Essentials of Genetics, Global Edition

606 Pages·2016·29.949 MB·English

by  William S. Klug, Michael R. Cummings, Charlotte A. Spencer, Michael A. Palladino

Brief Contents
1
Introduction to Genetics 17
2
Mitosis and Meiosis 28
3
Mendelian Genetics 47
4
Modification of Mendelian Ratios 69
5
Sex Determination and Sex Chromosomes 100
6
Chromosome Mutations: Variation in Number and Arrangement 115
7
Linkage and Chromosome Mapping in Eukaryotes 136
8
Genetic Analysis and Mapping in Bacteria and Bacteriophages 159
9
DNA Structure and Analysis 176
10
DNA Replication 196
11
Chromosome Structure and DNA Sequence Organization 215
12
The Genetic Code and Transcription 231
13
Translation and Proteins 254
14
Gene Mutation, DNA Repair, and Transposition 273
15
Regulation of Gene Expression 296
16
The Genetics of Cancer 323
17
Recombinant DNA Technology 338
18
Genomics, Bioinformatics, and Proteomics 361
19
Applications and Ethics of Genetic Engineering and Biotechnology 394
20
Developmental Genetics 419
21
Quantitative Genetics and Multifactorial Traits 438
22
Population and Evolutionary Genetics 457
Special TopicS in modern GeneTicS
1 Epigenetics 480
2 Emerging Roles of RNA 490
3 DNA Forensics 503
4 Genomics and Personalized Medicine 513
5 Genetically Modified Foods 523
6 Gene Therapy 535
Appendix Solutions to Selected problems and discussion Questions A-1
GloSSAry G-1
CreditS C-1
index i-1
ESSENTIALS
GENETICS
of
Ninth Edition
Global Edition
William S. Klug
The College of New Jersey
Michael R. Cummings
Illinois Institute of Technology
Charlotte A. Spencer
University of Alberta
Michael A. Palladino
Monmouth University
with contributions by
Darrell Killian
Colorado College
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Authorized adaptation from the United States edition, entitled Essentials of Genetics, 9th edition,
ISBN 978-0-134-04779-9, by William S. Klug, Michael R. Cummings, Charlotte A. Spencer, and Michael A.
Palladino, published by Pearson Education © 2016.
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About the Authors
William S. Klug  is an Emeritus Professor of Biology at The Charlotte A. Spencer  is a retired Associate Professor
College of New Jersey (formerly Trenton State College) in Ewing, from the Department of Oncology at the University of Alberta
New Jersey, where he served as Chair of the Biology Department in Edmonton, Alberta, Canada. She has also served as a facul-
for 17 years. He received his B.A. degree in Biology from Wabash ty member in the Department of Biochemistry at the Univer-
College in Crawfordsville, Indiana, and his Ph.D. from North- sity of Alberta. She received her B.Sc. in Microbiology from
western University in Evanston, Illinois. Prior to coming to The the University of British Columbia and her Ph.D. in Genet-
College of New Jersey, he was on the faculty of Wabash College ics from the University of Alberta, followed by postdoctoral
as an Assistant Professor, where he first taught genetics, as well training at the Fred Hutchinson Cancer Research Center in
as general biology and electron microscopy. His research inter- Seattle, Washington. Her research interests involve the regu-
ests have involved ultrastructural and molecular genetic stud- lation of RNA polymerase II transcription in cancer cells, cells
ies of development, utilizing oogenesis in Drosophila as a model infected with DNA viruses, and cells traversing the mitotic
system. He has taught the genetics course as well as the senior phase of the cell cycle. She has taught courses in biochem-
capstone seminar course in Human and Molecular Genetics to istry, genetics, molecular biology, and oncology, at both un-
undergraduate biology majors for over four decades. He was the dergraduate and graduate levels. In addition, she has written
recipient in 2001 of the first annual teaching award given at The booklets in the Prentice Hall Exploring Biology series, which
College of New Jersey, granted to the faculty member who “most are aimed at the undergraduate nonmajor level.
challenges students to achieve high standards.” He also received
the 2004 Outstanding Professor Award from Sigma Pi Interna- Michael A. Palladino  is Dean of the School of Sci-
tional, and in the same year, he was nominated as the Educator ence and Professor of Biology at Monmouth University in
of the Year, an award given by the Research and Development West Long Branch, New Jersey. He received his B.S. degree
Council of New Jersey. in Biology from Trenton State College (now known as The
College of New Jersey) and his Ph.D. in Anatomy and Cell
Michael R. Cummings  is Research Professor in the De- Biology from the University of Virginia. He directs an active
partment of Biological, Chemical, and Physical Sciences at Illi- laboratory of undergraduate student researchers study-
nois Institute of Technology, Chicago, Illinois. For more than 25 ing molecular mechanisms involved in innate immunity of
years, he was a faculty member in the Department of Biological mammalian male reproductive organs and genes involved
Sciences and in the Department of Molecular Genetics at the in oxygen homeostasis and ischemic injury of the testis.
University of Illinois at Chicago. He has also served on the fac- He has taught a wide range of courses for both majors and
ulties of Northwestern University and Florida State University. nonmajors and currently teaches genetics, biotechnol-
He received his B.A. from St. Mary’s College in Winona, Min- ogy, endocrinology, and laboratory in cell and molecular
nesota, and his M.S. and Ph.D. from Northwestern University biology. He has received several awards for research and
in Evanston, Illinois. In addition to this text and its companion teaching, including the 2009 Young Investigator Award of
volumes, he has also written textbooks in human genetics and the American Society of Andrology, the 2005 Distinguished
general biology for nonmajors. His research interests center on Teacher Award from Monmouth University, and the 2005
the molecular organization and physical mapping of the hetero- Caring Heart Award from the New Jersey Association for
chromatic regions of human acrocentric chromosomes. At the Biomedical Research. He is co-author of the undergradu-
undergraduate level, he teaches courses in Mendelian and mo- ate textbook Introduction to Biotechnology, Series Editor for
lecular genetics, human genetics, and general biology, and has the Benjamin Cummings Special Topics in Biology booklet
received numerous awards for teaching excellence given by uni- series, and author of the first booklet in the series, Under-
versity faculty, student organizations, and graduating seniors. standing the Human Genome Project.
3
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Contents
3.7 Laws of Probability Help to Explain Genetic Events 58
1 Introduction to Genetics 17 3.8 Chi-Square Analysis Evaluates the Influence of Chance
on Genetic Data 59
1.1 Genetics Has a Rich and Interesting History 18
3.9 Pedigrees Reveal Patterns of Inheritance of Human
1.2 Genetics Progressed from Mendel to DNA in Less Than a Traits 62
Century 19
3.10 Tay–Sachs Disease: The Molecular Basis of a Recessive
1.3 Discovery of the Double Helix Launched the Era of Disorder in Humans 64
Molecular Genetics 21
EXPLORING GENOMICS
1.4 Development of Recombinant DNA Technology Began
Online Mendelian Inheritance in Man 64
the Era of DNA Cloning 23
CASE STUDY:To test or not to test 65
1.5 The Impact of Biotechnology Is Continually Expanding 23
Insights and Solutions 65
1.6 Genomics, Proteomics, and Bioinformatics Are New and
Expanding Fields 24 Problems and Discussion Questions 67
1.7 Genetic Studies Rely on the Use of Model Organisms 25
1.8 We Live in the Age of Genetics 26 4 Modification of Mendelian Ratios 69
Problems and Discussion Questions 27
4.1 Alleles Alter Phenotypes in Different Ways 70
4.2 Geneticists Use a Variety of Symbols for Alleles 70
2 Mitosis and Meiosis 28 4.3 Neither Allele Is Dominant in Incomplete, or Partial,
Dominance 71
2.1 Cell Structure Is Closely Tied to Genetic Function 29
4.4 In Codominance, the Influence of Both Alleles in a
2.2 Chromosomes Exist in Homologous Pairs in Diploid Heterozygote Is Clearly Evident 72
Organisms 31
4.5 Multiple Alleles of a Gene May Exist in a Population 72
2.3 Mitosis Partitions Chromosomes into Dividing Cells 33
4.6 Lethal Alleles Represent Essential Genes 74
2.4 Meiosis Creates Haploid Gametes and Spores and
Evolving Concept of the Gene 74
Enhances Genetic Variation in Species 37
4.7 Combinations of Two Gene Pairs with Two Modes of
2.5 The Development of Gametes Varies in
Inheritance Modify the 9:3:3:1 Ratio 75
Spermatogenesis Compared to Oogenesis 40
4.8 Phenotypes Are Often Affected by More Than One Gene 76
2.6 Meiosis Is Critical to Sexual Reproduction
4.9 Complementation Analysis Can Determine If Two
in All Diploid Organisms 42
Mutations Causing a Similar Phenotype Are Alleles
2.7 Electron Microscopy Has Revealed the Physical Structure
of the Same Gene 80
of Mitotic and Meiotic Chromosomes 42
4.10 Expression of a Single Gene May Have Multiple Effects 82
EXPLORING GENOMICS 4.11 X-Linkage Describes Genes on the X Chromosome 82
PubMed: Exploring and Retrieving Biomedical Literature 43
4.12 In Sex-Limited and Sex-Influenced Inheritance, an
CASE STUDY:Triggering meiotic maturation of oocytes 44 Individual’s Sex Influences the Phenotype 84
Insights and Solutions 44 4.13 Genetic Background and the Environment Affect
Problems and Discussion Questions 45 Phenotypic Expression 86
4.14 Genomic (Parental) Imprinting and Gene Silencing 88
4.15 Extranuclear Inheritance Modifies Mendelian Patterns 89
3 Mendelian Genetics 47
GENETICS, TECHNOLOGY, AND SOCIETY
3.1 Mendel Used a Model Experimental Approach to Study Improving the Genetic Fate of Purebred Dogs 92
Patterns of Inheritance 48 CASE STUDY: Sudden blindness 93
3.2 The Monohybrid Cross Reveals How One Trait Is Insights and Solutions 94
Transmitted from Generation to Generation 48
Problems and Discussion Questions 95
3.3 Mendel’s Dihybrid Cross Generated a Unique F Ratio 52
2
3.4 The Trihybrid Cross Demonstrates That Mendel’s
5 Sex Determination and Sex
Principles Apply to Inheritance of Multiple Traits 55
3.5 Mendel’s Work Was Rediscovered in the Early Twentieth Chromosomes
100
Century 57
Evolving Concept of the Gene 58 5.1 X and Y Chromosomes Were First Linked to Sex
3.6 Independent Assortment Leads to Extensive Genetic Determination Early in the Twentieth Century 101
Variation 58 5.2 The Y Chromosome Determines Maleness in Humans 102
5
6 CONTENTS
5.3 The Ratio of Males to Females in Humans 7.6 Other Aspects of Genetic Exchange 153
Is Not 1.0 105
EXPLORING GENOMICS
5.4 Dosage Compensation Prevents Excessive Expression Human Chromosome Maps on the Internet 155
of X-Linked Genes in Humans and Other
CASE STUDY: Links to autism 155
Mammals 106
Insights and Solutions 165
5.5 The Ratio of X Chromosomes to Sets of Autosomes Can
Determine Sex 109 Problems and Discussion Questions 156
5.6 Temperature Variation Controls Sex Determination in
Reptiles 111
CASE STUDY: Not reaching puberty 112 8 Genetic Analysis and Mapping
Insights and Solutions 113
in Bacteria and Bacteriophages
159
Problems and Discussion Questions 113
8.1 Bacteria Mutate Spontaneously and Are Easily
Cultured 160
8.2 Genetic Recombination Occurs in Bacteria 160
6 Chromosome Mutations: Variation
8.3 Rec Proteins Are Essential to Bacterial
in Number and Arrangement 115 Recombination 166
8.4 The F Factor Is an Example of a Plasmid 167
6.1 Variation in Chromosome Number: Terminology and
8.5 Transformation Is Another Process Leading to Genetic
Origin 116
Recombination in Bacteria 168
6.2 Monosomy and Trisomy Result in a Variety of
8.6 Bacteriophages Are Bacterial Viruses 169
Phenotypic Effects 117
8.7 Transduction Is Virus-Mediated Bacterial DNA
6.3 Polyploidy, in Which More Than Two Haploid
Transfer 172
Sets of Chromosomes Are Present, Is Prevalent
in Plants 121 CASE STUDY: To treat or not to treat 174
6.4 Variation Occurs in the Composition and Arrangement Insights and Solutions 174
of Chromosomes 123 Problems and Discussion Questions 174
6.5 A Deletion Is a Missing Region of a
Chromosome 124
6.6 A Duplication Is a Repeated Segment
of a Chromosome 126 9 DNA Structure and Analysis 176
6.7 Inversions Rearrange the Linear Gene Sequence 128
6.8 Translocations Alter the Location of Chromosomal 9.1 The Genetic Material Must Exhibit Four
Segments in the Genome 129 Characteristics 177
6.9 Fragile Sites in Human Chromosomes Are Susceptible 9.2 Until 1944, Observations Favored Protein as the Genetic
to Breakage 131 Material 177
9.3 Evidence Favoring DNA as the Genetic Material Was
CASE STUDY: Changing the face of Down syndrome 133
First Obtained during the Study of Bacteria and
Insights and Solutions 133
Bacteriophages 178
Problems and Discussion Questions 134 9.4 Indirect and Direct Evidence Supports the Concept that
DNA Is the Genetic Material in Eukaryotes 183
9.5 RNA Serves as the Genetic Material in
Some Viruses 184
7 Linkage and Chromosome Mapping
9.6 The Structure of DNA Holds the Key to Understanding
in Eukaryotes 136 Its Function 184
Evolving Concept of the Gene 190
7.1 Genes Linked on the Same Chromosome Segregate 9.7 Alternative Forms of DNA Exist 190
Together 137
9.8 The Structure of RNA Is Chemically Similar to DNA, but
7.2 Crossing Over Serves as the Basis of Determining the Single-Stranded 190
Distance between Genes during Mapping 140
9.9 Many Analytical Techniques Have Been Useful during the
7.3 Determining the Gene Sequence during Mapping Investigation of DNA and RNA 191
Requires the Analysis of Multiple Crossovers 143
EXPLORING GENOMICS
7.4 As the Distance between Two Genes Increases, Mapping
Introduction to Bioinformatics: BLAST 193
Estimates Become More Inaccurate 149
CASE STUDY: Zigs and zags of the smallpox virus 194
Evolving Concept of the Gene 152
7.5 Chromosome Mapping Is Now Possible Using DNA Insights and Solutions 194
Markers and Annotated Computer Databases 152 Problems and Discussion Questions 194
CONTENTS 7
12.4 The Coding Dictionary Reveals the Function of the 64
10 DNA Replication and Triplets 238
Recombination 12.5 The Genetic Code Has Been Confirmed in Studies of
196
Bacteriophage MS2 239
10.1 DNA Is Reproduced by Semiconservative 12.6 The Genetic Code Is Nearly Universal 239
Replication 197 12.7 Different Initiation Points Create Overlapping
10.2 DNA Synthesis in Bacteria Involves Five Polymerases, as Genes 240
Well as Other Enzymes 201 12.8 Transcription Synthesizes RNA on a DNA
10.3 Many Complex Issues Must Be Resolved during DNA Template 241
Replication 204 12.9 RNA Polymerase Directs RNA Synthesis 241
10.4 A Coherent Model Summarizes DNA Replication 207 12.10 Transcription in Eukaryotes Differs from Prokaryotic
10.5 Replication Is Controlled by a Variety of Genes 208 Transcription in Several Ways 243
10.6 Eukaryotic DNA Replication Is Similar to Replication in 12.11 The Coding Regions of Eukaryotic Genes Are Interrupted
Prokaryotes, but Is More Complex 208 by Intervening Sequences Called Introns 246
10.7 The Ends of Linear Chromosomes Are Problematic Evolving Concept of the Gene 249
during Replication 210 12.12 RNA Editing May Modify the Final Transcript 249
GENETICS, TECHNOLOGY, AND SOCIETY GENETICS, TECHNOLOGY, AND SOCIETY
Telomeres: The Key to Immortality? 212 Fighting Disease with Antisense Therapeutics 250
CASE STUDY: Premature aging and DNA helicases 213 CASE STUDY: Cystic fibrosis 251
Insights and Solutions 213 Insights and Solutions 251
Problems and Discussion Questions 214 Problems and Discussion Questions 252
11 Chromosome Structure and DNA 13 Translation and Proteins 254
Sequence Organization
215
13.1 Translation of mRNA Depends on Ribosomes and
Transfer RNAs 255
11.1 Viral and Bacterial Chromosomes Are Relatively Simple
DNA Molecules 216 13.2 Translation of mRNA Can Be Divided into Three Steps
258
11.2 Mitochondria and Chloroplasts Contain DNA Similar to
Bacteria and Viruses 217 13.3 High-Resolution Studies Have Revealed Many Details
about the Functional Prokaryotic Ribosome 262
11.3 Specialized Chromosomes Reveal Variations in the
Organization of DNA 219 13.4 Translation Is More Complex in Eukaryotes 263
11.4 DNA Is Organized into Chromatin in Eukaryotes 221 13.5 The Initial Insight That Proteins Are Important in
Heredity Was Provided by the Study of Inborn Errors of
11.5 Eukaryotic Genomes Demonstrate Complex Sequence
Metabolism 263
Organization Characterized by Repetitive DNA 225
13.6 Studies of Neurospora Led to the One-Gene:One-Enzyme
11.6 The Vast Majority of a Eukaryotic Genome Does Not
Hypothesis 264
Encode Functional Genes 228
13.7 Studies of Human Hemoglobin Established That One
EXPLORING GENOMICS
Gene Encodes One Polypeptide 266
Database of Genomic Variants: Structural Variations in the Human
Evolving Concept of the Gene 267
Genome 228
13.8 Variation in Protein Structure Is the Basis of Biological
CASE STUDY: Art inspires learning 229
Diversity 267
Insights and Solutions 229
13.9 Proteins Function in Many Diverse Roles 270
Problems and Discussion Questions 230
CASE STUDY: Crippled ribosomes 271
Insights and Solutions 271
Problems and Discussion Questions 271
12 The Genetic Code and
Transcription
231
14 Gene Mutation, DNA Repair,
12.1 The Genetic Code Exhibits a Number of
Characteristics 232 and Transposition 273
12.2 Early Studies Established the Basic Operational Patterns
of the Code 232 14.1 Gene Mutations Are Classified in Various Ways 274
12.3 Studies by Nirenberg, Matthaei, and Others Deciphered 14.2 Spontaneous Mutations Arise from Replication Errors
the Code 233 and Base Modifications 277
8 CONTENTS
14.3 Induced Mutations Arise from DNA Damage Caused by 16.3 Cancer Cells Contain Genetic Defects Affecting Cell-
Chemicals and Radiation 279 Cycle Regulation 328
14.4 Single-Gene Mutations Cause a Wide Range of Human 16.4 Proto-oncogenes and Tumor-Suppressor Genes Are
Diseases 281 Altered in Cancer Cells 330
14.5 Organisms Use DNA Repair Systems to Detect and 16.5 Cancer Cells Metastasize and Invade Other Tissues 332
Correct Mutations 282 16.6 Predisposition to Some Cancers Can Be Inherited 332
14.6 The Ames Test Is Used to Assess the Mutagenicity of 16.7 Viruses and Environmental Agents Contribute
Compounds 303 to Human Cancers 333
14.7 Transposable Elements Move within the Genome and
GENETICS, TECHNOLOGY, AND SOCIETY
May Create Mutations 288
Breast Cancer: The Double-Edged Sword of Genetic Testing 334
CASE STUDY: Genetic dwarfism 292 CASE STUDY: Screening for cancer can save lives 335
Insights and Solutions 293 Insights and Solutions 335
Problems and Discussion Questions 293 Problems and Discussion Questions 336
15 Regulation of Gene Expression 296 17 Recombinant DNA Technology 338
15.1 Prokaryotes Regulate Gene Expression in Response to
17.1 Recombinant DNA Technology Began with
Both External and Internal Conditions 297
Two Key Tools: Restriction Enzymes and DNA Cloning
15.2 Lactose Metabolism in E. coli Is Regulated by an
Vectors 339
Inducible System 297
17.2 DNA Libraries Are Collections of Cloned Sequences 344
15.3 The Catabolite-Activating Protein (CAP) Exerts Positive
17.3 The Polymerase Chain Reaction Is a Powerful Technique
Control over the lac Operon 302
for Copying DNA 347
15.4 The Tryptophan (trp) Operon in E. coli Is a Repressible
17.4 Molecular Techniques for Analyzing DNA 349
Gene System 304
17.5 DNA Sequencing Is the Ultimate Way to Characterize
Evolving Concept of the Gene 304
DNA at the Molecular Level 352
15.5 Alterations to RNA Secondary Structure Also Contribute
17.6 Creating Knockout and Transgenic Organisms for
to Prokaryotic Gene Regulation 304
Studying Gene Function 354
15.6 Eukaryotic Gene Regulation Differs from That in
Prokaryotes 307 EXPLORING GENOMICS
15.7 Eukaryotic Gene Expression Is Influenced by Chromatin Manipulating Recombinant DNA: Restriction Mapping and Designing
PCR Primers 358
Modifications 308
15.8 Eukaryotic Transcription Regulation Requires Specific CASE STUDY: Should we worry about recombinant DNA
technology? 359
Cis-Acting Sites 310
15.9 Eukaryotic Transcription Initiation is Regulated by Insights and Solutions 359
Transcription Factors That Bind to Cis-Acting Sites 312 Problems and Discussion Questions 360
15.10 Activators and Repressors Interact with General
Transcription Factors and Affect Chromatin
Structure 313
18 Genomics, Bioinformatics, and
15.11 Posttranscriptional Gene Regulation Occurs at Many
Steps from RNA Processing to Protein Modification 315 Proteomics 361
15.12 RNA-Induced Gene Silencing Controls Gene Expression
in Several Ways 317 18.1 Whole-Genome Shotgun Sequencing Is a Widely
Used Method for Sequencing and Assembling Entire
GENETICS, TECHNOLOGY, AND SOCIETY Genomes 362
Quorum Sensing: Social Networking in the Bacterial World 318
18.2 DNA Sequence Analysis Relies on Bioinformatics
CASE STUDY: A mysterious muscular dystrophy 319 Applications and Genome Databases 364
Insights and Solutions 319 18.3 Genomics Attempts to Identify Potential Functions of
Problems and Discussion Questions 320 Genes and Other Elements in a Genome 366
18.4 The Human Genome Project Revealed Many Important
Aspects of Genome Organization in Humans 367
16 The Genetics of Cancer 323 18.5 After the Human Genome Project: What Is Next? 370
Evolving Concept of the Gene 374
16.1 Cancer Is a Genetic Disease at the Level 18.6 Comparative Genomics Analyzes and Compares
of Somatic Cells 324 Genomes from Different Organisms 376
16.2 Cancer Cells Contain Genetic Defects Affecting Genomic 18.7 Comparative Genomics Is Useful for Studying the
Stability, DNA Repair, and Chromatin Modifications 327 Evolution and Function of Multigene Families 381