Beruflich Dokumente
Kultur Dokumente
GENETIC ANALYSIS
About the Authors
Anthony J. F. Griffiths
University of British Columbia
Susan R. Wessler
University of California, Riverside
Sean B. Carroll
Howard Hughes Medical Institute
University of Wisconsin-Madison
John Doebley
University of Wisconsin-Madison
First printing
www.whfreeman.com
Contents in Brief Contents
Preface xiii Preface xiii
v
vi CONTENTS
3.1 Mendel’s Law of Independent Assortment 82 4.4 Centromere Mapping with Linear Tetrads 142
3.2 Working with Independent Assortment 86 4.5 Using the Chi-Square Test for Testing
Predicting progeny ratios 86
Linkage Analysis 143
Using the chi-square test on monohybrid and
4.6 Accounting for Unseen Multiple Crossovers 145
dihybrid ratios 89 A mapping function 146
Synthesizing pure lines 91 The Perkins formula 147
Hybrid vigor 93 4.7 Using Recombination-Based Maps in
3.3 The Chromosomal Basis of Independent Conjunction with Physical Maps 148
Assortment 94 4.8 The Molecular Mechanism of Crossing Over 150
Independent assortment in diploid organisms 95
Independent assortment in haploid organisms 96
5 The Genetics of Bacteria and
Independent assortment of combinations of
autosomal and X-linked genes 97
Their Viruses 169
Recombination 99 5.1 Working with Microorganisms 171
3.4 Polygenic Inheritance 101 5.2 Bacterial Conjugation 173
Discovery of conjugation 173
3.5 Organelle Genes: Inheritance Independent
of the Nucleus 103 Discovery of the fertility factor (F) 175
Patterns of inheritance in organelles 104 Hfr strains 176
Cytoplasmic segregation 105 Mapping of bacterial chromosomes 180
Cytoplasmic mutations in humans 107 F plasmids that carry genomic fragments 184
MtDNA in evolutionary studies 109 R plasmids 184
CONTENTS vii
6.3 Inferring Gene Interactions 222 Self-splicing introns and the RNA world 300
Sorting mutants using the complementation test 222 8.5 Small Functional RNAs that Regulate and
Analyzing double mutants of random mutations 226 Protect the Eukaryotic Genome 300
miRNAs are important regulators of
6.4 Penetrance and Expressivity 233 gene expression 300
siRNAs ensure genome stability 302
PART II FROM DNA TO PHENOTYPE
Similar mechanisms generate siRNA and miRNA 305
7 DNA: Structure and Replication 251
9 Proteins and Their Synthesis 309
7.1 DNA: The Genetic Material 252
Discovery of transformation 252 9.1 Protein Structure 311
Hershey–Chase experiment 254 9.2 The Genetic Code 314
7.2 The DNA Structure 256 Overlapping versus nonoverlapping codes 314
DNA structure before Watson and Crick 256 Number of letters in the codon 315
The double helix 258 Use of suppressors to demonstrate a
triplet code 315
7.3 Semiconservative Replication 261
Degeneracy of the genetic code 317
Meselson–Stahl experiment 262
The replication fork 263 Cracking the code 317
DNA polymerases 264 Stop codons 318
7.4 Overview of DNA Replication 266 9.3 tRNA: The Adapter 319
7.5 The Replisome: A Remarkable Replication Codon translation by tRNA 319
Machine 268 Degeneracy revisited 321
Unwinding the double helix 269 9.4 Ribosomes 322
Assembling the replisome: replication initiation 270 Ribosome features 324
viii CONTENTS
Translation initiation, elongation, and termination 325 Genetic analysis of the lac promoter 392
Nonsense suppressor mutations 328 Molecular characterization of the Lac
repressor and the lac operator 392
9.5 The Proteome 329
Alternative splicing generates protein isoforms 329 Polar mutations 393
12.5 Long-Term Inactivation of Genes in a 13.6 From Flies to Fingers, Feathers, and Floor Plates:
Chromatin Environment 435 The Many Roles of Individual Toolkit Genes 479
Mating-type switching and gene silencing 435 13.7 Development and Disease 480
Heterochromatin and euchromatin compared 437 Polydactyly 480
Position-effect variegation in Drosophila reveals Holoprosencephaly 481
genomic neighborhoods 437 Cancer as a developmental disease 481
Genetic analysis of PEV reveals proteins
necessary for heterochromatin formation 439 14 Genomes and Genomics 487
12.6 Gender-Specific Silencing of Genes and 14.1 The Genomics Revolution 489
Whole Chromosomes 441 14.2 Obtaining the Sequence of a Genome 490
Genomic imprinting explains some unusual Turning sequence reads into an assembled
patterns of inheritance 441 sequence 490
But what about Dolly and other cloned Whole-genome sequencing 492
mammals? 443 Traditional WGS 492
Silencing an entire chromosome: Next-generation whole-genome shotgun
X-chromosome inactivation 443 sequencing 493
12.7 Post-Transcriptional Gene Repression Whole-genome-sequence assembly 495
by miRNAs 444 Toward the personalized genome 497
14.3 Bioinformatics: Meaning from Genomic
13 The Genetic Control of Sequence 497
Development 449 The nature of the information content of DNA 498
Deducing the protein-encoding genes from
13.1 The Genetic Approach to Development 450 genomic sequence 498
13.2 The Genetic Toolkit for Drosophila
14.4 The Structure of the Human Genome 502
Development 452
Classification of genes by developmental function 453 14.5 Comparative Genomics 505
Phylogenetic inference 505
Homeotic genes and segmental identity 454
Of mice and humans 507
Organization and expression of Hox genes 455
Comparative genomics of chimpanzees and
The homeobox 457 humans 509
Clusters of Hox genes control development in Comparative genomics of humans 509
most animals 458 Conserved and ultraconserved noncoding
13.3 Defining the Entire Toolkit 461 elements 510
The anteroposterior and dorsoventral axes 462 Comparative genomics of nonpathogenic and
pathogenic E. coli 511
Expression of toolkit genes 463
14.6 Functional Genomics and Reverse
13.4 Spatial Regulation of Gene Expression in Genetics 512
Development 466
Ome, sweet ome 513
Maternal gradients and gene activation 466
Reverse genetics 516
Drawing stripes: integration of gap-protein inputs 468
Making segments different: integration of
PART III MUTATION, VARIATION,
Hox inputs 471
AND EVOLUTION
13.5 Posttranscriptional Regulation of Gene
Expression in Development 473 15 The Dynamic Genome:
RNA splicing and sex determination in Transposable Elements 523
Drosophila 473
Regulation of mRNA translation and cell lineage 15.1 Discovery of Transposable Elements
in C. elegans 475 in Maize 524
Translational control in the early embryo 475 McClintock’s experiments: the Ds element 524
Autonomous and nonautomous elements 527
miRNA control of developmental timing in
C. elegans and other species 478 Transposable elements: only in maize? 528
x CONTENTS
Error-prone repair: translesion DNA synthesis 575 Recombination and linkage disequilibrium 659
Repair of double-strand breaks 577 Gene drift and population size 661
S
ince its first edition in 1974, Introduction to Genetic Analysis has emphasized
the power and incisiveness of the genetic approach in biological research
and its applications. Over its many editions, the text has continuously
expanded its coverage as the power of traditional genetic analysis has been extended
with the introduction of recombinant DNA technology and then genomics. In the
tenth edition, we incorporate the practice of modern genetics into new chapters
on population genetics and the inheritance of complex traits.
xiii
xiv PREFACE
In addition, the final chapter on the evolution of genes and traits has been
heavily revised by Sean Carroll to reflect recent advances in understanding how
adaptations arise. New topics include the classic story of the evolution of malarial
resistance in humans, multistep evolutionary pathways, and the loss of characters
through adaptive changes in regulatory sequences (illustrated by pelvic reduction
in fish populations). Together, the chapter’s many lucid examples provide a firm
empirical foundation for the theory of evolution by natural selection.
IGF1
10-4
Significance threshold
P value
10-2
1
35 40 45 50 46 51 43 48 12 17 22 27 3 8
chr 15 chr 1 chr 2 chr 3 chr 34 chr 37
Position (Mb)
Figure 19-18 Results from an association-mapping experiment for body size in dogs.
Each dot in the plot represents the P value for a test of association between an SNP and
body size. Dots above the “threshold line” show evidence for a statistically significant
association. [Photo: Tetra Images/Corbis.]
1. unc-22 dsRNA synthesized in lab. 1. Transgene inserted into petunia cells. 1. Viral gene inserted into tobacco plant.
Antisense
Transgene Gene
dsRNA
Sense
Endogenous pigment gene
2. dsRNA injected into C. elegans embryos. 2. Adults grown from transformed cells 2. Plant exposed to virus but remains
have white sectors in flowers. healthy.
Micropipette
with dsRNA
solution
Conclusion: unc-22 gene silenced Conclusion: Transgene and Conclusion: viral gene silenced
endogenous pigment gene silenced
Modern Techniques: The techniques chapter appears earlier in the book, as Figure 8-21 Three experiments reveal
Chapter 10 in the tenth edition. This chapter introduces students to “retail” tech- key features of gene silencing.
niques commonly used in labs, while the first section of the genomics chapter
(Chapter 14) focuses on the “wholesale” techniques used for massive genome se-
quencing projects. Both chapters have been updated to include modern methods
used for solving genetics problems, including
Enduring Features
Coverage of Model Organisms
The tenth edition retains the enhanced coverage of model systems in formats that
are practical and flexible for both students and instructors.
PREFACE xvii
Chapter 1 introduces some key genetic model organisms and highlights some
of the successes achieved through their use.
Model Organism boxes presented in context where appropriate provide
additional information about the organism in nature and its use
experimentally.
A Brief Guide to Model Organisms, at the back of the book, provides quick
access to essential, practical information about the uses of specific model
organisms in research studies.
An Index to Model Organisms, on the endpapers at the back of the book,
provides chapter-by-chapter page references to discussions of specific
organisms in the text, enabling instructors and students to easily find and
assemble comparative information across organisms.
Problem Sets
No matter how clear the exposition, deep understanding requires the student to
personally engage with the material. Hence our efforts to encourage student
problem solving. Building on its focus on genetic analysis, the tenth edition pro-
vides students with opportunities to practice problem-solving skills—both in the
text and online through the following features:
Versatile Problem Sets. Problems span the full range of degrees of
difficulty. They are categorized according to level of difficulty—basic
or challenging.
NEW Working with the Figures. A new set of problems included at the back
of each chapter asks students pointed questions about figures in the chapter.
These questions encourage students to think about the figures and help them
to assess their understanding of key concepts.
Solved Problems. Found at the end of each chapter, these worked examples
illustrate how geneticists apply principles to experimental data.
Unpacking the Problems. A genetics problem draws on a complex matrix
of concepts and information. “Unpacking the Problem” helps students learn
to approach problem solving strategically, one step at a time, concept on
concept.
NEW Multiple-choice versions of the end-of-chapter
problems are available on our online GeneticsPortal for quick gradable
quizzing and easily gradable homework assignments. The Unpacking the
Problem tutorials from the text have been converted to in-depth online
tutorials and expanded to help students learn to solve problems and think
like a geneticist.
Electronic Resources
eBook
(ISBN: 1-4292-3257-9)
The eBook fully integrates the text and its interactive media in a format that fea-
tures a variety of helpful study tools (full-text, Google-style searching; note tak-
ing; bookmarking; highlighting; and more). Available as a stand-alone item or on
the GeneticsPortal.
Clicker Questions
Jump-start discussions, illuminate important points, and promote better concep-
tual understanding during lectures.
Layered PowerPoints
Illuminate challenging topics for students by deconstructing intricate genetic
concepts, sequences, and processes step-by-step in a visual format.
All Images from the Text
More than 500 illustrations can be downloaded as JPEGs and PowerPoints. Use
high-resolution images with enlarged labels to project clearly for lecture hall pre-
sentations. Additionally, these JPEG and PowerPoint files are available without
labels for easy customization in PowerPoint.
45 Step-Through and Continuous Play FLASH Animations
These animations were authored by Anthony Griffiths in conjunction with
BioStudio Visual Communications. The complete list of animations appears on
page xxi.
Assessment Bank
This resource brings together a wide selection of genetics problems for use
in testing, homework assignments, or in-class activities. Searchable by topic
and provided in MS Word format, the assessment bank offers a high level of
flexibility.
Student Solutions Manual
(ISBN: 1-4292-3255-2)
The Student Solutions Manual contains complete worked-out solutions to all
the problems in the textbook, including the “Unpacking the Problem” exercises.
Available on the GeneticsPortal and the Instructor’s Web site as easy-to-print
Word files.
xx PREFACE
Print Resources
Overhead Transparency Set (ISBN: 1-4292-7566-9)
The full-color overhead transparency set contains 150 key illustrations from the
text with enlarged labels that project more clearly for lecture hall presentation.
Understanding Genetics: Strategies for Teachers and
Learners in Universities and High Schools
(ISBN: 0-7167-5216-6)
Written by Anthony Griffiths and Jolie-Mayer Smith, this collection of articles
focuses on problem solving and describes methods for helping students improve
their ability to process and integrate new information.
Animations
Forty-five animations developed by Anthony Griffiths are fully integrated with
the content and figures in the text chapters. These animations are available on the
GeneticsPortal and the Book Companion site.
CHAPTER 1
Three-Dimensional Structure of Nuclear Chromosomes (Figure 1-10)
RFLP Analysis and Gene Mapping
CHAPTER 2
Mitosis (Chapter Appendix 2-1)
Meiosis (Chapter Appendix 2-2)
CHAPTER 3
Meiotic Recombination Between Unlinked Genes by Independent Assortment
(Figures 3-8 and 3-13)
CHAPTER 4
Meiotic Recombination Between Linked Genes by Crossing Over
(Figure 4-7)
A Mechanism of Crossing Over: A Heteroduplex Model (Figure 4-21)
A Mechanism of Crossing Over: Genetic Consequences of the Heteroduplex
Model
CHAPTER 5
Bacterial Conjugation and Mapping by Recombination (Figures 5-11 and 5-17)
CHAPTER 6
Interactions Between Alleles at the Molecular Level, RR: Wild-Type
Interactions Between Alleles at the Molecular Level, rr: Homozygous
Recessive, Null Mutation
Interactions Between Alleles at the Molecular Level, r`r`: Homozygous Recessive,
Leaky Mutation
Interactions Between Alleles at the Molecular Level, Rr: Heterozygous,
Complete Dominance
CHAPTER 7
DNA Replication: The Nucleotide Polymerization Process (Figure 7-15)
DNA Replication: Coordination of Leading and Lagging Strand Synthesis
(Figure 7-20)
DNA Replication: Replication of a Chromosome (Figure 7-24)
CHAPTER 8
Transcription (Figure 8-4)
CHAPTER 9
Translation: Peptide-Bond Formation (Figure 9-2)
Translation: The Three Steps of Translation (Figure 9-16)
Nonsense Suppression at the Molecular Level: The rodns Nonsense Mutation
(Figure 9-18)
Nonsense Suppression at the Molecular Level: The tRNA Nonsense Suppressor
(Figure 9-18)
Nonsense Suppression at the Molecular Level: Nonsense Suppression of the rodns
Allele (Figure 9-18)
CHAPTER 10
Polymerase Chain Reaction (Figure 10-3)
Finding Specific Cloned Genes by Functional Complementation: Functional
Complementation of the Gal– Yeast Strain and Recovery of the Wild-Type
GAL gene
xxii PREFACE
Acknowledgments
We extend our thanks and gratitude to our colleagues who reviewed this edition
and whose insights and advice were most helpful:
Mirjana M. Brockett, Georgia Institute of Technology Andrew G. McCubbin, Washington State University
Judy Brusslan, California State University, Long Beach Debra M. McDonough, University of New England
Michael A. Buratovich, Spring Arbor University R. A. McGowan, Memorial University of Newfoundland,
Soochin Cho, Creighton University St. John’s
Matthew H. Collier, Wittenberg University Thomas M. McGuire, Penn State University, Abington
Erin J. Cram, Northeastern University Leilani M. Miller, Santa Clara University
Kenneth A. Curr, California State University, East Bay Erin R. Morris, Baker University
Ann Marie Davison, Kwantlen Polytechnic University Rebecca J. Mroczek-Williamson, University of Arkansas,
Kim Dej, McMaster University Fort Smith
Michael Deyholos, University of Alberta Todd C. Nickle, Mount Royal University
Christine M. Fleet, Emory & Henry College Thomas R. Peavy, California State University, Sacramento
Kimberly Gallagher, University of Pennsylvania Michael Perlin, University of Louisville
Michael A. Gilchrist, University of Tennessee, Knoxville Lynn A. Petrullo, College of New Rochelle
Jamie Lyman Gingerich, University of Wisconsin, Eau-Claire David K. Peyton, Morehead State University
Paul Goldstein, University of Texas, El Paso Jeffrey L. Reinking, SUNY, New Paltz
Julie Goodliffe, University of North Carolina, Charlotte Turk Rhen, University of North Dakota
Thomas A. Grigliatti, University of British Columbia Inder Saxena, University of Texas at Austin
Bruce Haggard, Hendrix College Daniel Schoen, McGill University
Jody L. Hall, Brown University David Scott, South Carolina State University
Mike Harrington, University of Alberta Rebecca L. Seipelt, Middle Tennessee State University
Donna Hazelwood, Dakota State University Bin Shuai, Wichita State University
Deborah Hettinger, Texas Lutheran University Elaine A. Sia, University of Rochester
Brian A. Hyatt, Bethel University Loren C. Skow, Texas A&M University
Glenn H. Kageyama, California State Polytechnic University, Christopher Somers, University of Regina
Pomona Marc Spingola, University of Missouri, St. Louis
Pamela Kalas, University of British Columbia Michael Stock, Grant MacEwan College,
Kathleen Karrer, Marquette University City Centre Campus
Elena L. Keeling, California Polytechnic State University Jared L. Strasburg, Washington University
Michele C. Kieke, Concordia University, St. Paul Aram D. Stump, Adelphi University
Dubear Kroening, University of Wisconsin, Fox Valley Dan Szymanski, Purdue University
James A. Langeland, Kalamazoo College Frans E. Tax, University of Arizona
Janine LeBlanc-Straceski, Merrimack College Justin Thackeray, Clark University
Brenda G. Leicht, University of Iowa Laura G. Vallier, Hofstra University
Steven W. L’ Hernault, Emory University Jacob Varkey, Humboldt State University
Stefan Maas, Lehigh University Michael K. Watters, Valparaiso University
Jeffrey Marcus, Western Kentucky University Marta L. Wayne, University of Florida
Michael Martin, John Carroll University Darla J. Wise, Concord University
Sean Carroll would like to thank Leanne Olds for help with the artwork for
Chapters 11, 12, 13, 14, and 20. John Doebley would like to thank his University of
Wisconsin colleagues Bill Engels, Carter Denniston, and Jim Crow, who shaped his
approach to teaching genetics.
The authors also thank the team at W. H. Freeman for their hard work and
patience. In particular we thank our developmental editor, Susan Moran; exec-
utive editor Susan Winslow; senior project editor Mary Louise Byrd; and copy
editor Karen Taschek. We also thank Paul Rohloff, production coordinator; Diana
Blume, art director; Marsha Cohen, who designed the page layouts; Bill Page and
Janice Donnola, illustration coordinators; Bianca Moscatelli, photo editor; Aaron
Gass, media editor; Anna Bristow and Brittany Murphy, supplements editors; and
Brittany Murphy and Heidi Bamatter, editorial assistants. Finally, we especially
appreciate the marketing and sales efforts of Debbie Clare, associate director of
marketing, and the entire sales force.
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The Genetics 1
Revolution in the
Life Sciences
KEY QUESTIONS
s 7HAT CONSTITUTES BIOLOGICAL
INFORMATION AND HOW DOES IT
GENERATE FORM FROM RANDOM
ENVIRONMENTAL COMPONENTS
s (OW IS LIFE ABLE TO PERSIST DOWN
THE GENERATIONS
s 7HAT IS THE BASIS OF HEREDITARY
VARIATION
s (OW DID SPECIES ARISE ON THE
PLANET
s (OW HAS GENETICS AFFECTED
HUMAN SOCIETY
s (OW DOES GENETIC RESEARCH
WORK
s (OW WILL GENETICS AFFECT OUR
FUTURE
'ENETIC VARIATION IN THE COLOR OF CORN KERNELS %ACH KERNEL REPRESENTS A SEPARATE INDIVIDUAL
WITH A DISTINCT GENETIC MAKEUP 4HE PHOTOGRAPH SYMBOLIZES THE HISTORY OF HUMANITYS INTEREST OUTLINE
IN HEREDITY (UMANS WERE BREEDING CORN THOUSANDS OF YEARS BEFORE THE ADVENT OF THE MODERN
DISCIPLINE OF GENETICS %XTENDING THIS HERITAGE CORN TODAY IS ONE OF THE MAIN RESEARCH ORGANISMS
1.1 4HE NATURE OF BIOLOGICAL
IN CLASSICAL AND MOLECULAR GENETICS ;William Sheridan, University of North Dakota; photograph INFORMATION
by Travis Amos.]
1.2 (OW INFORMATION BECOMES
BIOLOGICAL FORM
O
ur planet Earth is teeming with life (Figure 1-1), and this living world 1.3 'ENETICS AND EVOLUTION
has been a subject of great curiosity and investigation since the dawn of
1.4 'ENETICS HAS PROVIDED A
civilization. However, in the last 60 years a revolution has taken place
in our understanding of the living world, and the foundation for this revolution
POWERFUL NEW APPROACH TO
has been the discoveries of genetic research. Today, most of the main questions BIOLOGICAL RESEARCH
of biology have been answered through genetics, largely through an under- 1.5 -ODEL ORGANISMS HAVE BEEN
standing of molecular and cellular mechanisms centered on DNA. The molecule CRUCIAL IN THE GENETICS REVOLUTION
deoxyribonucleic acid (DNA) is the central topic of interest to geneticists, but
it has also become a kind of logo for all of the life sciences. The unfolding of our
1.6 'ENETICS CHANGES SOCIETY
understanding of the nature of DNA and how it operates has not only provided 1.7 'ENETICS AND THE FUTURE
1
2 CHAPTER 1 The Genetics Revolution in the Life Sciences
basic answers to the core questions of all areas of biology, but has also led to
spectacular applications in many areas of human endeavor such as medicine
and agriculture.
In this chapter, we will provide an overview of the genetics revolution and how
it has come to pass. In doing so, we will see how the old mists have parted, leaving
us with a clear view of the central processes of life at the cell, organism, and popu-
lation levels.
First, we need to define genetics. Broadly, genetics is the study of all aspects of
genes. In turn, genes are defined as the fundamental units of biological informa-
tion. They can be thought of as the words in the language of the living process.
Much was known about genes before the discovery of DNA, but now we know that
in virtually all cases, genes are composed of DNA. Thus, the discovery of DNA led
biological science into a realm called molecular genetics. By and large, molecu-
lar genetics deals with genes one or a few at a time. However, more recent techno-
logical innovations have led to genomics, the study of complete gene sets (called
genomes). Hence, information can be analyzed not only at the level of the
“words,” but at the more complex level of life’s “sentences” and “grammar.” Today,
the term genetics embraces both molecular genetics and genomics.
the nuclei of cells (Figure 1-2). Chromosomes were considered likely informa-
tion carriers because they are passed intact from generation to generation
through precisely orchestrated nuclear divisions called meiosis and mitosis.
In the 1940s, several lines of research showed that the ele-
ment that carries biological information within the chromo- DNA is biological information
somes is the molecule DNA. Eventually, the detailed
molecular structure of DNA was elucidated by James Watson
and Francis Crick in the 1950s. They inferred from this struc-
ture that DNA contains information written in a genetic
code. DNA is a linear series of four molecular building blocks
called nucleotides. The specific sequence of nucleotides con-
stitutes the language of the code. DNA, as part of the chromo-
some, is passed intact from one generation to the next, so all
cells in each generation contain the same set of DNA with the
same information-containing nucleotide sequence. Hence,
one of the big secrets of life had been answered: the architec-
tural blueprint for life is DNA. This discovery was to be a key
step in the genetics revolution. In order to see how DNA
plays its role, we need to understand its structure and its
arrangement in cells.
Before embarking on our broad view of the current state
of genetics, it is worth mentioning that most of the items cov-
ered in this chapter will be revisited for detailed treatment in
subsequent chapters; the treatment here is descriptive rather
than analytical, and the goal is to provide a general overview
of the subject.
to the base facing it in the opposite strand to constitute the “rungs” of the
Complementary base pairing
ladder: adenine in one strand is always paired with thymine in the other,
O 5´ whereas guanine is always paired with cytosine. This bonding specificity
P O is based on complementarity of shape and charge. It is the sequence of A,
3´ O
O H N O T, G, and C on one strand that represents the coded information carried
H
5´ CH2 by the DNA molecule (Figure 1-4).
O T N H N A O 4´
2´
3´ O 1´ 3´
4´ 1´ 2´ -ESSAGE $
.! IS BIOLOGICAL INFORMATION ENCODED AS A SEQUENCE OF
O O
5´ CH2 NUCLEOTIDES $.! IS A DOUBLE HELIX OF TWO NUCLEOTIDE CHAINS HELD TOGETHER BY
P O COMPLEMENTARY PAIRING OF ! WITH 4 AND OF '