Principles of Genetics 7th Edition Robert H Tamarin

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 file:///C|/Documents%20and%20Settings/Admin/Desktop/genni.txt

Table of Contents

Part I Genetics and the Scientific Method

1 Introduction

Part II Mendelism and the Chromosomal Theory

2 Mendel’s Principles
3 Mitosis and Meiosis
4 Probability and Statistics
5 Sex Determination, Sex Linkage, and Pedigree Analysis
6 Linkage and Mapping in Eukaryotes
7 Linkage and Mapping in Prokaryotes and Bacterial Viruses
8 Cytogenetics

Part III Molecular Genetics

9 Chemistry of the Gene

10 Gene Expression: Transcription
11 Gene Expression: Translation
12 DNA: Its Mutation, Repair and Recombination
13 Genomics, Biotechnology, and Recombinant DNA
14 Gene Expression: Control in Prokaryotes and Phages
15 The Eukaryotic Chromosome
16 Gene Expression: Control in Eukaryotes
17 Non-Mendelian Inheritance

Part IV Quantitative and Evolutionary Genetics

18 Quantitative Inheritance
19 Population Genetics: The Hardy-Weinberg Equilibrium and Mating Systems
20 Population Genetics: Processes That Change Allelic Frequencies
21 Evolution and Speciation

Appendixes

A. Brief Answers to Selected Exercises, Problems, and Critical Thinking Questions

B. Suggestions for Further Reading

file:///C|/Documents%20and%20Settings/Admin/Desktop/genni.txt [9/11/2008 5:06:57 PM]

Tamarin: Principles of Front Matter Preface © The McGraw−Hill
Genetics, Seventh Edition Companies, 2001

PREFACE

he twentieth century began with the redis- in their use. A comprehensive glossary and index will

T covery of Mendel’s rules of inheritance and

ended with the complete sequence of the hu-
man genome, one of the most monumental
scientific accomplishments of all time. What
lies in the future? What will the twenty-first century, the
century of genomics, bring? Will geneticists a hundred
years from now speak of a complete cure for cancer,
help maintain continuity if the instructor chooses to
change the order of the chapters from the original.
An understanding of genetics is crucial to advance-
ments in medicine, agriculture, and many industries. Ge-
netic controversies—such as the pros and cons of the
Human Genome Project, the potential ethical and med-
ical risks of recombinant DNA and cloning of mammals,
heart disease, and mental illness? Will we have a cure for and human behavioral genetic issues such as the degree
autoimmune diseases such as diabetes and arthritis? Will of inheritance of homosexuality, alcoholism, and intelli-
aging be slowed or even prevented? Will we have a com- gence—have captured the interest of the general public.
plete understanding of the process of development and a Throughout this text, we examine the implications for
concurrent elimination of birth defects and developmen- human health and welfare of the research conducted
tal problems? Will genetics put an end to world hunger? in universities and research laboratories around the
How will we live, and what will be the quality of our world; boxed material in the text gives insight into ge-
lives? The students who now are taking genetics will netic techniques, controversies, and breakthroughs.
learn the answers to these questions as time progresses. Because genetics is the first analytical biology course
Some students will contribute to the answers. for many students, some may have difficulty with its
The science of genetics includes the rules of inheri- quantitative aspects. There is no substitute for work with
tance in cells, individuals, and populations and the mo- pad and pencil. This text provides a larger number of
lecular mechanisms by which genes control the growth, problems to help the student learn and retain the mate-
development, and appearance of an organism. No area of rial. All problems within the body of the text and a selec-
biology can truly be appreciated or understood without tion at the end of the chapters should be worked through
an understanding of genetics because genes not only as they are encountered. After the student has worked
control cellular processes, they also determine the out the problems, he or she can refer to the answer sec-
course of evolution. Genetic concepts provide the frame- tion in Appendix A. We provide solved problems at the
work for the study of modern biology. end of each chapter to help.
This text provides a balanced treatment of the ma- In this text, we stress critical thinking, an approach
jor areas of genetics in order to prepare the student for that emphasizes understanding over memorization, ex-
upper-level courses and to help share in the excitement perimental proof over the pronouncements of authori-
of research. Most readers of this text will have taken a ties, problem solving over passive reading, and active
general biology course and will have had some back- participation in lectures. The latter is best accomplished
ground in cell biology and organic chemistry. For an un- if the student reads the appropriate text chapter before
derstanding of the concepts in this text, however, the coming to lecture rather than after. That way the student
motivated student will need to have completed only an can use the lecture to gain insight into difficult material
introductory biology course and have had some chem- rather than spending the lecture hectically transcribing
istry and algebra in high school. the lecturer’s comments onto the notebook page.
Genetics is commonly divided into three areas: classi- For those students who wish to pursue particular
cal, molecular, and population, although molecular ad- topics, a reference section in the back of the text pro-
vancements have blurred these distinctions. Many genetics vides chapter-by-chapter listings of review articles and ar-
teachers feel that a historical approach provides a sound ticles in the original literature. Although some of these
introduction to the field and that a thorough grounding articles might be difficult for the beginner to follow, each
in Mendelian genetics is necessary for an understanding is a landmark paper, a comprehensive summary, or a pa-
of molecular and population genetics—an approach this per with some valuable aspect. Some papers may contain
text follows. Other teachers, however, may prefer to be- an insightful photograph or diagram. Some magazines
gin with molecular genetics. For this reason, the chapters and journals are especially recommended for the student
have been grouped as units that allow for flexibility to look at periodically, including Scientific American,

xiii
 Tamarin: Principles of Front Matter Preface © The McGraw−Hill
Genetics, Seventh Edition Companies, 2001

xiv Preface

Science, and Nature, because they contain nontechnical • The material in chapter 3 on Genetic Control of the
summaries as well as material at the cutting edge of ge- Cell Cycle has been upgraded to a chapter section on
netics. Some articles are included to help the instructor the Cell Cycle.
find supplementary materials related to the concepts in • Molecular material throughout the book has been
this book. Photographs of selected geneticists also are in- completely updated to include such subjects as nu-
cluded. Perhaps the glimpse of a face from time to time merous DNA repair polymerases and their function-
will help add a human touch to this science. ing; base-flipping; TRAP control of attenuation; and
The World Wide Web also can provide a valuable re- chromatosomes.
source. The textbook has its own website: www.
mhhe.com/tamarin7. In addition, the student can find
much material of a supplemental nature by “surfing” the LEARNING AIDS FOR
web. Begin with a search engine such as: www. THE STUDENT
yahoo.com, or www.google.com and type in a key word.
Follow the links from there. Remember that the material To help the student learn genetics, as well as enjoy the
on the web is “as is”; it includes a lot of misinformation. material, we have made every effort to provide pedagog-
Usually, content from academic, industrial, and organiza- ical aids.These aids are designed to help organize the ma-
tional sources is relatively reliable; however, caveat emp- terial and make it understandable to students.
tor—buyer beware. Often in surfing for scientific key
words, the student will end up at a scientific journal or • Study Objectives Each chapter begins with a set of
clearly defined, page-referenced objectives. These ob-
book that does not have free access. Check with the uni-
jectives preview the chapter and highlight the most
versity librarian to see if access might be offered to that
important concepts.
journal or book. The amount of information that is accu-
• Study Outline The chapter topics are provided in
rate and free is enormous. Be sure to budget the amount
an outline list. These headings consist of words or
of time spent on the Internet.
phrases that clearly define what the various sections
of the chapter contain.
• Boldface Terms Throughout the chapter, all new
terms are presented in boldface, indicating that each
NEW TO THIS EDITION is defined in the glossary at the end of the book.
• Boxed Material In most chapters, short topics
Since the last edition of this text, many exciting discover- have been set aside in boxed readings, outside the
ies have been made in genetics. All chapters have been main body of the chapter. These boxes fall into four
updated to reflect those discoveries. In particular: categories: Historical Perspectives, Experimental
Methods, Biomedical Applications, and Ethics
• The chapter on Recombinant DNA Technology has and Genetics. The boxed material is designed to
been revised to be a chapter on Genomics, Biotech-
supplement each chapter with entertaining, interest-
nology, and Recombinant DNA (sixth edition chapter
ing, and relevant topics.
12 has become chapter 13 in this edition). The chap-
• Full Color Art and Graphics Many genetic con-
ter includes new material on the completion of the
cepts are made much clearer with full-color illustra-
Human Genome Project, bioinformatics, proteomics,
tions and the latest in molecular computer models to
and the latest techniques in creating cDNA and
help the student visualize and interpret difficult
knockout mice.
concepts. We’ve added thirty new photographs and
• The chapter on Control of Transcription in Eukary- over a hundred new and modified line drawings to
otes (sixth edition chapter 15 has become chapter
this edition.
16 in this edition) has been completely reorganized
• Summary Each chapter summary recaps the study
and rewritten to emphasize signal transduction, spe-
objectives at the beginning of the chapter. Thus, the
cific transcription factors, methylation, and chro-
student can determine if he or she has gained an un-
matin remodeling in control of gene expression; as in
derstanding of the material presented in the study ob-
the last edition, there are specific sections on
jectives and reinforce them with the summary.
Drosophila and plant development, cancer, and im-
• Solved Problems From two to four problems are
munogenetics.
worked out at the end of each chapter to give the stu-
• For better continuity, the chapter on Mutation, Re- dent practice in solving and understanding basic
combination, and DNA Repair has been moved to fol-
problems related to the material.
low the chapters on Transcription and Translation
• Exercises and Problems At the end of the chap-
(sixth edition chapter 16 has become chapter 12 in
ter are numerous problems to test the student’s
this edition).
Tamarin: Principles of Front Matter Preface © The McGraw−Hill
Genetics, Seventh Edition Companies, 2001

Preface xv

understanding of the material. These problems are lenging concepts in the course and makes them more
grouped according to the sections of the chapter. An- understandable through the presentation of full-
swers to the odd-numbered problems are presented color, narrated animations and interactive exercises.
in Appendix A, with the even-numbered problems an- The text indicates related topics on the CD with the
swered only in the Student Study Guide so that the following icon:
student and instructor can be certain that the student
is gaining an understanding of the material.
• Student Study Guide. This study guide features key
• Critical Thinking Questions Two critical think- concepts, problem-solving hints, practice problems,
ing questions at the end of each chapter are designed
terms, study questions, and answers to even-numbered
to help the student develop an ability to evaluate and
questions in the text. (ISBN 0072334207)
solve problems. The answer to the first critical think-
ing question can be found in Appendix A, and the an- • Laboratory Manual of Genetics 4/e, by A. M. Win-
chester and P. J. Wejksnora, University of Wisconsin–
swer to the second question is in the Student Study
Milwaukee. This manual for the genetics laboratory
Guide.
features classical and molecular biology exercises
that give students the opportunity to apply the scien-
tific method to “real”—not simulated—lab investiga-
tions. (ISBN 0697122875)
A N C I L L A R Y M AT E R I A L S • Case Workbook in Human Genetics, 2/e, by Ricki
Lewis, SUNY–Albany. The Workbook includes
For the Instructor thought-provoking case studies in human genetics,
• Website. Visit us at www.mhhe.com/tamarin7. with many examples gleaned from the author’s expe-
Here instructors will find jpeg files of the line draw- riences as a practicing genetic counselor. (ISBN
ings and tables suitable for downloading into Power- 0072325305) Also included is the Answer Key. (ISBN
Point, quizzes for study support, and links to genetic 0072439009)
sites. In addition, instructors will also find a link to
our hugely successful PageOut: The Course Web-
site Development Center, where instructors can
create a professional-looking, customized course AC K N OW L E D G M E N T S
website. It’s incredibly easy to use, and you need not
know html coding. I would like to thank many people for their encourage-
• Visual Resource Library (VRL). This Windows- and ment and assistance in the production of this Seventh
Macintosh-compatible CD-ROM has all the line draw- Edition. I especially thank Brian Loehr, my Developmen-
ings and tables from the text suitable for PowerPoint tal Editor, for continuous support, enthusiasm, and help
presentations. (ISBN 0072334266) in improving the usability of the text. It was also a plea-
• Instructor’s Manual with Test Item File. Available on sure to work with many other dedicated and creative
the website, the Instructor’s Manual contains out- people at McGraw-Hill during the production of this
lines, key words, summaries, instructional hints, and book, especially James M. Smith, Thomas Timp, Gloria
supplemental aids. The Test Item File contains 35 to Schiesl, David Hash, Sandy Ludovissy, Carrie Burger, and
50 objective questions with answers for each chap- Jodi Banowetz. I wish to thank Dr. Michael Gaines of the
ter. (ISBN 0072334215) University of Miami for many comments that helped me
• Test Item File on MicroTest III Classroom Testing improve the textbook and Marion Muskiewicz, Refer-
Software is an easy-to-use CD-ROM test generator also ence Librarian at the University of Massachusetts Lowell,
offered free upon request to adopters of this text.The who was an enormous help in my efforts to use the uni-
software requires no programming experience and is versity’s electronic library. Many reviewers greatly
compatible with Windows or Macintosh systems. helped improve the quality of this edition. I specifically
(ISBN 0072334231). wish to thank the following:

For the Student Reviewers of the Seventh Edition

• Website. Visit us at www.mhhe.com/tamarin7. John Belote
Here the student will find quizzes for study support,
Syracuse University
web exercises and resources, and links to genetic sites.
• Genetics: From Genes to Genomes CD-ROM, by Ann Douglas Coulter
E. Reynolds, University of Washington. Packaged free Saint Louis University
with every text, this CD-ROM covers the most chal-
Explore https://ebookultra.com
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 Tamarin: Principles of Front Matter Preface © The McGraw−Hill
Genetics, Seventh Edition Companies, 2001

xvi Preface

James M. Freed John R. Ellison

Ohio Wesleyan University Texas A&M University
Elliott S. Goldstein Elliott S. Goldstein
Arizona State University Arizona State University
Keith Hartberg Keith Hartberg
Baylor University Baylor University
Vincent Henrich David R. Hyde
University of North Carolina at Greensboro University of Notre Dame
Mitrick A. Johns Pauline A. Lizotte
Northern Illinois University Northwest Missouri State University
Philip Mathis James J. McGivern
Middle Tennessee State University Gannon University
Bruce McKee Gregory J. Phillips
University of Tennessee Iowa State University
Elbert Myles Mark Sanders
Tennessee State University University of California–Davis
John Osterman Ken Spitze
University of Nebraska–Lincoln University of Miami
Uwe Pott Joan M. Stoler
University of Wisconsin–Green Bay Massachusetts General Hospital, Harvard Medical
Ken Spitze School
University of Miami Robert J. Wiggers
Randall G. Terry Stephen F. Austin State University
University of Montana Ronald B. Young
Michael Wooten University of Alabama
Auburn University Lastly, thanks are due to the many students, particu-
larly those in my Introductory Genetics, Population Biol-
Reviewers of the Sixth Edition ogy, Evolutionary Biology, and Graduate Seminar courses,
Edward Berger who have helped clarify points, find errors, and discover
Dartmouth new and interesting ways of looking at the many topics
collectively called genetics.
Deborah C. Clark
Middle Tennessee State University ROBERT H. TAMARIN
Lowell, Massachusetts
Tamarin: Principles of I. Genetics and the 1. Introduction © The McGraw−Hill
Genetics, Seventh Edition Scientific Method Companies, 2001

1
INTRODUCTION STUDY OBJECTIVES
1. To examine a brief overview of the modern history
of genetics 3
2. To gain an overview of the topics included in this book—the
syllabus of genetics 4
3. To analyze the scientific method 5
4. To look at why certain organisms and techniques have been
used preferentially in genetics research 7

STUDY OUTLINE
A Brief Overview of the Modern History of Genetics 3
Before 1860 3
1860–1900 3
1900–1944 3
1944–Present 4
The Three General Areas of Genetics 4
How Do We Know? 5
Why Fruit Flies and Colon Bacteria? 7
Techniques of Study 8
Classical, Molecular, and Evolutionary Genetics 9
Classical Genetics 9
Molecular Genetics 10
Evolutionary Genetics 13
Summary 14
Box 1.1 The Lysenko Affair 6

Chameleon, Cameleo pardalis.

(© Art Wolfe/Tony Stone Images.)

2
Tamarin: Principles of I. Genetics and the 1. Introduction © The McGraw−Hill
Genetics, Seventh Edition Scientific Method Companies, 2001

A Brief Overview of the Modern History of Genetics 3

enetics is the study of inheritance in all of its 1860-1900

G manifestations, from the distribution of hu-

man traits in a family pedigree to the bio-
chemistry of the genetic material in our
chromosomes—deoxyribonucleic acid, or
DNA. It is our purpose in this book to introduce and de-
scribe the processes and patterns of inheritance. In this
chapter, we present a broad outline of the topics to be
The period from 1860 to 1900 encompasses the publica-
tion of Gregor Mendel’s work with pea plants in 1866 to
the rediscovery of his work in 1900. It includes the dis-
coveries of chromosomes and their behavior—insights
that shed new light on Mendel’s research.
From 1879 to 1885, with the aid of new staining tech-
niques, W. Flemming described the chromosomes—first
covered as well as a summary of some of the more im-
noticed by C. von Nägeli in 1842—including the way they
portant historical advancements leading to our current
split during division, and the separation of sister chromatids
understanding of genetics.
and their movement to opposite poles of the dividing cell
during mitosis. In 1888, W. Waldeyer first used the term
chromosome. In 1875, O. Hertwig described the fusion of
sperm and egg to form the zygote. In the 1880s, Theodor
A BRIEF OVERVIEW OF Boveri, as well as K. Rabl and E. van Breden, hypothesized
THE MODERN HISTORY that chromosomes are individual structures with continuity
OF GENETICS from one generation to the next despite their “disappear-
ance” between cell divisions. In 1885, August Weismann
For a generation of students born at a time when incred-
stated that inheritance is based exclusively in the nucleus.
ible technological advances are commonplace, it is valu-
In 1887, he predicted the occurrence of a reductional di-
able to see how far we have come in understanding the
vision, which we now call meiosis. By 1890, O. Hertwig and
mechanisms of genetic processes by taking a very brief,
T. Boveri had described the process of meiosis in detail.
encapsulated look at the modern history of genetics. Al-
though we could discuss prehistoric concepts of animal
and plant breeding and ideas going back to the ancient 1900-1944
Greeks, we will restrict our brief look to events begin-
From 1900 to 1944, modern genetics flourished with the
ning with the discovery of cells and microscopes. For our
development of the chromosomal theory, which showed
purposes, we divide this recent history into four periods:
before 1860, 1860–1900, 1900–1944, and 1944 to the
present.

Before 1860
Before 1860, the most notable discoveries paving the
way for our current understanding of genetics were
the development of light microscopy, the elucidation of
the cell theory, and the publication in 1859 of Charles
Darwin’s The Origin of Species. In 1665, Robert Hooke
coined the term cell in his studies of cork. Hooke saw, in
fact, empty cells observed at a magnification of about
thirty power. Between 1674 and 1683, Anton van
Leeuwenhoek discovered living organisms (protozoa and
bacteria) in rainwater. Leeuwenhoek was a master lens
maker and produced magnifications of several hundred
power from single lenses (fig. 1.1). More than a hundred
years passed before compound microscopes could equal
Leeuwenhoek’s magnifications. In 1833, Robert Brown
(the discoverer of Brownian motion) discovered the nu-
clei of cells, and between 1835 and 1839, Hugo von Mohl
described mitosis in nuclei.This era ended in 1858, when
Rudolf Virchow summed up the concept of the cell the-
ory with his Latin aphorism omnis cellula e cellula: all
cells come from preexisting cells. Thus, by 1858, biolo- Figure 1.1 One of Anton van Leeuwenhoek’s microscopes,
gists had an understanding of the continuity of cells and ca. 1680. This single-lensed microscope magnifies up to 200x.
knew of the cell’s nucleus. (© Kathy Talaro/Visuals Unlimited, Inc.)
 Tamarin: Principles of I. Genetics and the 1. Introduction © The McGraw−Hill
Genetics, Seventh Edition Scientific Method Companies, 2001

4 Chapter One Introduction

that chromosomes are linear arrays of genes. In addition, cleases, the enzymes that opened up our ability to ma-
the foundations of modern evolutionary and molecular nipulate DNA through recombinant DNA technology. In
genetics were derived. 1972, Paul Berg was the first to create a recombinant
In 1900, three biologists working independently— DNA molecule.
Hugo de Vries, Carl Correns, and Erich von Tschermak— Since 1972, geneticists have cloned numerous genes.
rediscovered Mendel’s landmark work on the rules of in- Scientists now have the capability to create transgenic
heritance, published in 1866, thus beginning our era of organisms, organisms with functioning foreign genes. For
modern genetics. In 1903, Walter Sutton hypothesized example, we now have farm animals that produce phar-
that the behavior of chromosomes during meiosis ex- maceuticals in their milk that are harvested easily and in-
plained Mendel’s rules of inheritance, thus leading to the expensively for human use. In 1997, the first mammal
discovery that genes are located on chromosomes. In was cloned, a sheep named Dolly. The sequence of the
1913, Alfred Sturtevant created the first genetic map, us- entire human genome was determined in 2000; we will
ing the fruit fly. He showed that genes existed in a lin- spend the next century mining its information in the
ear order on chromosomes. In 1927, L. Stadler and newly created field of genomics, the study of the com-
H. J. Muller showed that genes can be mutated artificially plete genetic complement of an organism. Although no
by X rays. inherited disease has yet been cured by genetic interven-
Between 1930 and 1932, R. A. Fisher, S. Wright, and tion, we are on the verge of success in numerous dis-
J. B. S. Haldane developed the algebraic foundations for eases, including cancer.
our understanding of the process of evolution. In 1943, The material here is much too brief to convey any of
S. Luria and M. Delbrück demonstrated that bacteria have the detail or excitement surrounding the discoveries of
normal genetic systems and thus could serve as models modern genetics. Throughout this book, we will expand
for studying genetic processes. on the discoveries made since Darwin first published his
book on evolutionary theory in 1859 and since Mendel
was rediscovered in 1900.
1944-Present
The period from 1944 to the present is the era of molec-
ular genetics, beginning with the demonstration that
DNA is the genetic material and culminating with our THE THREE GENERAL AREAS
current explosion of knowledge due to recombinant OF GENETICS
DNA technology.
In 1944, O. Avery and colleagues showed conclu- Historically, geneticists have worked in three different ar-
sively that deoxyribonucleic acid—DNA—was the ge- eas, each with its own particular problems, terminology,
netic material. James Watson and Francis Crick worked tools, and organisms. These areas are classical genetics,
out the structure of DNA in 1953. Between 1968 and molecular genetics, and evolutionary genetics. In classi-
1973, W. Arber, H. Smith, and D. Nathans, along with their cal genetics, we are concerned with the chromosomal
colleagues, discovered and described restriction endonu- theory of inheritance; that is, the concept that genes are

Table 1.1 The Three Major Areas of Genetics_Classical, Molecular, and Evolutionary_
and the Topics They Cover

Classical Genetics Molecular Genetics Evolutionary Genetics

Mendel’s principles Structure of DNA Quantitative genetics

Meiosis and mitosis Chemistry of DNA Hardy-Weinberg equilibrium
Sex determination Transcription Assumptions of equilibrium
Sex linkage Translation Evolution
Chromosomal mapping DNA cloning and genomics Speciation
Cytogenetics (chromosomal changes) Control of gene expression
DNA mutation and repair
Extrachromosomal inheritance
Tamarin: Principles of I. Genetics and the 1. Introduction © The McGraw−Hill
Genetics, Seventh Edition Scientific Method Companies, 2001

How Do We Know? 5

located in a linear fashion on chromosomes and that the

Observation
relative positions of genes can be determined by their
frequency in offspring. Molecular genetics is the study of
the genetic material: its structure, replication, and ex-
pression, as well as the information revolution emanating
Hypothesis
from the discoveries of recombinant DNA techniques
(genetic engineering, including the Human Genome Proj-
ect). Evolutionary genetics is the study of the mecha-
nisms of evolutionary change, or changes in gene fre-
Prediction
quencies in populations. Darwin’s concept of evolution
by natural selection finds a firm genetic footing in this Support
area of the study of inheritance (table 1.1).
Today these areas are less clearly defined because of
advances made in molecular genetics. Information com- Experiment
ing from the study of molecular genetics allows us to un-
derstand better the structure and functioning of chromo- Refute
somes on the one hand and the mechanism of natural
selection on the other. In this book we hope to bring to- New hypothesis
gether this information from a historical perspective.
From Mendel’s work in discovering the rules of inheri-
Figure 1.2 A schematic of the scientific method. An
tance (chapter 2) to genetic engineering (chapter 13) to
observation leads the researcher to propose a hypothesis, and
molecular evolution (chapter 21), we hope to present a
then to make predictions from the hypothesis and to test these
balanced view of the various topics that make up
predictions by experiment. The results of the experiment either
genetics.
support or refute the hypothesis. If the experiment refutes the
hypothesis, a new hypothesis must be developed. If the
experiment supports the hypothesis, the researcher or others
design further experiments to try to disprove it.
HOW DO WE KNOW?
Genetics is an empirical science, which means that our time, the longer-necked giraffes would survive and
information comes from observations of the natural reproduce better than the shorter-necked ones. Thus,
world. The scientific method is a tool for understanding longer necks would come to predominate. Any genetic
these observations (fig. 1.2). At its heart is the experi- mutations (changes) that introduced greater neck length
ment, which tests a guess, called a hypothesis, about how would be favored.
something works. In a good experiment, only two types To test Lamarck’s hypothesis, you might begin by de-
of outcomes are possible: outcomes that support the hy- signing an experiment. You could do the experiment on
pothesis and outcomes that refute it. Scientists say these giraffes to test Lamarck’s hypothesis directly; however, gi-
outcomes provide strong inference. raffes are difficult to acquire, maintain, and breed. Re-
For example, you might have the idea that organisms member, though, that you are testing a general hypothe-
can inherit acquired characteristics, an idea put forth by sis about the inheritance of acquired characteristics
Jean-Baptiste Lamarck (1744–1829), a French biologist. rather than a specific hypothesis about giraffes. Thus, if
Lamarck used the example of short-necked giraffes evolv- you are clever enough, you can test the hypothesis with
ing into the long-necked giraffes we know of today. He almost any organism. You would certainly choose one
suggested that giraffes that reached higher into trees to that is easy to maintain and manipulate experimentally.
get at edible leaves developed longer necks. They passed Later, you can verify the generality of any particular con-
on these longer necks to their offspring (in small incre- clusions with tests on other organisms.
ments in each generation), leading to today’s long-necked You might decide to use lab mice, which are relatively
giraffes. An alternative view, evolution by natural selec- inexpensive to obtain and keep and have a relatively
tion, was put forward in 1859 by Charles Darwin. Ac- short generation time of about six weeks, compared with
cording to the Darwinian view, giraffes normally varied the giraffe’s gestation period of over a year. Instead of
in neck length, and these variations were inherited. looking at neck length, you might simply cut off the tip of
Giraffes with slightly longer necks would be at an advan- the tail of each mouse (in a painless manner), using short-
tage in reaching edible leaves in trees. Therefore, over ened tails as the acquired characteristic. You could then
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