniques A-1
Experiment 22A DNA Isolated from 24.3 Sources of New Genetic Appendix B
Malignant Mouse Cells Can Transform
Variation 689 Solutions to Even-Numbered
Normal Mouse Cells into Malignant
Problems A-8
Cells 616
25
QUANTITATIVE Glossary G-1
23 GENETICS 700
Credits C-1
DEVELOPMENTAL
GENETICS 637 25.1 Quantitative Traits 700 Index I-1
23.1 Overview of Animal 25.2 Polygenic Inheritance 705
Development 637 Experiment 25A Polygenic Inheritance
Explains DDT Resistance
23.2 Invertebrate Development 640
in Drosophila 708
Experiment 23A Heterochronic
25.3 Heritability 711
Mutations Disrupt the Timing of
Experiment 25B Heritability of Dermal
Developmental Changes
Ridge Count in Human Fingerprints
in C. elegans 650
Is Very High 716
ABOUT THE AUTHOR
Robert J. Brooker is a professor in the Department of
Genetics, Cell Biology, and Development at the University of
Minnesota–Minneapolis. He received his B.A. in biology from
Wittenberg University in 1978 and his Ph.D. in genetics from
Yale University in 1983. At Harvard, he conducted postdoctoral
studies on the lactose permease, which is the product of the lacY
gene of the lac operon. He continues his work on transporters at
the University of Minnesota. Dr. Brooker’s laboratory primarily
investigates the structure, function, and regulation of iron
transporters found in bacteria and C. elegans. At the University of
Minnesota he teaches undergraduate courses in biology, genetics,
and cell biology.
DEDICATION
To my wife, Deborah, and our children, Daniel, Nathan, and Sarah
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P R E F A C E
::
I
Red White
P generation
n the fourth edition of Genetics: Analysis & Principles, the
content has been updated to reflect current trends in the field. In CRCR x CWCW
addition, the presentation of the content has been improved in
a way that fosters active learning. As an author, researcher, and
teacher, I want a textbook that gets students actively involved Gametes CR CW
in learning genetics. To achieve this goal, I have worked with a
talented team of editors, illustrators, and media specialists who
have helped me to make the fourth edition of Genetics: Analy-
sis & Principles a fun learning tool. The features that we feel are Pink
most appealing to students, and which have been added to or
F generation
improved on in the fourth edition, are the following. 1
(cid:129) Interactive exercises Education specialists have crafted CRCW
interactive exercises in which the students can make their
own choices in problem-solving activities and predict what
the outcomes will be. Previously, these exercises focused Gametes CR or CW
on inheritance patterns and human genetic diseases. (For
example, see Chapters 4 and 22 .) For the fourth edition, Self-fertilization
we have also added many new interactive exercises for the
molecular chapters.
Sperm
(cid:129) Animations Our media specialists have created over F generation CR CW
2
50 animations for a variety of genetic processes. These
animations were made specifically for this textbook and use
the art from the textbook. The animations make many of CR
the figures in the textbook “come to life.” CRCR CRCW
Egg
(cid:129) Experiments As in the previous editions, each chapter
(beginning with Chapter 2 ) incorporates one or two experi-
CW
ments that are presented according to the scientific method.
CRCW CWCW
These experiments are not “boxed off” from the rest of
the chapter. Rather, they are integrated within the chapters
and flow with the rest of the text. As you are reading the FIGURE 4.3 Incomplete dominance in the
experiments, you will simultaneously explore the scientific four-o’clock plant, Mirabilis jalapa.
method and the genetic principles that have been discovered Genes → Traits When two different homozygotes (CRCR and
using this approach. For students, I hope this textbook helps CWCW) are crossed, the resulting heterozygote, CRCW, has an intermediate phenotype
of pink flowers. In this case, 50% of the functional protein encoded by the CR allele
you to see the fundamental connection between scientific
is not sufficient to produce a red phenotype.
analysis and principles. For both students and instructors, I
expect that this strategy makes genetics much more fun to
explore.
(cid:129) Art The art has been further refined for clarity and com-
pleteness. This makes it easier and more fun for students to
Sometimes they are questions that the students might ask
study the illustrations without having to go back and forth
themselves when they are learning about genetics.
between the art and the text.
(cid:129) Engaging text As in previous editions, a strong effort has Overall, an effective textbook needs to accomplish three
been made in the fourth edition to pepper the text with goals. First, it needs to provide comprehensive, accurate, and up-
questions. Sometimes these are questions that scientists to-date content in its field. Second, it needs to expose students to
considered when they were conducting their research. the techniques and skills they will need to become successful in
vii
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