Table of Contents
| 1 | An Introduction to Genetics | 1 |
| Pt. 1 | Heredity and the Phenotype | 17 |
| 2 | Cell Division and Chromosomes | 18 |
| 3 | Mendelian Genetics | 50 |
| 4 | Modification of Mendelian Ratios | 79 |
| 5 | Linkage, Crossing Over, and Chromosome Mapping | 115 |
| 6 | Recombination and Mapping in Bacteria and Bacteriophages | 149 |
| 7 | Extensions of Genetic Analysis | 180 |
| 8 | Extranuclear Inheritance | 208 |
| 9 | Chromosome Variation and Sex Determination | 221 |
| Pt. 2 | Molecular Basis of Heredity | 261 |
| 10 | Structure and Analysis of DNA and RNA | 262 |
| 11 | DNA Replication and Recombination | 298 |
| 12 | Storage and Expression of Genetic Information | 324 |
| 13 | Proteins: The End Product of Genetic Expression | 364 |
| 14 | Gene Mutation, DNA Repair, and Transposable Elements | 389 |
| 15 | Recombinant DNA Technology | 428 |
| 16 | Applications of Recombinant DNA Technology | 461 |
| 17 | Genomic Organization of DNA | 490 |
| Pt. 3 | Advanced Topics in Genetic Analysis | 521 |
| 18 | Regulation of Gene Expression in Bacteria and Phages | 522 |
| 19 | Regulation of Gene Expression in Eukaryotes | 544 |
| 20 | Developmental Genetics | 567 |
| 21 | Genetics and Cancer | 591 |
| 22 | Genetic Basis of the Immune Response | 614 |
| 23 | The Genetics of Behavior | 637 |
| 24 | Population Genetics | 659 |
| 25 | Genetics and Evolution | 681 |
| Appendix A: Experimental Methods | |
| Appendix B: Glossary | |
| Appendix C: Solutions to Selected Even-Numbered Problems and Discussion Questions | |
| Credits | |
| Index | |
Forewords & Introductions
PREFACE:
Preface
Concepts of Genetics is now completing its second decade of providing support to students as they study one of the most fascinating scientific disciplines. Certainly no subject area has had a more sustained impact on shaping our knowledge of the living condition. As a result of discoveries over the past 50 years, we now understand with reasonable clarity the underlying genetic mechanisms that explain how organisms develop into and then function as adults. We also better understand the basis of biological diversity and have greater insight into the evolutionary process.
Advances in genetic technology are now having a profound impact on our knowledge of human genetics. More than any other event, the launching of the Human Genome Project in 1990 symbolizes our commitment to the pursuit of such knowledge. As we enter the twentyfirst century, the application of genetics to the betterment of the human condition will become commonplace. While this era will be filled with the excitement of scientific discovery, many accompanying problems and controversies will also face us. Of these, how we utilize our knowledge of the nucleotide sequence of the human genome promises to be the most significant. This growing body of information has already generated many legal and ethical issues. Currently, the implications of genetic testing and gene therapy are becoming important societal concerns.
As geneticists and students of genetics, the thrill of being part of this era must be balanced by a strong sense of responsibility and careful attention to the many related issues that will undoubtedly arise. The formulation of proper laws and policies will dependona comprehensive knowledge of genetics and measured responses to these issues. As a result, here has never been a higher premium or greater need for a useful and up-to-date genetics textbook.
The sixth edition of Concepts of Genetics, as with all past efforts, has been designed to achieve five major goals:
- To establish a conceptual framework that represents a sound approach to learning and facilitates the comprehension of the vast amount of information constituting the field of genetics.
- To emphasize the rich history of scientific discovery and analytical thought, so prevalent in genetics, that provides an opportunity for students to hone their problemsolving abilities and to explore how we know what we know.
- To cover material with a clear, crisp organizational format, both within each chapter and throughout the chapter sequence, to facilitate effective use of the text.
- To present carefully designed figures with strong pedagogic value that teach rather than merely illustrate complex topics and analytical experiments.
- To provide students with clearly written, straightforward explanations that elucidate difficult, complex topics without oversimplifying presentations.
Writing a textbook that achieves these goals and having the opportunity to continually improve on each effort has been a labor of love for us. The creation of each of the six editions is a reflection not only of our passion for teaching genetics, but the constructive feedback and encouragement provided over the past two decades from adopters, reviewers, and our students.
FEATURES OF THIS EDITION
New Conceptual Headings
The organization of each chapter has been revised to include conceptual headings that initiate the discussion of all major topics. These are presented using the narrative style. Collectively, these are presented at the onset of each chapter in a section entitled Key Concepts. This organizational feature reinforces our emphasis on conceptual issues in genetics. This is in keeping not only with the title of this textbook, but with our ardent belief that learning depends on the understanding of concepts, which are not easily forgotten, in contrast with countless details that may be memorized but not always retained. We believe that this new approach to the organization of each chapter will pay substantial dividends in improving the pedagogic value of Concepts of Genetics.
New Organization of the Table of Contents
As the field of genetics has expanded, many topics have taken on greater importance and stature within the discipline. Once again, as with previous editions, this has necessitated substantial reorganization, which is apparent in the Table of Contents. This edition continues to reflect our belief that coverage of the more classical studies of genetics, which we refer to as Heredity and the Phenotype, should initiate the text as Part I At the urging of many reviewers, we have included coverage of bacterial and phage genetics, as well as extranuclear inheritance in this section. Additionally, this initial part of the text now includes a new chapter, Sex Determination and Sex Chromosomes (Chapter 9). Part I then concludes with the chapter on Chromosome Mutations.
Part 11, entitled The Molecular Basis of Inheritance, includes a group of chapters that center around the storage, expression, and regulation of genetic information. These topics collectively form what is classically thought of as molecular genetics. While we have decided to keep many DNA-related topics in Part 11, those who wish to begin their courses at this point in the text should not have difficulty doing so. Because the chapters have been written to be independent of the information presented in Part 1, many adopters have used this approach successfully in previous editions.
With the strong foundation of DNA structure and function in hand, Part III includes four chapters that address the newest area of study in genetics, entitled Genomics. This section opens with coverage of how DNA clones are constructed and analyzed, and follows with chapters that address how DNA sequences and genes are organized within chromosomes. Part III concludes with the consideration of the applications and ethics of genetic technology. Obviously, Part III represents the "cutting edge" of genetics and contains the most exciting, far-reaching findings in this field.
We conclude the text by returning to the consideration of organisms and populations in Part IV. Initial chapters in this section address the role of genetic information in development, cancer, and behavior. We conclude our coverage with a consideration of the genetics of populations and the role of genetics in the process of evolution.
While we realize that no Table of Contents in a diverse field such as genetics can satisfy everyone, the present organization can be used in all first courses in undergraduate genetics, whether offered in a semester, quarter, or two-quarter format. The Parts and Chapters within the text are written so as to be used interchangeably, providing flexibility for the instructor.
Revised Art Program
In this edition, we have substantially refined the entire art program. Every figure has been scrutinized, and most have been either redrawn or revised. A number of the most complex figures have been more carefully and closely linked to their accompanying discussion. In conjunction with many new striking photographs, the text provides an attractive and pedagogically superior illustration program to enhance and support discussions of every major topic.
As in past editions, our goal has been to create figures that facilitate learning. In many cases, "flowchart" figures have been developed to illustrate complex experimental approaches or genetic processes that are significant to our understanding of genetics.
Revisions and Modernization
The entire text has been revised with an eye to providing clarity and coverage of the latest cutting-edge topics in genetics. One of the most significant paradigms in genetic investigation involves the utilization of DNA cloning technology to study many areas of biology. In this edition, this trend is reflected particularly in the revision and modernization of the information presented in Part III—Genomics. For example, Chapter 20—Organization of Genes in Chromosomes, is a new chapter that includes the most recent findings involving our understanding of genomic organization in a wide range of organisms.
While many areas of genetics have been enhanced by the ability to clone and analyze DNA, human genetics has benefited more than any other field. The expansion of knowledge about our own species represents a second major trend in genetics. As a result, we have increased our coverage of human genetics throughout the text. This is particularly evident in two chapters, Applications and Ethics of Genetic Technology (Chapter 2 1) and Genetics and Cancer (Chapter 2 3), both of which heavily emphasize human genetics. These chapters convey some of the vast amount of information provided by the Human Genome Project. Two other chapters important to the understanding of genetics have been revised substantially: Population Genetics (Chapter 25) and Genetics and Evolution (Chapter 26). The revisions reflect recent progress in these fields, as well as the influence of molecular genetics on these disciplines. This effort is in keeping with the important role population and evolutionary biology plays in the study of biology.
New Essays
In keeping with the influence of genetics on our everyday lives, we have revised and expanded the essays entitled Genetics, Technology, and Society, which appear near the end of most chapters. A number of these are new to this edition, including "Genetics and Society at the Millenium" in Chapter 1 and "Gene Pools and Endangered Species" in Chapter 25. These essays, which contain accompanying references, can readily serve as the basis of class discussions in genetics courses.
New Media Resources
Concepts of Genetics takes a significant step forward with the integration of interactive media resources into the text. This is done through the GenCDX Student CDROM, the textspecific companion website (http-J/www.pr-enhaU.com/klug), and the "Genetics MediaLab" sections at the end of most of the chapters. The integration of these resources provides both instructors and students with new learning tools to help explain difficult concepts, and to explore the applications and implications of genetics in a broader context.
The GenCDX Student CD-ROM, designed specifically for this edition, contains over two dozen animations and tutorials, and is the starting point for the web problems in the Genetics MediaLabs. These animations and tutorials are taken directly from the text's art program and cover a wide range of topics, including Mendelian inheritance, gene mutation, transcription, translation, gene cloning and population genetics. The tutorials consist of an introduction, followed by an animation with accompanying text, self-test questions and a conclusion. Each is followed by a set of related questions with hints and explanations accompanying each answer. Icons placed in selected text figures link the tutorials to concepts in the book. The GenCDX Student CD-ROM also contains a set of selfgrading chapter problems, "Genetics, Technology and Society" exercises, Chapter Search Terms, a Student Bulletin Board and other resources.
The Companion Website for the text provides access to the websites for the MediaLab investigations, linked web destinations in genetics, links to genetics newsgroups, web navigation tips, additional self-grading problems, and other resources.
The "Genetics MediaLab" section at the end of each chapter contains a list of chapter-specific resources available on the GenCDX Student CD-ROM and the companion website. This section also contains several interactive web-linked problems designed to enhance the topics presented in the chapter. To complete these problems, students must actively participate in exercises such as virtual experiments in Drosophila genetics, bacterial conjugation and the nature of human genetic disorders. Each of these problems has been selected and screened for pedagogical value by genetics instructors. For reference, the estimated time required to solve the problem is noted at the beginning of the exercise.
Coupled to the book, these media resources provide a powerful tool for teaching and learning genetics. It is our hope that both students and instructors will explore these resources and make them an integral part of a genetics course.
Read an Excerpt
PREFACE:
Preface
Concepts of Genetics is now completing its second decade of providing support to students as they study one of the most fascinating scientific disciplines. Certainly no subject area has had a more sustained impact on shaping our knowledge of the living condition. As a result of discoveries over the past 50 years, we now understand with reasonable clarity the underlying genetic mechanisms that explain how organisms develop into and then function as adults. We also better understand the basis of biological diversity and have greater insight into the evolutionary process.
Advances in genetic technology are now having a profound impact on our knowledge of human genetics. More than any other event, the launching of the Human Genome Project in 1990 symbolizes our commitment to the pursuit of such knowledge. As we enter the twentyfirst century, the application of genetics to the betterment of the human condition will become commonplace. While this era will be filled with the excitement of scientific discovery, many accompanying problems and controversies will also face us. Of these, how we utilize our knowledge of the nucleotide sequence of the human genome promises to be the most significant. This growing body of information has already generated many legal and ethical issues. Currently, the implications of genetic testing and gene therapy are becoming important societal concerns.
As geneticists and students of genetics, the thrill of being part of this era must be balanced by a strong sense of responsibility and careful attention to the many related issues that will undoubtedly arise. The formulation of proper laws and policies will depend ona comprehensive knowledge of genetics and measured responses to these issues. As a result, here has never been a higher premium or greater need for a useful and up-to-date genetics textbook.
The sixth edition of Concepts of Genetics, as with all past efforts, has been designed to achieve five major goals:
- To establish a conceptual framework that represents a sound approach to learning and facilitates the comprehension of the vast amount of information constituting the field of genetics.
- To emphasize the rich history of scientific discovery and analytical thought, so prevalent in genetics, that provides an opportunity for students to hone their problemsolving abilities and to explore how we know what we know.
- To cover material with a clear, crisp organizational format, both within each chapter and throughout the chapter sequence, to facilitate effective use of the text.
- To present carefully designed figures with strong pedagogic value that teach rather than merely illustrate complex topics and analytical experiments.
- To provide students with clearly written, straightforward explanations that elucidate difficult, complex topics without oversimplifying presentations.
Writing a textbook that achieves these goals and having the opportunity to continually improve on each effort has been a labor of love for us. The creation of each of the six editions is a reflection not only of our passion for teaching genetics, but the constructive feedback and encouragement provided over the past two decades from adopters, reviewers, and our students.
FEATURES OF THIS EDITION
New Conceptual Headings
The organization of each chapter has been revised to include conceptual headings that initiate the discussion of all major topics. These are presented using the narrative style. Collectively, these are presented at the onset of each chapter in a section entitled Key Concepts. This organizational feature reinforces our emphasis on conceptual issues in genetics. This is in keeping not only with the title of this textbook, but with our ardent belief that learning depends on the understanding of concepts, which are not easily forgotten, in contrast with countless details that may be memorized but not always retained. We believe that this new approach to the organization of each chapter will pay substantial dividends in improving the pedagogic value of Concepts of Genetics.
New Organization of the Table of Contents
As the field of genetics has expanded, many topics have taken on greater importance and stature within the discipline. Once again, as with previous editions, this has necessitated substantial reorganization, which is apparent in the Table of Contents. This edition continues to reflect our belief that coverage of the more classical studies of genetics, which we refer to as Heredity and the Phenotype, should initiate the text as Part I At the urging of many reviewers, we have included coverage of bacterial and phage genetics, as well as extranuclear inheritance in this section. Additionally, this initial part of the text now includes a new chapter, Sex Determination and Sex Chromosomes (Chapter 9). Part I then concludes with the chapter on Chromosome Mutations.
Part 11, entitled The Molecular Basis of Inheritance, includes a group of chapters that center around the storage, expression, and regulation of genetic information. These topics collectively form what is classically thought of as molecular genetics. While we have decided to keep many DNA-related topics in Part 11, those who wish to begin their courses at this point in the text should not have difficulty doing so. Because the chapters have been written to be independent of the information presented in Part 1, many adopters have used this approach successfully in previous editions.
With the strong foundation of DNA structure and function in hand, Part III includes four chapters that address the newest area of study in genetics, entitled Genomics. This section opens with coverage of how DNA clones are constructed and analyzed, and follows with chapters that address how DNA sequences and genes are organized within chromosomes. Part III concludes with the consideration of the applications and ethics of genetic technology. Obviously, Part III represents the "cutting edge" of genetics and contains the most exciting, far-reaching findings in this field.
We conclude the text by returning to the consideration of organisms and populations in Part IV. Initial chapters in this section address the role of genetic information in development, cancer, and behavior. We conclude our coverage with a consideration of the genetics of populations and the role of genetics in the process of evolution.
While we realize that no Table of Contents in a diverse field such as genetics can satisfy everyone, the present organization can be used in all first courses in undergraduate genetics, whether offered in a semester, quarter, or two-quarter format. The Parts and Chapters within the text are written so as to be used interchangeably, providing flexibility for the instructor.
Revised Art Program
In this edition, we have substantially refined the entire art program. Every figure has been scrutinized, and most have been either redrawn or revised. A number of the most complex figures have been more carefully and closely linked to their accompanying discussion. In conjunction with many new striking photographs, the text provides an attractive and pedagogically superior illustration program to enhance and support discussions of every major topic.
As in past editions, our goal has been to create figures that facilitate learning. In many cases, "flowchart" figures have been developed to illustrate complex experimental approaches or genetic processes that are significant to our understanding of genetics.
Revisions and Modernization
The entire text has been revised with an eye to providing clarity and coverage of the latest cutting-edge topics in genetics. One of the most significant paradigms in genetic investigation involves the utilization of DNA cloning technology to study many areas of biology. In this edition, this trend is reflected particularly in the revision and modernization of the information presented in Part III—Genomics. For example, Chapter 20—Organization of Genes in Chromosomes, is a new chapter that includes the most recent findings involving our understanding of genomic organization in a wide range of organisms.
While many areas of genetics have been enhanced by the ability to clone and analyze DNA, human genetics has benefited more than any other field. The expansion of knowledge about our own species represents a second major trend in genetics. As a result, we have increased our coverage of human genetics throughout the text. This is particularly evident in two chapters, Applications and Ethics of Genetic Technology (Chapter 2 1) and Genetics and Cancer (Chapter 2 3), both of which heavily emphasize human genetics. These chapters convey some of the vast amount of information provided by the Human Genome Project. Two other chapters important to the understanding of genetics have been revised substantially: Population Genetics (Chapter 25) and Genetics and Evolution (Chapter 26). The revisions reflect recent progress in these fields, as well as the influence of molecular genetics on these disciplines. This effort is in keeping with the important role population and evolutionary biology plays in the study of biology.
New Essays
In keeping with the influence of genetics on our everyday lives, we have revised and expanded the essays entitled Genetics, Technology, and Society, which appear near the end of most chapters. A number of these are new to this edition, including "Genetics and Society at the Millenium" in Chapter 1 and "Gene Pools and Endangered Species" in Chapter 25. These essays, which contain accompanying references, can readily serve as the basis of class discussions in genetics courses.
New Media Resources
Concepts of Genetics takes a significant step forward with the integration of interactive media resources into the text. This is done through the GenCDX Student CDROM, the textspecific companion website (http-J/www.pr-enhaU.com/klug), and the "Genetics MediaLab" sections at the end of most of the chapters. The integration of these resources provides both instructors and students with new learning tools to help explain difficult concepts, and to explore the applications and implications of genetics in a broader context.
The GenCDX Student CD-ROM, designed specifically for this edition, contains over two dozen animations and tutorials, and is the starting point for the web problems in the Genetics MediaLabs. These animations and tutorials are taken directly from the text's art program and cover a wide range of topics, including Mendelian inheritance, gene mutation, transcription, translation, gene cloning and population genetics. The tutorials consist of an introduction, followed by an animation with accompanying text, self-test questions and a conclusion. Each is followed by a set of related questions with hints and explanations accompanying each answer. Icons placed in selected text figures link the tutorials to concepts in the book. The GenCDX Student CD-ROM also contains a set of selfgrading chapter problems, "Genetics, Technology and Society" exercises, Chapter Search Terms, a Student Bulletin Board and other resources.
The Companion Website for the text provides access to the websites for the MediaLab investigations, linked web destinations in genetics, links to genetics newsgroups, web navigation tips, additional self-grading problems, and other resources.
The "Genetics MediaLab" section at the end of each chapter contains a list of chapter-specific resources available on the GenCDX Student CD-ROM and the companion website. This section also contains several interactive web-linked problems designed to enhance the topics presented in the chapter. To complete these problems, students must actively participate in exercises such as virtual experiments in Drosophila genetics, bacterial conjugation and the nature of human genetic disorders. Each of these problems has been selected and screened for pedagogical value by genetics instructors. For reference, the estimated time required to solve the problem is noted at the beginning of the exercise.
Coupled to the book, these media resources provide a powerful tool for teaching and learning genetics. It is our hope that both students and instructors will explore these resources and make them an integral part of a genetics course.
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