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GENE
&
T I
ANALYSIS
C SPRINCIPLES

6e

Robert J. Brooker
 Sixth Edition

ROBERT J. BROOKER
University of Minnesota
GENETICS: ANALYSIS & PRINCIPLES, SIXTH EDITION

Published by McGraw-Hill Education, 2 Penn Plaza, New York NY 10121. Copyright © 2018 by McGraw-Hill Education. All rights reserved.
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Library of Congress Cataloging-in-Publication Data

Names: Brooker, Robert J.

Title: Genetics : analysis & principles / Robert J. Brooker, University of Minnesota.
Description: Sixth edition. | New York NY : McGraw-Hill Education, [2018] |
Includes index.
Identifiers: LCCN 2016034541| ISBN 9781259616020 (alk. paper) | ISBN 1259616029 (alk. paper)
Subjects: LCSH: Genetics.
Classification: LCC QH430 .B766 2018 | DDC 576.5--dc23 LC record available at https://lccn.loc.gov/2016034541
2013035482

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www.mhhe.com
B R I E F C O N T E N T S

P A R T I INTRODUCTION
1 Overview of Genetics   1
12
P A R T I I PATTERNS OF INHERITANCE 13
2 Mendelian Inheritance  18
14
3 Chromosome Transmission During Cell
15
Division and Sexual Reproduction   46
4 Extensions of Mendelian Inheritance   76
PA R T I V M
 OLECULAR PROPERTIES
OF GENES
Gene Transcription and RNA Modification   278
Translation of mRNA   306
Gene Regulation in Bacteria   336
Gene Regulation in Eukaryotes I:
Transcriptional and Translation
Regulation  361

5 Non-Mendelian Inheritance  102 16 Gene Regulation in Eukaryotes II:

Epigenetics  388
6 Genetic Linkage and Mapping in
Eukaryotes  127 17 Non-coding RNAs  411

7 Genetic Transfer and Mapping in 18 Genetics of Viruses   433

Bacteria  155
19 Gene Mutation and DNA Repair   461
8 Variation in Chromosome Structure
20 Recombination, Immunogenetics, and
and Number  177
Transposition  491

PA R T I I I M
 OLECULAR STRUCTURE AND
P A R T V GENETIC TECHNOLOGIES
REPLICATION OF THE GENETIC
MATERIAL 21 Molecular Technologies  511
9 Molecular Structure of DNA and RNA   208 22 Biotechnology  539
10 Chromosome Organization and Molecular 23 Genomics I: Analysis of DNA   563
Structure  229
24 Genomics II: Functional Genomics, Proteomics,
11 DNA Replication  252 and Bioinformatics  589

PA R T V I G
 ENETIC ANALYSIS
OF INDIVIDUALS AND
POPULATIONS

25 Medical Genetics and Cancer   611

26 Developmental Genetics  643

27 Population Genetics  675

28 Complex and Quantitative Traits   707

29 Evolutionary Genetics  732

v
T A B L E O F C O N T E N T S

Preface  ix 4.2
4.3
Dominant and Recessive Alleles 78
Environmental Effects on Gene
Expression 80
8 VARIATION IN CHROMOSOME
STRUCTURE AND NUMBER 177

PA R T I 4.4 Incomplete Dominance, Overdominance, 8.1 Microscopic Examination of Eukaryotic

and Codominance 81 Chromosomes 177
INTRODUCTION 1
4.5 X-Linked Inheritance 86 8.2 Changes in Chromosome Structure:

1 OVERVIEW OF GENETICS 1
4.6

4.7
Sex-Influenced and Sex-Limited
Inheritance 88
Lethal Alleles 90
8.3
8.4
An Overview 180
Deletions and Duplications 181
Inversions and Translocations 187
1.1 The Molecular Expression of Genes   3 8.5 Changes in Chromosome Number:
4.8 Pleiotropy 91
1.2 The Relationship Between Genes and An Overview 192
4.9 Gene Interactions 92
Traits  6 8.6 Variation in the Number of
1.3 Fields of Genetics 11 Chromosomes Within a Set:
1.4 The Science of Genetics 13
5 NON-MENDELIAN
INHERITANCE 102
8.7
Aneuploidy 193
Variation in the Number of Sets of
Chromosomes 195
5.1 Maternal Effect 103
5.2 Epigenetic Inheritance: Dosage 8.8 Natural and Experimental Mechanisms
PA R T I I Compensation 106 That Produce Variation in Chromosome
Number 198
PATTERNS OF INHERITANCE 18 5.3 Epigenetic Inheritance: Genomic
Imprinting 112

2 MENDELIAN INHERITANCE 18
5.4 Extranuclear Inheritance 116 PA R T I I I
MOLECULAR STRUCTURE AND
2.1
2.2
2.3
Mendel’s Study of Pea Plants 19
Law of Segregation 22
Law of Independent Assortment 26
6 GENETIC LINKAGE AND MAPPING
IN EUKARYOTES 127
REPLICATION OF THE GENETIC
MATERIAL  208

9
6.1 Overview of Linkage 127
2.4 Studying Inheritance Patterns in MOLECULAR STRUCTURE OF DNA
6.2 Relationship Between Linkage and
Humans 32 AND RNA 208
Crossing Over 129
2.5 Probability and Statistics 34
6.3 Genetic Mapping in Plants and 9.1 Identification of DNA as the Genetic
Animals 135 Material 208

3 CHROMOSOME TRANSMISSION
DURING CELL DIVISION AND
SEXUAL REPRODUCTION 46
6.4

6.5
Genetic Mapping in Haploid
Eukaryotes 142
Mitotic Recombination 145
9.2

9.3
Overview of DNA and RNA
Structure 211
Nucleotide Structure 212
3.1 General Features of Chromosomes 46 9.4 Structure of a DNA Strand 214

7
3.2 Cell Division 50 GENETIC TRANSFER AND 9.5 Discovery of the Double Helix 215
3.3 Mitosis and Cytokinesis 53 MAPPING IN BACTERIA 155 9.6 Structure of the DNA Double Helix 218
3.4 Meiosis 57 9.7 RNA Structure 222
7.1 Overview of Genetic Transfer in
3.5 Sexual Reproduction 61

10
Bacteria 156 CHROMOSOME
3.6 The Chromosome Theory of Inheritance
7.2 Bacterial Conjugation 157 ORGANIZATION AND
and Sex Chromosomes 64 MOLECULAR STRUCTURE 229
7.3 Conjugation and Mapping via Hfr

4
Strains 161
EXTENSIONS OF MENDELIAN 10.1 Organization of Sites Along Bacterial
7.4 Bacterial Transduction 166 Chromosomes 229
INHERITANCE 76
7.5 Bacterial Transformation 168 10.2 Structure of Bacterial Chromosomes 230
4.1 Overview of Simple Inheritance 7.6 Medical Relevance of Bacterial Genetic 10.3 Organization of Sites Along Eukaryotic
Patterns 77 Transfer 170 Chromosomes 234

vi
 TABLE OF CONTENTS vii

10.4 Sizes of Eukaryotic Genomes and 14.3 Regulation of the trp Operon 349 18.3 Bacteriophage λ Reproductive
Repetitive Sequences 235 14.4 Translational and Posttranslational Cycle 444
10.5 Structure of Eukaryotic Chromosomes in Regulation 353 18.4 HIV Reproductive Cycle 450
Nondividing Cells 237 14.5 Riboswitches 354
10.6 Structure of Eukaryotic Chromosomes
During Cell Division 243 19 GENE MUTATION AND DNA

15
REPAIR 461
GENE REGULATION IN

11
EUKARYOTES I:
19.1 Effects of Mutations on Gene Structure
TRANSCRIPTIONAL AND
DNA REPLICATION 252 TRANSLATION
and Function 462
REGULATION 361 19.2 Random Nature of Mutations 468
11.1 Structural Overview of DNA 19.3 Spontaneous Mutations 470
Replication 252 15.1 Regulatory Transcription Factors 362 19.4 Induced Mutations 475
11.2 Bacterial DNA Replication: The 15.2 Chromatin Remodeling, Histone 19.5 DNA Repair 479
Formation of Two Replication Forks at Variants, and Histone
the Origin of Replication 256 Modification 369
11.3 Bacterial DNA Replication: Synthesis of
New DNA Strands 259
11.4 Bacterial DNA Replication: Chemistry
15.3 DNA Methylation 376
15.4 Insulators 378
20 RECOMBINATION,
IMMUNOGENETICS, AND
TRANSPOSITION 491
15.5 The ENCODE Project 379
and Accuracy 266 20.1 Homologous Recombination 491
15.6 Regulation of Translation 380
11.5 Eukaryotic DNA Replication 268 20.2 Immunogenetics 497
20.3 Transposition 499
PA R T I V
MOLECULAR PROPERTIES
16 GENE REGULATION IN
EUKARYOTES II:
EPIGENETICS 388
PA R T V
OF GENES 278
16.1 Overview of Epigenetics 388 GENETIC TECHNOLOGIES 511

12 GENE TRANSCRIPTION AND

RNA MODIFICATION 278
16.2
16.3
Epigenetics and Development 393
Paramutation 398

12.1
12.2
Overview of Transcription 278
Transcription in Bacteria 281
16.4 Epigenetics and Environmental
Agents 400
16.5 Role of Epigenetics in Cancer 405
21 MOLECULAR
TECHNOLOGIES 511

12.3 Transcription in Eukaryotes 286 21.1 Gene Cloning Using Vectors 512
21.2 Polymerase Chain Reaction 519
12.4
12.5
RNA Modification 291
A Comparison of Transcription
and RNA Modification in Bacteria
17 NON-CODING RNAs 411
21.3
21.4
DNA Sequencing 524
Gene Mutagenesis 526
and Eukaryotes 300 17.1 Overview of Non-coding RNAs 412 21.5 Blotting Methods to Detect Gene
17.2 Non-coding RNAs: Effects on Chromatin Products 529

13 TRANSLATION OF mRNA 306

Structure and Transcription 416
17.3 Non-coding RNAs: Effects on
Translation, mRNA Degradation, and
21.6 Methods for Analyzing DNA- and RNA-
Binding Proteins 531

13.1 The Genetic Basis for Protein

Synthesis 306
13.2 The Relationship Between the Genetic
RNA Modifications 417
17.4 Non-coding RNAs and Protein
Targeting 422
22 BIOTECHNOLOGY 539

Code and Protein Synthesis 309 22.1 Uses of Microorganisms in

13.3 Experimental Determination of the
Genetic Code 315
13.4 Structure and Function of tRNA 319
13.5 Ribosome Structure and Assembly 322
13.6 Stages of Translation 324
17.5 Non-coding RNAs and Genome Biotechnology 539
Defense 423 22.2 Genetically Modified Animals 542
17.6 Role of Non-coding RNAs in Human 22.3 Reproductive Cloning and Stem
Disease 427 Cells 546
22.4 Genetically Modified Plants 551

14 GENE REGULATION IN 18 GENETICS OF VIRUSES 433

22.5 Human Gene Therapy 555

BACTERIA 336

14.1 Overview of Transcriptional

Regulation 337
18.1 Virus Structure and Genetic
Composition 433 23 GENOMICS I: ANALYSIS
OF DNA 563

18.2 Overview of Viral Reproductive 23.1 Overview of Chromosome

14.2 Regulation of the lac Operon 339 Cycles 438 Mapping 564
viii TABLE OF CONTENTS

23.2 Cytogenetic Mapping via

Microscopy 564
23.3 Linkage Mapping via Crosses 567
25.3
25.4
25.5
Genetic Testing and Screening 621
Prions 623
Genetic Basis of Cancer 624
28 COMPLEX AND QUANTITATIVE
TRAITS 707

23.4 Physical Mapping via Cloning and DNA 25.6 Personalized Medicine 634 28.1 Overview of Complex and Quantitative
Sequencing 570 Traits 707
23.5 Genome-Sequencing Projects 574
23.6 Metagenomics 582 26 DEVELOPMENTAL
GENETICS 643
28.2 Statistical Methods for Evaluating
Quantitative Traits 709
28.3 Polygenic Inheritance 712

24
26.1 Overview of Animal Development 643 28.4 Identification of Genes that Control
GENOMICS II: FUNCTIONAL
GENOMICS, PROTEOMICS, AND 26.2 Invertebrate Development 647 Quantitative Traits 715
BIOINFORMATICS 589 26.3 Vertebrate Development 659 28.5 Heritability 717
26.4 Plant Development 662 28.6 Selective Breeding 722
24.1 Functional Genomics 590

29
26.5 Sex Determination in Animals 666
24.2 Proteomics 595

27
24.3 Bioinformatics 600 EVOLUTIONARY GENETICS 732

POPULATION GENETICS 675 29.1 Origin of Species 733

PA R T V I 29.2 Phylogenetic Trees 738
27.1 Genes in Populations and the Hardy-
Weinberg Equation 675 29.3 Molecular Evolution 746
GENETIC ANALYSIS OF
INDIVIDUALS AND 27.2 Overview of Microevolution 680 *Appendix A: Experimental Techniques
POPULATIONS 611 27.3 Natural Selection 681 can be found on the website for this
textbook: www.mhhe.com/
27.4 Genetic Drift 689

25
brookergenetics6e
MEDICAL GENETICS AND 27.5 Migration 692
CANCER 611 Appendix B
27.6 Nonrandom Mating 692
Solutions to Even-Numbered
25.1 Inheritance Patterns of Genetic 27.7 Sources of New Genetic Variation 694 Problems and All Comprehension
Diseases 612 and Concept Check Questions B-1
25.2 Detection of Disease-Causing Alleles via Glossary G-1
Haplotypes 618
Index I-1

ABOUT THE AUTHOR

Robert J. Brooker is a professor in the Department of Genetics,

Cell Biology, and Development and the Department of Biology
Teaching and Learning at the University of Minnesota–
Minneapolis. He received his B.A. in biology from Wittenberg
University in 1978 and his Ph.D. in genetics from Yale University
in 1983. At Harvard, he conducted postdoctoral studies on the
lactose permease, which is the product of the lacY gene of the lac
operon. He continued to work on transporters at the University of
Minnesota with an emphasis on the structure, function, and
regulation of iron transporters found in bacteria and C. elegans. At
the University of Minnesota, he teaches undergraduate courses in
biology and genetics.

DEDICATION

To my wife, Deborah, and our children, Daniel, Nathan, and Sarah

P R E FAC E

I
students may be provided with online lectures, “flipping the class-
room” typically gives students more responsibility for understanding
the textbook material on their own. Along these lines, Genetics:
Analysis & Principles, Sixth Edition, is intended to provide students
n the sixth edition of Genetics: Analysis & Principles, the with a resource that can be effectively used outside of the classroom.
content has been updated to reflect current trends in the field. In Here are several of the key pedagogical features:
addition, the presentation of the content has been improved in a
∙ 
NEW! A new feature called Genetic TIPS provides a
way that fosters active learning. As an author, researcher, and
teacher, I want a textbook that gets students actively involved in consistent approach to help students solve problems in
learning genetics. To achieve this goal, I have worked with a genetics. This approach has three components. First, the
­talented team of editors, illustrators, and media specialists who student is made aware of the Topic at hand. Second, the
have helped me to make the sixth edition of Genetics: Analysis & question is evaluated with regard to the Informaiton that is
Principles a fun learning tool. available to the student. Finally, the student is guided
Overall, an effective textbook needs to accomplish four through one or more Problem-Solving Strategies to tackle
goals.14 First, it needsCHAP toT provide
E R 1 :: comprehensive,
OVERVIEW OF GENETICS accurate, and up- the question.
to-date content in its field. Second, it needs to expose students to
the techniques and skills they will need to become successful in
that field. words,
Third, what an effective
scientifictextbook
question should
was thehave pedagogical
researcher trying
features, such to answer?
as formative assessment, that foster student learn- GENETIC TIPS THE QUESTION: All of the Genetic TIPS
begin with a question. As an example, let’s consider the following
ing. And 3. finally,
Next, the figure follows
it should inspirethe experimental
students so theysteps wantthe to scientist
pursue question:
that field as took to test the
a career. Thehypothesis.
hard workEach thatfeatured
has gone experiment con-
into the sixth The coding strand of DNA in a segment of a gene is as follows:
edition oftains two parallel
Genetics: Analysisillustrations labeled has
& Principles Experimental
been aimed Levelat ATG GGC CTT AGC. This strand carries the information to make a
achieving and Conceptual
all four of theseLevel. goals!The Experimental Level helps you region of a polypeptide with the amino acid sequence, methionine-
to understand the techniques followed. The Conceptual glycine-leucine-serine. What would be the consequences if a mutation
Level helps you to understand what is actually happening changed the second cytosine (C) in this sequence to an adenine (A)?
at each step in the procedure.
FLIPPING
4. The raw data THE for eachCLASSROOM
experiment are then presented. T OPIC: What topic in genetics does this question address? The
5. Last, an interpretation of the data is offered within the text. topic is gene expression. More specifically, the question is about
A recent trend in science education is the phenomenon that is some- the relationship between a gene sequence and the genetic code.
The rationale behind this approach is that it enables you to see the
timesexperimental
called “flipping the classroom.” This phrase refers to the idea
process from beginning to end. As you read through
that some of the activities that usedwillto be done I NFORMATION: What information do you know based on the
the chapters, the experiments help youintoclass
see theare relationship
now done
question and your understanding of the topic? In the question,
outside of class, and vice versa.
between science and scientific theories. For example, instead of spending
the entire As class time lecturing over textbook and other materials, you are given the base sequence of a short segment of a gene and
a student of genetics, you will be given the opportunity told that one of the bases has been changed. From your understanding
sometoofinvolve
the classyourtimemind is in
spent engaging students
the experimental process. in various
As you are activi-
read- of the topic, you may remember that a polypeptide sequence is
ties, ing
such anas problem solving,
experiment, you mayworking through
find yourself case about
thinking studies, and
different determined by reading the mRNA (transcribed from a gene) in
designing experiments.
approaches This approach
and alternative is called
hypotheses. activepeople
Different learning.can For
view groups of three bases called codons.
manytheinstructors,
same datathe and classroom
arrive at hasverybecome
differentmore learner centered
conclusions. As you
rather teacherthrough
progress centered. theAexperiments
learner-centeredin thisclassroom
book, youprovides
will enjoy a P ROBLEM-SOLVING S TRATEGY: Compare and contrast.
rich genetics
environment far morein which
if youstudents can interact
try to develop your own withskills
eachatother and
formulat- One strategy to solve this problem is to compare the mRNA
withingtheirhypotheses,
instructors.designingInstructorsexperiments,
and fellow students often provide
and interpreting data. sequence (transcribed from this gene) before and after the mutation:
Also, some
formative of the questions in the
assessment—immediate problem
feedback thatsets are aimed
helps at refin-
each student Original: AUG GGC CUU AGC
ing these
understand skills.
if his or her learning is on the right track. Mutant: AUG GGC AUU AGC
Finally,
What are some it is worthwhile
advantages to of point
activeoutlearning?
that science is a social
Educational
discipline. As you develop your skills at scrutinizing
studies reveal that active learning usually promotes greater learning experiments, ANSWER: The mutation has changed the sequence of bases in the
gains.it In
is fun to discuss
addition, activeyour ideasoften
learning with focuses
other people,
on skillincluding
developmentfellow mRNA so that the third codon has changed from CUU to AUU.
students and faculty members. Keep in mind that you do not need Because codons specify amino acids, this may change the third
rather than on the memorization of facts that are easily forgotten.
to “know all the answers” before you enter into a scientific discus- amino acid to something else. Note: If you look ahead to Chapter 13
Students become trained to “think like scientists” and to develop a
(see Table 13.1), you will see that CUU specifies leucine, whereas
skill sion. Instead,
set that enables it isthem
moretorewarding to view
apply scientific science asAan
reasoning. ongoing
common AUU specifies isoleucine. Therefore, you would predict that the mu-
and never-ending dialogue.
concern among instructors who are beginning to try out active learn- tation would change the third amino acid from leucine to isoleucine.
ing is that they think they will have less time to teach and therefore
will cover
Genetic less material.
TIPS Will However,
HelpthisYou maytonot be the case.
Improve Although
Your
Problem-Solving Skills Throughout Chapters 2 through 29, each chapter will contain sev- ix
As your progress through this textbook, your learning will involve eral Genetic TIPS. Some of these will be within the chapter itself
two general goals: and some will precede the problem sets that are at the end of each
x PREFACE
∙ 
Genes → Traits: Because genetics is such a broad discipline,
ranging from the molecular level to populations, many
instructors have told us that it is a challenge for students to
see both “the forest and the trees.” It is commonly mentioned
that students often have trouble connecting the concepts they
have learned in molecular genetics with the traits that occur
at the level of a whole organism (i.e., What does
transcription have to do with blue eyes?). To try to make this
connection more meaningful, certain figure legends in each
chapter, designated Genes → Traits, remind students that
molecular and cellular phenomena ultimately lead to the
traits that are observed in each species (see Figure 14.8).
∙ 
Learning Outcomes: Each section of every chapter begins
with a set of learning outcomes. These outcomes help
students understand what they should be able to do once they
have mastered the material in that section.
∙ 
Formative Assessment: When students are expected to learn
textbook material on their own, it is imperative that they are
regularly given formative assessment so they can gauge
whether they are mastering the material. Formative
assessment is a major feature of this textbook and is bolstered
by Connect—a state-of-the art digital assignment and
assessment platform. In Genetics: Analysis & Principles, Sixth
Examples of Specific Content Changes
Edition, formative assessment is provided in multiple ways.
to Individual Chapters
1. As mentioned, a new feature called Genetic TIPS is
aimed at helping students refine their problem solving
skills.
2. Each section of every chapter ends with multiple-choice
questions. Also, compared with the previous edition, many
chapters in the sixth edition are divided into more sections
that are shorter in length. Formative assessment at the end
of each section allows students to evaluate their mastery of
the material before moving on to the next section.
3. Most figures have Concept Check questions so students
can determine if they understand the key points in the
figure.
4. Extensive end-of chapter questions continue to provide
students with feedback regarding their mastery of the
material.
5. The textbook material is supported by digital learning
tools found in Connect. Questions and activities are
assignable in Connect, and students also have access to
our valuable adaptive study tool, SmartBook. With this
tool, students are repeatedly given questions regarding
the textbook material, and depending on their answers,
they may advance ahead in their reading, or they are
given specific advice on what textbook material to go
back and review.
Overall, the pedagogy of Genetics: Analysis & Principles,
sixth edition, has been designed to foster student learning. Instead of
being a collection of facts and figures, Genetics: Analysis & Prin-
ciples, Sixth Edition, by Rob Brooker, is intended to be an engaging
and motivating textbook in which formative assessment allows stu-
dents to move ahead and learn the material in a productive way. We
welcome your feedback so we can make future editions even better!
SIGNIFICANT CONTENT CHANGES
IN THE SIXTH EDITION
∙ 
NEW! A new problem-solving feature called Genetic TIPS
has been added to the sixth edition. The Genetic TIPS are
found within each chapter and three or four are found at the
end of each chapter.
∙ 
NEW! The topic of Epigenetics has been expanded to a
whole chapter, which is now Chapter 16.
∙ 
NEW! A new chapter on non-coding RNA has been added,
which is Chapter 17. This long-overdue chapter is in
response to a remarkable explosion in our appreciation for
the roles of non-coding RNAs in many aspects of molecular
biology. Note: Although two new chapters have been added
to this edition, the overall page length of the sixth edition is
not longer than the fifth edition.
∙ 
NEW! CRISPR-Cas systems: The role of the CRISPR-Cas
system in providing prokaryotes with a genome defense
mechanism is described in Chapter 17, and its use by
researchers to mutate genes is described in Chapter 21.
∙ 
Chapter 2. Mendelian Inheritance: Several Genetic TIPS
have been added to help students work through problem-
solving strategies involving Mendelian inheritance.
∙ 
Chapter 3. Chromosome Transmission During Cell Division
and Sexual Reproduction: The discussion of the random
alignment of homologs during metaphase of meiosis I was
expanded.
∙ 
Chapter 4. Extensions of Mendelian Inheritance: The topic
of gene interaction was streamlined to focus primarily on
examples in which the underlying molecular mechanisms are
known.
∙ 
Chapter 5. Non-Mendelian Inheritance:A common
misconception among students is that you can use a Punnett
square to deduce nonMendelian inheritance patterns.
Throughout the chapter, this misconception has been laid to
rest, and students are given effective strategies to predict
offspring genotypes and phenotypes.
∙ 
Chapter 6. Genetic and Linkage Mapping in Eukaryotes:
When looking at experiments involving linkage, student
often find it very difficult to identify the recombinant
offspring. In various parts of the chapter, a strong effort has
been made to make it clear that recombinant offspring have
inherited a chromosome that is the product of a crossover.
Along these same lines, a new figure (see Figure 6.6) has
been added involving the experiments of Curt Stern showing
that recombinant offspring carry chromosomes that are the
product of a crossover. Also, Figure 6.8 has been revised to
emphasis this point.
∙ 
Chapter 7. Genetic Transfer and Mapping in Bacteria:
Figure 7.13 is a new figure showing the increase in methicillin
resistance in certain Staphylococcus aureus strains.

∙ 
Chapter 8. Variation in Chromosome Structure and Number:
Several Genetic TIPS have been added to help students solve
problems that involve changes in chromosome structure and
chromosome number.
∙ 
Chapter 9. Molecular Structure of DNA and RNA: The
section on the discovery of the DNA double helix has been
streamlined to focus on the key experiments.
∙ 
Chapter 10. Chromosome Organization and Molecular
Structure: The topic of bacterial chromosome structure has
been updated, which includes a new figure (see Figure 10.3)
and a discussion of microdomains.
∙ 
Chapter 11. DNA Replication: A new figure has been added
on the initiation of DNA replication in eukaryotes (see
Figure 11.20).
∙ 
Chapter 12. Gene Transcription and RNA Modification: The
information on alternative splicing has been moved to this
chapter.
∙ 
Chapter 13. Translation of mRNA: Several Genetic TIPS
have been added to help students understand the relationship
between the genetic code and the synthesis of polypeptides.
∙ 
Chapter 14. Gene Regulation in Bacteria: The information
on catabolite activator protein has been updated.
∙ 
Chapter 15. Gene Regulation in Eukaryotes I: Transcriptional
and Translation Regulation: The material on eukaryotic gene
regulation is now divided into two chapters. Chapter 15
focuses on transcriptional and translational regulation.
∙ 
Chapter 16. Gene Regulation in Eukaryotes II: Epigenetics:
This topic has now been expanded to an entire chapter. A
new subsection has been added on the role of epigenetics in
vernalization, which is the process in which some plant
species require an exposure to cold in order to flower the
following spring. Also, a new section has been added on the
intriguing topic of paramutation.
∙ 
Chapter 17. Non-coding RNA: This new chapter begins
with an overview of the general functions of non-coding
RNAs, and then the subsequent sections explore certain
topics in greater detail, such as their role in chromatin
modification, transcription, translation, protein targeting, and
genome defense (e.g., the CRISPR-Cas system).
∙ 
Chapter 18. Genetics of Viruses: The material on the
integration of phage λ has been added to this chapter, along
with a brief discussion of Zika virus. Also, information on
the origin of HIV and the occurrence of HIV infection
worldwide and in the US has been updated.
∙ 
Chapter 19. Gene Mutation and DNA Repair: The information
on the mismatch repair system has been updated.
∙ 
Chapter 20. Recombination, Immunogenetics, and
Transposition: Section 20.2 has been revised to focus on
immunogenetics.
PREFACE xi
∙ 
Chapter 21. DNA Technologies: A new subsection has
been added on gene mutagenesis, which includes a
description of the Crispr-Cas system for inactivating
and mutating genes.
∙ 
Chapter 22. Biotechnology: Several Genetic TIPS have been
added to help students appreciate the uses of molecular
techniques in biotechnology.
∙ 
Chapter 23. Genomics I: Analysis of DNA: The information
has been updated regarding completed genome sequences
and other aspects of genomics.
∙ 
Chapter 24. Genomics II: Functional Genomics, Proteomics,
and Bioinformatics: A new subsection has been added
on the method called RNA-Seq (see Figure 24.3). The
Bioinformatics section has been reorganized with an emphasis
on gene prediction and homology.
∙ 
Chapter 25. Medical Genetics and Cancer: Several
Genetic TIPS have been added to help students
understand how mutations play a role in certain
diseases, including cancer.
∙ 
Chapter 26. Developmental Genetics: The information on
Hox genes in development, and the role of the SRY gene is
human sex determination, have been updated.
∙ 
Chapter 27. Population Genetics: The topic of inbreeding
has been expanded.
∙ 
Chapter 28. Complex and Quantitative Traits: The topic of
the identification of QTLs is now found in its own
subsection.
∙ 
Chapter 29. Evolutionary Genetics: The cladistics method
for constructing a phylogenetic tree is compared with the
UPGMA method.
Suggestions Welcome!
It seems very appropriate to use the word evolution to describe the
continued development of this textbook. I welcome any and all
comments. The refinement of any science textbook requires input
from instructors and their students. These include comments re-
garding writing, illustrations, supplements, factual content, and
topics that may need greater or less emphasis. You are invited to
contact me at:
Dr. Rob Brooker
Dept. of Genetics, Cell Biology, and Development
University of Minnesota
6-160 Jackson Hall
321 Church St.
Minneapolis, MN 55455
brook005@umn.edu
612-624-3053
 ®

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xiv PREFACE

components that need to be assembled to produce a book. I would

ACKNOWLEDGMENTS
The production of a textbook is truly a collaborative effort, and I
am deeply indebted to many people. All six editions of this text-
book went through multiple rounds of rigorous revision that in-
volved the input of faculty, students, editors, and educational and
media specialists. Their collective contributions are reflected in
the final outcome.
Deborah Brooker (Freelance Developmental Editor) metic-
ulously read the new material, analyzed every figure, and offered
extensive feedback. Her attention to detail in this edition and pre-
vious editions has profoundly contributed to the accuracy and
clarity of this textbook. I would also like to thank Jane Hoover
(Freelance Copy Editor) for understanding the material and work-
ing extremely hard to improve the text’s clarity. Her efforts are
truly appreciated.
I would particularly like to acknowledge the many people at
McGraw-Hill Education whose skills and insights are amazing.
My highest praise goes to Elizabeth Sievers (Lead Product Devel-
oper), who carefully checks all aspects of textbook development
and makes sure that all of the pieces of the puzzle are in place.
I am also grateful to Justin Wyatt (Brand Manager) for overseeing
this project. I would like to thank other people at McGraw-Hill
who have played key roles in producing an actual book and the
supplements that go along with it. In particular, Jayne Klein (Senior
Content Project Manager), has done a superb job of managing the
also like to thank Carrie Burger (Content Licensing Specialist),
who acted as an interface between me and the photo company. In
addition, my gratitude goes to David Hash (Designer), who pro-
vided much input into the internal design of the book as well as
created an awesome cover. Finally, I would like to thank Patrick
Reidy (Executive Marketing Manager), whose major efforts begin
when the sixth edition comes out!
I would also like to extend my thanks to everyone at Aptara
who worked with great care in the paging of the book, making sure
that the figures and relevant text are as close to each other as pos-
sible. Likewise, the people at Photo Affairs, Inc. have done a great
job of locating many of the photographs that have been used in the
sixth edition.
Finally, I want to thank the many scientists who reviewed
the chapters of this textbook with special attention to the new
Chapter 17, Non-coding RNAs. Their broad insights and construc-
tive suggestions were an overriding factor that shaped its final
content and organization. I am truly grateful for their time and
compassion.
∙ 
Susan Carpenter University of California, Santa Cruz
∙ 
Johnny El-Rady University of South Florida
∙ 
Terri McElhinny Michigan State University
∙ 
Douglas Wendell Oakland University
∙ 
Jeremy Wilusz University of Pennsylvania Perelman School
of Medicine

REVIEWERS Reggie Cobb, Nash Community College Thomas Peavy, California State University–
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 PA RT I I N T RO D U C T I O N

CHAPTER OUTLINE
1.1 The Molecular Expression
of Genes
1.2 The Relationship Between
Genes and Traits
1.3 Fields of Genetics
1.4 The Science of Genetics

1
CC (for “carbon copy”
or “copy cat”), the first
cloned pet. In 2002, the
cat shown here was
­produced by cloning,
a procedure described
in Chapter 22.
© Corbis

OVERVIEW OF GENETICS
Hardly a week goes by without a major news story involving a
genetic breakthrough. The increasing pace of genetic discoveries
has become staggering. The Human Genome Project is a case in
point. This project began in the United States in 1990, when the
National Institutes of Health and the Department of Energy
joined forces with international partners to decipher the massive
amount of information contained in our genome—the DNA
found within all of our chromosomes (Figure 1.1). Remarkably,
in only a decade, they determined the DNA sequence (the order
of the bases A, T, G, and C) of over 90% of the human genome.
The completed sequence, published in 2003, has an accuracy
greater than 99.99%; less than one mistake was made in every
10,000 base pairs!
In 2008, a more massive undertaking, called the 1000 Ge-
nomes Project, was launched to establish a detailed understand-
ing of human genetic variation. In this international project,
researchers set out to determine the DNA sequence of at least
1000 anonymous participants from around the globe. In 2015,
the sequencing of over 2500 genomes was described in the jour-
nal Nature.
Studying the human genome allows us to explore fundamen-
tal details about ourselves at the molecular level. The results of the
Human Genome Project and the 1000 Genomes Project have shed
considerable light on basic questions, like how many genes we
have, how genes direct the activities of living cells, how species
evolve, how single cells develop into complex tissues, and how
defective genes cause disease. Furthermore, such understanding
may lend itself to improvements in modern medicine by leading to
better diagnoses of diseases and the development of new treat-
ments for them.
The journey to unravel the mysteries within our genes has
­involved the invention of many new technologies. For example, re-
searchers have developed genetic techniques to produce medicines,
such as human insulin, that would otherwise be difficult or impos-
sible to make. Human insulin is synthesized in strains of Esche-
richia coli bacteria that have been genetically altered by the addition
of genes that encode the polypeptides that form this hormone. The
bacteria are grown in a laboratory and make large amounts of hu-
man insulin. As discussed in Chapter 22, the insulin is purified and
administered to many people with insulin-dependent diabetes.

1
2 C H A P T E R 1 :: OVERVIEW OF GENETICS

Chromosomes
DNA, the molecule of life
Cell
The adult human body
is composed of trillions
of cells.

Most human cells contain

the following:
Gene
• 46 human chromosomes,
found in 23 pairs

C G

T A
T A
• 2 meters of DNA

G
C G

T A

T A
• Approximately 22,000

T A
genes coding for

A T

C G
proteins that perform

A T
most life functions
DNA
• Approximately 3 billion
DNA base pairs per set mRNA
of chromosomes,
containing the bases A,
T, G, and C

Amino acid

Protein (composed of amino acids)

FI G U RE 1.1 The human genome. The human genome is a complete set of human chromosomes. People have two sets of chromosomes—one
set from each parent—which are found in the cell nucleus. The Human Genome Project revealed that each set of chromosomes is composed of a DNA
sequence that is approximately 3 billion nucleotide base pairs long. Estimates suggest that each set contains about 22,000 different genes that encode
­proteins. As discussed later, most genes are first transcribed into mRNA and then the mRNA is used to make proteins. This figure emphasizes the DNA
found in the cell nucleus. Humans also have a small amount of DNA in their mitochondria, which has also been sequenced.
CONCEPT CHECK: How might a better understanding of our genes be used in the field of medicine?

New genetic technologies are often met with skepticism and

sometimes even with disdain. An example is mammalian cloning.
In 1997, Ian Wilmut and his colleagues created clones of sheep,
using mammary cells from an adult animal (Figure 1.2). More
recently, such cloning has been achieved in several mammalian
species, including cows, mice, goats, pigs, and cats. In 2002, the
first pet was cloned, a cat named CC (for “carbon copy” or “copy
cat”; see photo at the beginning of the chapter). The cloning of
mammals provides the potential for many practical applications.
With regard to livestock, cloning would enable farmers to use
cells from their best individuals to create genetically homoge-
neous herds. This could be advantageous in terms of agricultural
yield, although such a genetically homogeneous herd may be
more susceptible to certain diseases. However, people have be-
come greatly concerned with the possibility of human cloning.
This prospect has raised serious ethical questions. Within the past F I G URE 1 . 2 The cloning of a mammal. The lamb in the front
few years, legislation has been introduced that involves bans on is Dolly, the first mammal to be cloned. She was cloned from the cells
human cloning. of a Finn Dorset (a white-faced sheep). The sheep in the back is Dolly’s
Finally, genetic technologies provide the means to modify surrogate mother, a Blackface ewe. A description of how Dolly was
the traits of animals and plants in ways that would have been ­produced is presented in Chapter 22.
­unimaginable just a few decades ago. Figure 1.3a illustrates © Roslin Institute/Phototake

a striking example in which scientists introduced a gene from CONCEPT CHECK: What ethical issues may be associated with human cloning?

(a) GFP expressed in mice
GFP
(b) GFP expressed in the gonads of a male mosquito
FI GURE 1.3 The introduction of a jellyfish gene into
­laboratory mice and mosquitoes. (a) A gene that naturally occurs
in jellyfish encodes a protein called green fluorescent protein (GFP).
The GFP gene was cloned and introduced into mice. When these mice
are exposed to UV light, GFP emits a bright green color. These mice
glow green, just like the jellyfish! (b) The GFP gene was introduced
next to a gene sequence that causes the expression of GFP only in the
gonads of male mosquitoes. This allows researchers to identify and
sort males from females.
(a): © Advanced Cell Technology, Inc., Worcester, Massachusetts; (b): Photo taken by Flaminia
Catteruccia, Jason Benton and Andrea Crisanti, and assembled by www.luciariccidesign.com
CONCEPT CHECK: Why is it useful to sort male mosquitoes from females?
jellyfish into mice. Certain species of jellyfish emit a “green
glow” produced by a gene that encodes a bioluminescent protein
called green fluorescent protein (GFP). When exposed to blue or
ultraviolet (UV) light, the protein emits a striking green-colored
light. Scientists were able to clone the GFP gene from a sample
of jellyfish cells and then introduce this gene into laboratory
mice. The green fluorescent protein is made throughout the cells
of their bodies. As a result, their skin, eyes, and organs give off
an eerie green glow when exposed to UV light. Only their fur
does not glow.
The expression of green fluorescent protein allows research-
ers to identify particular proteins in cells or specific body parts.
1.1 THE MOLECULAR EXPRESSION OF GENES 3
For example, Andrea Crisanti and colleagues have altered mosqui-
toes to express GFP only in the gonads of males (Figure 1.3b).
This enables the researchers to identify and sort males from fe-
males. Why is this useful? Researchers can produce a population
of mosquitoes and then sterilize the males. The ability to rapidly
sort males and females makes it possible to release the sterile
males without the risk of releasing additional females. The release
of sterile males may be an effective means of controlling mosquito
populations because females mate only once before they die.
­Mating with a sterile male prevents a female from producing off-
spring. In 2008, Osamu Shimomura, Martin Chalfie, and Roger
Tsien received the Nobel Prize in chemistry for the discovery and
the development of GFP, which has become a widely used tool
in biology.
Overall, as we move forward in the twenty-first century, the
excitement level in the field of genetics is high, perhaps higher
than it has ever been. Nevertheless, new genetic knowledge and
technologies will also create many ethical and societal challenges.
In this chapter, we begin with an overview of genetics and then
explore the various fields of genetics and their experimental
­approaches.
1.1 THE MOLECULAR
EXPRESSION OF GENES
Learning Outcomes:
1. Describe the biochemical composition of cells.
2. Explain how proteins are largely responsible for cell struc-
ture and function.
3. Outline how DNA stores the information to make proteins.
Genetics is the branch of biology that deals with heredity and
variation. It stands as the unifying discipline in biology by
­allowing us to understand how life can exist at all levels of
­complexity, ranging from the molecular to the population level.
Genetic variation is the root of the natural diversity that we
­observe among members of the same species and among differ-
ent species.
Genetics is centered on the study of genes. A gene is classi-
cally defined as a unit of heredity. At the molecular level, a gene
is a segment of DNA that produces a functional product. The func-
tional product of most genes is a polypeptide, which is a linear
sequence of amino acids that folds into units that constitute pro-
teins. In addition, genes are commonly described according to the
way they affect traits, which are the characteristics of an organ-
ism. In humans, for example, we speak of traits such as eye color,
hair texture, and height. The ongoing theme of this textbook is the
relationship between genes and traits. As an organism grows and
develops, its collection of genes provides a blueprint that deter-
mines its traits.
In this section, we examine the general features of life,
beginning with the molecular level and ending with popula-
tions of organisms. As will become apparent, genetics is the
4 C H A P T E R 1 :: OVERVIEW OF GENETICS Plant cell

common thread that explains the existence of life and its conti-
nuity from generation to generation. For most students, this
chapter should serve as an overview of topics they have learned
in other introductory courses such as General Biology. Even so,
it is usually helpful to see the “big picture” of genetics before
delving into the finer details that are covered in Chapters 2
through 29.
Nucleus

Living Cells Are Composed of Biochemicals

To fully understand the relationship between genes and traits, we
need to begin with an examination of the composition of living
organisms. Every cell is constructed from intricately organized
chemical substances. Small organic molecules such as glucose
and amino acids are produced from the linkage of atoms via
chemical bonds. The chemical properties of organic molecules
are essential for cell vitality in two key ways. First, the breaking
of chemical bonds during the degradation of small molecules pro- Chromosome
vides energy to drive cellular processes. A second important
function of these small organic molecules is their role as the Proteins
building blocks for the synthesis of larger molecules. Four impor-
tant categories of larger molecules are nucleic acids (i.e., DNA
and RNA), proteins, carbohydrates, and lipids. Three of these—
nucleic acids, proteins, and carbohydrates—form macromolecules
that are composed of many repeating units of smaller building
blocks. RNA, proteins, and some carbohydrates are made from DNA
hundreds or even thousands of repeating building blocks. DNA is
the largest macromolecule found in living cells. A single DNA
molecule can be composed of a linear sequence of hundreds of
millions of building blocks called nucleotides!
The formation of cellular structures relies on the interac-
tions of molecules and macromolecules. For example, nucleo-
tides are connected together to make DNA, which is a constituent
of chromosomes (Figure 1.4). In addition, DNA is associated
with many proteins that provide organization to the structure of
chromosomes. Within a eukaryotic cell, the chromosomes are Nucleotides
contained in a compartment called the cell nucleus. The nucleus
is bounded by a double membrane composed of lipids and pro-
teins that shields the chromosomes from the rest of the cell. The NH2
organization of chromosomes within a cell nucleus protects the Cytosine N
H Guanine
chromosomes from mechanical damage and provides a single O
O– H
compartment for genetic activities such as gene transcription. As O N H
N
N
O P O CH2 H
a general theme, the formation of large cellular structures arises O–
O O– H2 N N
N
from interactions among different molecules and macromole- H
H H
H O P O CH2
O
cules. These cellular structures, in turn, are organized to make a OH H
O–
H H
H H
complete living cell.
OH H

Each Cell Contains Many Different
Proteins That Determine Cell Structure
and Function
To a great extent, the characteristics of a cell depend on the types
of proteins that it makes. The entire collection of proteins that a
cell makes at a given time is called its proteome. The range of
functions among different types of proteins is truly remarkable.
Some proteins help determine the shape and structure of a given
F I G URE 1 . 4 Molecular organization of a living cell. Cellular
structures are constructed from smaller building blocks. In this example,
DNA is formed from the linkage of nucleotides to produce a very long
macromolecule. The DNA associates with proteins to form a chromosome.
The chromosomes are located within a membrane-bound organelle called
the nucleus, which, along with many different types of organelles, is
found within a complete cell.
photo: © Biophoto Associates/Science Source
CONCEPT CHECK: Is DNA a small molecule, a macromolecule, or an organelle?
 1.1 THE MOLECULAR EXPRESSION OF GENES 5

cell. For example, the protein known as tubulin assembles into

large structures known as microtubules, which provide the cell
with internal structure and organization. Other proteins are in-
serted into cell membranes and aid in the transport of ions and
small molecules across the membrane. Enzymes, which acceler-
ate chemical reactions, are a particularly important category of
proteins. Some enzymes play a role in the breakdown of molecules
or macromolecules into smaller units. These are known as cata-
bolic enzymes and are important in the utilization of energy.
­Alternatively, anabolic enzymes and accessory proteins function
in the synthesis of molecules and macromolecules throughout the
cell. The construction of a cell greatly depends on its proteins that
are involved in anabolism because these are required to synthesize
all cellular macromolecules.
Molecular biologists have come to realize that the functions
of proteins underlie the cellular characteristics of every organism.
At the molecular level, proteins can be viewed as the active par-
ticipants in the enterprise of life.

DNA Stores the Information for Protein Synthesis

The genetic material of living organisms is composed of a sub-
stance called deoxyribonucleic acid, abbreviated DNA. The
DNA stores the information needed for the synthesis of all cellular
proteins. In other words, the main function of the genetic blueprint
is to code for the production of proteins in the correct cell, at the
proper time, and in suitable amounts. This is an extremely compli-
cated task because living cells make thousands of different pro-
teins. Genetic analyses have shown that a typical bacterium can
make a few thousand different proteins, and estimates for the
­numbers produced by complex eukaryotic species range in the
tens of thousands.
DNA’s ability to store information is based on its structure.
DNA is composed of a linear sequence of nucleotides. Each
nucleotide contains one of four nitrogen-containing bases: ade- F I G URE 1 . 5 A micrograph of the 46 chromosomes found in a
nine (A), thymine (T), guanine (G), or cytosine (C). The linear cell from a ­human male.
order of these bases along a DNA molecule contains information © CNRI/Science Source

similar to the way that groups of letters of the alphabet represent CONCEPT CHECK: Which types of macromolecules are found in chromosomes?
words. For example, the “meaning” of the sequence of bases
ATGGGCCTTAGC differs from that of TTTAAGCTTGCC.
DNA sequences within most genes contain the information to
as a karyotype. The DNA of an average human chromosome is an
direct the order of amino acids within polypeptides according to
extraordinarily long, linear, double-stranded structure that con-
the genetic code. In the code, a three-base sequence specifies
tains well over a hundred million nucleotides. Along the immense
one particular amino acid among the 20 possible choices. One
length of a chromosome, the genetic information is parceled into
or more polypeptides form a functional protein. In this way, the
functional units known as genes. An average-sized human chro-
DNA can store the information to specify the proteins made by
mosome is expected to contain about 1000 different protein-­
an organism.
encoding genes.
DNA Sequence Amino Acid Sequence

ATG GGC CTT AGC Methionine Glycine Leucine Serine The Information in DNA Is Accessed During
TTT AAG CTT GCC Phenylalanine Lysine Leucine Alanine the Process of Gene Expression
To synthesize its proteins, a cell must be able to access the informa-
In living cells, the DNA is found within large structures known as tion that is stored within its DNA. The process of using a gene se-
chromosomes. Figure 1.5 is a micrograph of the 46 chromosomes quence to affect the characteristics of cells and organisms is referred
contained in a cell from a human male; this type of image is known to as gene expression. At the molecular level, the information
6 C H A P T E R 1 :: OVERVIEW OF GENETICS
DNA
Gene
Transcription
RNA (messenger RNA)
Translation
Protein
(sequence of
amino acids)
Functioning of proteins within living
cells influences an organism’s traits.
FI GURE 1.6 Gene expression at the molecular
level. The ­expression of a gene is a multistep process.
During transcription, one of the DNA strands is used as a
template to make an RNA strand. During translation, the
RNA strand is used to specify the sequence of amino ­acids within a
polypeptide. One or more polypeptides produce a protein that functions
within the cell, thereby influencing an organism’s traits.
CONCEPT CHECK: Where is the information to make a polypeptide stored?
within genes is accessed in a stepwise process (Figure 1.6). In the
first step, known as transcription, the DNA sequence within a gene
is copied into a nucleotide sequence of ribonucleic acid (RNA).
Protein-encoding genes (also called structural genes) carry the
information for the amino acid sequence of a polypeptide. When a
protein-encoding gene is transcribed, the first product is an RNA
molecule known as messenger RNA (mRNA). During polypeptide
synthesis—a process called translation—the sequence of nucleo-
tides within the mRNA determines the sequence of amino acids in a
polypeptide. One or more polypeptides then fold and assemble into
a functional protein. The synthesis of functional proteins ultimately
determines an organism’s traits. As discussed further in Chapter 12
(look ahead to Figure 12.1), the pathway of gene expression from
DNA to RNA to protein is called the central dogma of genetics
(also called the central dogma of molecular biology). It forms a
cornerstone of our understanding of genetics at the molecular level.
1.1 COMPREHENSION QUESTIONS
1. Which of the following is not a constituent of a cell’s proteome?
a. An enzyme
b. A cytoskeletal protein
c. A transport protein in the plasma membrane
d. An mRNA
2. A gene is a segment of DNA that has the information to produce
a functional product. The functional product of most genes is
a. DNA.
b. mRNA.
c. a polypeptide.
d. all of the above.
3. The function of the genetic code is to
a. promote transcription.
b. specify the amino acids within a polypeptide.
c. alter the sequence of DNA.
d. none of the above.
4. The process of transcription directly results in the synthesis of
a. DNA.
b. RNA.
c. a polypeptide.
d. all of the above.
1.2 THE RELATIONSHIP
BETWEEN GENES AND TRAITS
Learning Outcomes:
1. Outline how the expression of genes leads to an organism’s
traits.
2. Define genetic variation.
3. Discuss the relationship between genes and traits.
4. Describe how genes are transmitted in sexually reproducing
species.
5. Explain the process of evolution.
A trait is any characteristic that an organism displays. In genetics,
we often focus our attention on morphological traits, those that
affect the appearance, form, and structure of an organism. The
color of a flower and the height of a pea plant are morphological
traits. Geneticists frequently study these types of traits because
they are easy to evaluate. For example, an experimenter can sim-
ply look at a plant and tell if it has red or white flowers. However,
not all traits are morphological. Physiological traits affect the
ability of an organism to function. For example, the rate at which
a bacterium metabolizes a sugar such as lactose is a physiological
trait. Like morphological traits, physiological traits are con-
trolled, in part, by the expression of genes. Behavioral traits
­affect the ways an organism responds to its environment. An ex-
ample is the mating calls of bird species. In animals, the nervous
system plays a key role in governing such traits. In this section,
we will examine the relationship between the expression of genes
and an organism’s traits.
The Molecular Expression of Genes Leads
to an Organism’s Traits
A complicated, yet very exciting, aspect of genetics is that our
observations and theories span four levels of biological organization:
 1.2 THE RELATIONSHIP BETWEEN GENES AND TRAITS 7

molecules, cells, organisms, and populations. This can make it

­difficult to appreciate the relationship between genes and traits.
To understand this connection, we need to relate the following
phenomena: Pigmentation gene Pigmentation gene
(dark allele) (light allele)
1. Genes are expressed at the molecular level. In other found in a dark butterfly found in a light butterfly
words, gene transcription and translation lead to the Transcription and translation
­production of a particular protein, which is a molecular
process.
2. Proteins often function at the cellular level. The function
of a protein within a cell affects the structure and workings
of that cell.
3. An organism’s traits are determined by the characteristics Highly functional Poorly functional
of its cells. We do not have microscopic vision, yet when pigmentation enzyme pigmentation enzyme
we view morphological traits, we are really observing the
properties of an individual’s cells. For example, a red (a) Molecular level
flower has its color because its cells make a red pigment.
The trait of red flower color is an observation at the
­organism level. Yet the trait is rooted in the molecular Pigment
characteristics of the organism’s cells. molecule
4. A species is a group of organisms that maintains a
Wing cells
­distinctive set of attributes in nature. The occurrence of a
trait within a species is an observation at the population
level. Along with learning how a trait occurs, we also Lots of pigment made Little pigment made
want to ­understand why a trait becomes prevalent in a
particular species. In many cases, researchers discover (b) Cellular level
that a trait ­predominates within a population because
it promotes the reproductive success of the members
of the population. This leads to the evolution of
­beneficial traits.
As a schematic example to illustrate the four levels of genetics,
Figure 1.7 shows the trait of pigmentation in butterflies. One
member of this species is dark-colored and the other is very light. Dark butterfly Light butterfly
Let’s consider how we can explain this trait at the molecular,
­cellular, organism, and ­population levels. (c) Organism level
At the molecular level, we need to understand the nature of
the gene or genes that govern this trait. As shown in Figure 1.7a, a
gene, which we will call the pigmentation gene, is responsible for
the amount of pigment produced. The pigmentation gene exists in
two different versions. Alternative versions of a specific gene are
called alleles. In this example, one allele confers a dark pigmenta-
tion and the other causes a light pigmentation. Each of these al-
leles encodes a protein that functions as a pigment-synthesizing
enzyme. However, the DNA sequences of the two alleles differ
slightly from each other. This difference in the DNA sequence
leads to a variation in the structure and function of the respective
pigmentation enzymes.
At the cellular level (Figure 1.7b), the functional differences
Dark butterflies are usually Light butterflies are usually
between the two pigmentation enzymes affect the amount of pig- in forested regions. in unforested regions.
ment produced. The allele causing dark pigmentation, which is
shown on the left, encodes an enzyme that functions very well. (d) Population level
Therefore, when this gene is expressed in the cells of the wings,
a large amount of pigment is made. By comparison, the allele F I G URE 1 . 7 The relationship between genes and traits at the
causing light pigmentation encodes an enzyme that functions (a) molecular, (b) cellular, (c) organism, and (d) population levels.
poorly. Therefore, when this allele is the only pigmentation gene CONCEPT CHECK: Which butterfly has a more active pigment-producing
expressed, little pigment is made. ­enzyme, the dark- or light-colored one?
8 C H A P T E R 1 :: OVERVIEW OF GENETICS

At the organism level (Figure 1.7c), the amount of pig-

ment in the wing cells governs the color of the wings. If the
pigment cells produce high amounts of pigment, the wings are
dark-colored. If the pigment cells produce little pigment, the
wings are light.
Finally, at the population level (Figure 1.7d), geneticists
would like to know why a species of butterfly would contain some
members with dark wings and other members with light wings.
One possible explanation is differential predation. The butterflies
with dark wings might avoid being eaten by birds if they happen to
live within the dim light of a forest. The dark wings would help to
camouflage the butterfly if it were perched on a dark surface such
as a tree trunk. In contrast, the lightly colored wings would be an
advantage if the butterfly inhabited a brightly lit meadow. Under
these conditions, a bird may be less likely to notice a light-colored
butterfly that is perched on a sunlit surface. A population geneti-
cist might study this species of butterfly and find that the dark-
colored members usually live in forested areas and the light-colored
members reside in unforested regions.

Inherited Differences in Traits Are

Due to Genetic Variation
In Figure 1.7, we considered how gene expression leads to vari- F I G URE 1 . 8 Two dyeing poison frogs (Dendrobates tinctorius)
ation in a trait of organisms, using the example of dark- versus showing different morphs within a single species.
light-colored wings in butterflies. Variation in traits among (Top): © Mark Smith/Science Source; (Bottom): © Dante Fenolio/Science Source

members of the same species is very common. For example,
some people have brown hair and others have blond hair; some
petunias have white flowers and others have purple flowers.
These are examples of genetic variation. This term describes
the differences in inherited traits among individuals within
a population.
In large populations that occupy a wide geographic range,
genetic variation can be quite striking. Morphological differ-
ences have often led geneticists to misidentify two members of
the same species as belonging to separate species. As an exam-
ple, Figure 1.8 shows two dyeing poison frogs that are members
of the same species, Dendrobates tinctorius. They display dra-
matic differences in their markings. Such contrasting forms
within a single species are termed morphs. You can easily imag-
ine how someone might m ­ istakenly conclude that these frogs are
not members of the same species.
Changes in the nucleotide sequence of DNA underlie the
genetic variation that we see among individuals. Throughout this
textbook, we will routinely examine how variation in the genetic
material results in changes in an organism’s traits. At the molecu-
lar level, genetic variation can be attributed to different types of
modifications.
1. Small or large differences can occur within gene sequences.
When such changes initially occur, they are called gene
mutations. Mutations result in genetic variation in which
a gene is found in two or more alleles, as previously
described in Figure 1.7. In many cases, gene mutations
alter the expression or function of a protein that a gene
­encodes.
CONCEPT CHECK: Why do these two frogs look so different?
2. Major alterations can also occur in the structure of a
­chromosome. A large segment of a chromosome can be
lost, rearranged, or reattached to another chromosome.
3. Variation may also occur in the total number of chromo-
somes. In some cases, an organism may inherit one too
many or one too few chromosomes. In other cases, it may
inherit an extra set of chromosomes.
Variations of sequences within genes are a common source of ge-
netic variation among members of the same species. In humans,
familiar examples of variation involve genes for eye color, hair
texture, and skin pigmentation. Chromosome variation—a change
in chromosome structure or number (or both)—is also found, but
this type of change is often detrimental. Many human genetic dis-
orders are the result of chromosomal alterations. The most com-
mon example is Down syndrome, which is due to the presence of
an extra chromosome (Figure 1.9a). By comparison, chromosome
variation in plants is common and often leads to plants with supe-
rior characteristics, such as increased resistance to disease. Plant
breeders have frequently exploited this observation. Cultivated
varieties of wheat, for example, have many more chromosomes
than the wild species (Figure 1.9b).
Traits Are Governed by Genes
and by the Environment
In our discussion thus far, we have considered the role that genes
play in determining an organism’s traits. Another critical factor is

(a) (b)
FI GURE 1.9 Examples of chromosome variation. (a) A person
with Down syndrome. She has 47 chromosomes rather than the common
number of 46, because she has an extra copy of chromosome 21. (b) A
wheat plant. Cultivated wheat is derived from the contributions of three
wild species with two sets of chromosomes each, producing an organism
with six sets of chromosomes.
(a): © Stockbyte/Alamy RF; (b): © Brand X Pictures/PunchStock RF
CONCEPT CHECK: Do these examples constitute variation in chromosome
structure or variation in chromosome number?
the environment—the surroundings in which an organism ex-
ists. A variety of factors in an organism’s environment pro-
foundly affect its morphological and physiological features. For
example, a person’s diet greatly influences many traits such
as height, weight, and even intelligence. Likewise, the amount
of sunlight a plant receives affects its growth rate and the color
of its flowers.
An interesting example of the interplay between genes and
the environment involves the human genetic disease phenylke-
tonuria (PKU). Humans have a gene that encodes an enzyme
known as phenylalanine hydroxylase. Most people have two
functional copies of this gene. People with one or two func-
tional copies of the gene can eat foods containing the amino
acid phenylalanine and metabolize it properly. A rare variation
in the gene that encodes phenylalanine hydroxylase results in a
nonfunctional version of this enzyme. Individuals with two cop-
ies of this rare, inactive allele cannot metabolize phenylalanine
properly. When given a standard diet containing phenylalanine,
individuals with this disorder are unable to break down this
amino acid. Phenylalanine accumulates and is converted into
phenylketones, which are detected in the urine. Individuals with
PKU can manifest a variety of detrimental traits, including men-
tal impairment, underdeveloped teeth, and foul-smelling urine.
Fortunately, through routine newborn screening in the United
States, PKU is now diagnosed early. Part of the treatment is a
diet that restricts phenylalanine, which is present in high-­protein
foods such as eggs, meat, and dairy products. Restricting phe-
nylalanine allows the affected child to develop normally.
PKU provides a dramatic example of how the environment and
1.2 THE RELATIONSHIP BETWEEN GENES AND TRAITS 9
an individual’s genes can interact to influence the traits of the
organism.
During Reproduction, Genes Are Passed
from Parent to Offspring
Now that we have considered how genes and the environment
govern the outcome of traits, we can turn to the issue of inheritance.
How are traits passed from parents to offspring? The foundation
for our understanding of inheritance came from Gregor Mendel’s
study of pea plants in the nineteenth century. His work revealed
that the genetic determinants that govern traits, which we now call
genes, are passed from parent to offspring as discrete units. We can
predict the outcome of many genetic crosses based on Mendel’s
laws of inheritance.
The inheritance patterns identified by Mendel can be ex-
plained by the existence of chromosomes and their behavior dur-
ing cell division. Like Mendel’s pea plants, sexually reproducing
species are commonly diploid. This means they contain two
copies of each chromosome, one from each parent. The two cop-
ies are called homologs of each other. Because genes are located
within chromosomes, diploid organisms have two copies of most
genes. Humans, for example, have 46 chromosomes, which are
found in homologous pairs (Figure 1.10a). With the exception
of the sex chromosomes (X and Y), each homologous pair con-
tains the same kinds of genes. For example, both copies of ­human
chromosome 12 carry the gene that encodes phenylalanine
­hydroxylase, which was discussed previously. Therefore, an indi-
vidual has two copies of this gene, which may or may not be
identical alleles.
Most cells of the human body that are not directly involved
in sexual reproduction contain 46 chromosomes. These cells are
called somatic cells. In contrast, the gametes—sperm and egg
cells—contain half that number (23) and are termed haploid
­(Figure 1.10b). The union of gametes during fertilization re-
stores the diploid number of chromosomes. The primary advan-
tage of sexual reproduction is that it enhances genetic variation.
For example, a tall person with blue eyes and a short person with
brown eyes may have short offspring with blue eyes or tall off-
spring with brown eyes. Therefore, sexual reproduction can re-
sult in new combinations of two or more traits that differ from
those of either parent.
The Genetic Composition of a Species Evolves over
the Course of Many Generations
As we have just seen, sexual reproduction has the potential to
enhance genetic variation. This can be an advantage for a popu-
lation of individuals as they struggle to survive and compete
within their natural environment. The term biological evolu-
tion, or simply, evolution, refers to the phenomenon that the
genetic makeup of a population changes from one generation to
the next.
As suggested by Charles Darwin, the members of a species
are in competition with one another for essential resources.
Random genetic changes (i.e., mutations) occasionally occur
­
10 C H A P T E R 1 :: OVERVIEW OF GENETICS

1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8

9 10 11 12 13 14 15 16 9 10 11 12 13 14 15 16

17 18 19 20 21 22 XX 17 18 19 20 21 22 X

(a) Chromosomal composition found (b) Chromosomal composition found in

in human somatic cells of females a human gamete (23 chromosomes)
(46 chromosomes)

FI G U RE 1.1 0 The complement of human chromosomes in somatic cells and gametes. (a) A schematic drawing of the 46 chromosomes
of a human. With the exception of the sex chromosomes, these are always found in homologous pairs. (b) The chromosomal composition of a gamete,
which contains only 23 chromosomes, one from each pair. This gamete contains an X chromosome. Half of the gametes from human males contain a
Y chromosome instead of the X chromosome.
CONCEPT CHECK: The leaf cells of a corn plant contain 20 chromosomes each. How many chromosomes are found in a gamete made by a corn plant?

within an individual’s genes, and sometimes these changes lead to

a modification of traits that promote reproductive success. For 1.2 COMPREHENSION QUESTIONS
example, over the course of many generations, random gene muta- 1. Gene expression can be viewed at which of the following levels?
tions have lengthened the snout and extended the tongue of the
a. Molecular and cellular levels
anteater, enabling it to probe into the ground and feed on ants.
When a mutation creates a new allele that is beneficial, the allele b. Organism level
may become prevalent in future generations because the individu- c. Population level
als carrying the allele are more likely to reproduce and pass the d. All of the above
beneficial ­allele to their offspring. This process is known as 2. Variation in the traits of organisms may be attributable to
­natural selection. In this way, a species becomes better adapted to a. gene mutations.
its environment. b. alterations in chromosome structure.
Over a long period of time, the accumulation of many
c. variation in chromosome number.
genetic changes may lead to rather striking modifications in a
species’ characteristics. As an example, Figure 1.11 depicts d. all of the above.
the evolution of the modern-day horse. Over time, a variety of 3. A human skin cell has 46 chromosomes. A human sperm cell has
morphological changes occurred, including an increase in a. 23.
size, fewer toes, and modified jaw structure. The changes can b. 46.
be attributed to natural selection. Over North America, where c. 92.
much of horse evolution occurred, large areas of dense forests d. none of the above.
were replaced with grasslands. The increase in size and
4. Evolutionary change caused by natural selection results in
changes in foot structure enabled horses to escape predators
species with
more easily and travel greater distances in search of food.
­Natural selection favored the changes seen in horses’ teeth, a. greater complexity.
because such changes allowed them to eat grasses and other b. less complexity.
types of vegetation that are more abrasive and require more c. greater reproductive success in their native environment.
chewing. d. the ability to survive longer.
 1.3 FIELDS OF GENETICS 11

Equus
0 Hippidium and other genera

Nannippus
Stylohipparion
5 Pliohippus

Hipparion Neohipparion

10

Sinohippus Megahippus
Calippus

Archaeohippus
20
Merychippus
Millions of years ago (mya)

Anchitherium Hypohippus

Parahippus

Miohippus

Mesohippus
40

Paleotherium Epihippus

Propalaeotherium

Pachynolophus Orohippus

55
Hyracotherium

FI GURE 1.11 The evolutionary changes that led to the modern horse genus, Equus. Three important morphological changes that occurred
were larger size, fewer toes, and a shift toward a jaw structure suited for grazing.
CONCEPT CHECK: According to the theory of evolution, why have these changes occurred in horse populations over the course of many generations?

as biochemistry, biophysics, cell biology, mathematics, microbiol-

1.3 FIELDS OF GENETICS
Learning Outcome:
1. Compare and contrast the three major fields of genetics:
transmission, molecular, and population genetics.
Genetics is a broad discipline encompassing molecular, cellular,
organism, and population biology. Many scientists who are inter-
ested in genetics have been trained in supporting disciplines such
ogy, population biology, ecology, agriculture, and medicine.
­Experimentally, geneticists often focus their efforts on model
­organisms—organisms studied by many different researchers so
they can compare their results and determine scientific principles
that apply more broadly to other species. Figure 1.12 shows some
common examples, including Escherichia coli (a bacterium), Sac-
charomyces cerevisiae (a yeast), Drosophila melanogaster (fruit
fly), Caenorhabditis elegans (a nematode worm), Mus musculus
(mouse), and Arabidopsis thaliana (a flowering plant). Model
12 C H A P T E R 1 :: OVERVIEW OF GENETICS
0.3 μm
(a) Escherichia coli (b) Saccharomyces cerevisiae
156 μm
(d) Caenorhabditis elegans (e) Mus musculus
FI G U RE 1.1 2 Examples of model organisms studied by geneticists. (a) Escherichia coli (a bacterium), (b) Saccharomyces cerevisiae (a yeast),
(c) Drosophila melanogaster (fruit fly), (d) Caenorhabditis elegans (a nematode worm), (e) Mus musculus (mouse), and (f) Arabidopsis thaliana
(a flowering plant).
(a): Peggy S. Hayes & Elizabeth H. White, M.S./CDC; (b): © SCIMAT/Science Source; (c): © Steve Hopkin/ardea.com; (d): © Sinclair Stammers/Science Source; (e): © J-M. Labat/Science
Source; (f): © WILDLIFE GmbH/Alamy
CONCEPT CHECK: Can you think of another example of a model organism?
o­ rganisms offer experimental advantages over other species. For
example, E. coli is a very simple organism that can be easily grown
in the laboratory. By limiting their work to a few such model or-
ganisms, researchers can more easily unravel the genetic mecha-
nisms that govern the traits of a given species. Furthermore, the
genes found in model organisms often function in a similar way to
those found in humans.
The study of genetics has been traditionally divided into
three areas—transmission, molecular, and population genetics—
although overlap is found among them. In this section, we will
examine the three fields of genetics.
Transmission Genetics Explores the Inheritance
Patterns of Traits as They Are Passed from Parents
to Offspring
A scientist working in the field of transmission genetics examines
the relationship between the transmission of genes from parent to
offspring and the outcome of the offspring’s traits. For example,
how can two brown-eyed parents produce a blue-eyed child? Or
why do tall parents tend to produce tall children, but not always?
Our modern understanding of transmission genetics began with
the studies of Gregor Mendel. His work provided the conceptual
framework for transmission genetics. In particular, he originated
the idea that factors, which we now call genes, are passed as dis-
crete units from parents to offspring via sperm and egg cells. Since
7 μm
(c) Drosophila melanogaster
(f) Arabidopsis thaliana
Mendel’s pioneering studies in the 1860s, our knowledge of ge-
netic transmission has skyrocketed. Many patterns of genetic
transmission are more complex than the simple Mendelian pat-
terns that are described in Chapter 2. The additional complexities
of transmission genetics are examined in Chapters 3 through 8.
Experimentally, the fundamental approach of a transmission
geneticist is the genetic cross—the breeding of two selected indi-
viduals and the subsequent analysis of their offspring in an attempt
to understand how traits are passed from parents to offspring. In
the case of experimental organisms, the researcher chooses two
parents with particular traits and then categorizes the offspring
according to the traits they possess. In many cases, this analysis is
quantitative in nature. For example, an experimenter may cross
two tall pea plants and obtain 100 offspring that fall into two cat-
egories: 75 tall and 25 dwarf. As we will see in Chapter 2, the ratio
of tall to dwarf offspring provides important information concern-
ing the inheritance pattern of the height trait.
Molecular Genetics Focuses on a Biochemical
Understanding of the Hereditary Material
The goal of molecular genetics, as the name of the field implies, is
to understand how the genetic material works at the molecular
level. In other words, molecular geneticists want to understand
the molecular features of DNA and how these features underlie the
expression of genes. The experiments of molecular geneticists are

usually conducted within the confines of a laboratory. Their ef-
forts frequently progress to a detailed analysis of DNA, RNA, and
proteins, using a variety of techniques that are described through-
out Parts III, IV, and V of this textbook.
Molecular geneticists often study mutant genes that have
abnormal function. This is called a genetic approach to the
study of a research question. In many cases, researchers analyze
the effects of gene mutations that eliminate the function of a
gene. This is called a loss-of-function mutation, and the result-
ing version of the gene is called a loss-of-function allele. By
studying the effects of such mutations, the role of the functional,
nonmutant gene is often revealed. For example, let’s suppose
that a particular plant species produces purple flowers. If a loss-
of-function mutation within a given gene causes a plant of that
species to produce white flowers, one would suspect that the
role of the functional gene involves the production of purple
pigmentation.
Studies within molecular genetics interface with other
­disciplines such as biochemistry, biophysics, and cell biology. In
addition, advances within molecular genetics have shed consider-
able light on the areas of transmission and population genetics. Our
quest to understand molecular genetics has spawned a variety of
modern molecular technologies and computer-based approaches.
Furthermore, discoveries within molecular genetics have had wide-
spread applications in agriculture, medicine, and biotechnology.
Population Genetics Is Concerned with Genetic
Variation and Its Role in Evolution
The foundations of population genetics arose during the first few
decades of the twentieth century. Although many scientists of this
era did not accept the findings of Mendel or Darwin, the theories
of population genetics provided a compelling way to connect the
two viewpoints. Mendel’s work and that of many succeeding ge-
neticists gave insight into the nature of genes and how they are
transmitted from parents to offspring. The work of Darwin pro-
vided a natural explanation for the variation in characteristics ob-
served among the members of a species. To relate these two
phenomena, population geneticists have developed mathematical
theories to explain the prevalence of certain alleles within popula-
tions of individuals. This work helps us understand how processes
such as natural selection have resulted in the prevalence of indi-
viduals that carry particular alleles.
Population geneticists are particularly interested in genetic
variation and how that variation is related to an organism’s envi-
ronment. In this field, the frequencies of alleles within a popula-
tion are of central importance.
1.3 COMPREHENSION QUESTIONS
1. Which of the following is not a model organism?
a. Mus musculus (laboratory mouse)
b. Escherichia coli (a bacterium)
c. Saccharomyces cerevisiae (a yeast)
d. Sciurus carolinensis (gray squirrel)
1.4 THE SCIENCE OF GENETICS 13
2. A person studying the rate of transcription of a particular gene is
working in the field of
a. molecular genetics.
b. transmission genetics.
c. population genetics.
d. None of the above are correct.
1.4 THE SCIENCE OF GENETICS
Learning Outcomes:
1. Discuss how genetics is an experimental science.
2. Outline different strategies for solving problems in genetics.
Science is a way of knowing about our natural world. The science
of genetics allows us to understand how the expression of our
genes produces the traits that we possess. In this section, we will
consider how scientists attempt to answer questions via experi-
mentation. We will also consider general approaches for solving
problems.
Genetics Is an Experimental Science
Researchers typically follow two general types of scientific ap-
proaches: hypothesis testing and discovery-based science. In
­hypothesis testing, also called the scientific method, scientists
follow a series of steps to reach verifiable conclusions about the
world. Although scientists arrive at their theories in different
ways, the scientific method provides a way to validate (or invali-
date) a particular hypothesis. Alternatively, research may also in-
volve the collection of data without a preconceived hypothesis.
For example, researchers might analyze the genes found in cancer
cells to identify those that have become mutant. In this case, the
scientists may not have a hypothesis about which particular genes
may be involved. The collection and analysis of data without the
need for a preconceived hypothesis is called discovery-based
­science or, simply, discovery science.
In traditional science textbooks, the emphasis often lies on
the product of science. That is, many textbooks are aimed primar-
ily at teaching the student about the observations scientists have
made and the hypotheses they have proposed to explain those ob-
servations. Along the way, the student is provided with many bits
and pieces of experimental techniques and data. Likewise, this
textbook also provides you with many observations and hypothe-
ses. However, it attempts to go one step further. Most chapters
contain one or two experiments that have been “dissected” into
five individual components to help you to understand the entire
scientific process:
1. Background information is provided so that you can appre-
ciate earlier observations that were known prior to conduct-
ing the experiment.
2. Most experiments involve hypothesis testing. In those
cases, the figure presenting the experiment states the
­hypothesis the scientists were trying to test. In other
14 C H A P T E R 1 :: OVERVIEW OF GENETICS
words, what scientific question was the researcher trying
to answer?
3. Next, the figure follows the experimental steps the scientist
took to test the hypothesis. Each featured experiment con-
tains two parallel illustrations labeled Experimental Level
and Conceptual Level. The Experimental Level helps you
to understand the techniques followed. The Conceptual
Level helps you to understand what is actually happening
at each step in the procedure.
4. The raw data for each experiment are then presented.
5. Last, an interpretation of the data is offered within the text.
The rationale behind this approach is that it enables you to see the
experimental process from beginning to end. As you read through
the chapters, the experiments will help you to see the relationship
between science and scientific theories.
As a student of genetics, you will be given the opportunity
to involve your mind in the experimental process. As you are read-
ing an experiment, you may find yourself thinking about different
approaches and alternative hypotheses. Different people can view
the same data and arrive at very different conclusions. As you
progress through the experiments in this book, you will enjoy
­genetics far more if you try to develop your own skills at formulat-
ing hypotheses, designing experiments, and interpreting data.
Also, some of the questions in the problem sets are aimed at refin-
ing these skills.
Finally, it is worthwhile to point out that science is a social
discipline. As you develop your skills at scrutinizing experiments,
it is fun to discuss your ideas with other people, including fellow
students and faculty members. Keep in mind that you do not need
to “know all the answers” before you enter into a scientific discus-
sion. Instead, it is more rewarding to view science as an ongoing
and never-ending dialogue.
Genetic TIPS Will Help You to Improve Your
Problem-Solving Skills
As your progress through this textbook, your learning will involve
two general goals:
∙ You will gather foundational knowledge. In other words,
you will be able to describe core concepts in genetics. For
example, you will be able to explain how DNA replication
occurs and describe the proteins that are involved in this
process.
∙ You will develop problem-solving skills that allow you
to apply that foundational knowledge in different ways.
For example, you will learn how to use statistics to de-
termine if a genetic hypothesis is consistent with experi-
mental data.
The combination of foundational knowledge and problem-­solving
skills will enable you not only to understand genetics, but also to
apply your knowledge in different situations. To help you develop
these skills, Chapters 2 through 29 contain solved problems
named Genetic TIPS, which stands for Topic, Information, and
Problem-solving Strategy. These solved problems follow a con-
sistent pattern.
GENETIC TIPS THE QUESTION: All of the Genetic TIPS
begin with a question. As an example, let’s consider the following
question:
The coding strand of DNA in a segment of a gene is as follows:
ATG GGC CTT AGC. This strand carries the information to make a
region of a polypeptide with the amino acid sequence, methionine-
glycine-leucine-serine. What would be the consequences if a mutation
changed the second cytosine (C) in this sequence to an adenine (A)?
T OPIC: What topic in genetics does this question address? The
topic is gene expression. More specifically, the question is about
the relationship between a gene sequence and the genetic code.
I NFORMATION: What information do you know based on the
question and your understanding of the topic? In the question,
you are given the base sequence of a short segment of a gene and
told that one of the bases has been changed. From your understanding
of the topic, you may remember that a polypeptide sequence is
determined by reading the mRNA (transcribed from a gene) in
groups of three bases called codons.
P ROBLEM-SOLVING S TRATEGY: Compare and contrast.
One strategy to solve this problem is to compare the mRNA
sequence (transcribed from this gene) before and after the mutation:
Original: AUG GGC CUU AGC
Mutant:   AUG GGC AUU AGC
ANSWER: The mutation has changed the sequence of bases in the
mRNA so that the third codon has changed from CUU to AUU.
­Because codons specify amino acids, this may change the third
amino acid to something else. Note: If you look ahead to Chapter 13
(see Table 13.1), you will see that CUU specifies leucine, whereas
AUU specifies isoleucine. Therefore, you would predict that the mu-
tation would change the third amino acid from leucine to isoleucine.
Throughout Chapters 2 through 29, each chapter will contain sev-
eral Genetic TIPS. Some of these will be within the chapter itself
and some will precede the problem sets that are at the end of each
chapter. Though there are many different problem-solving strate-
gies, Genetic TIPS will focus on ten strategies that will help you to
solve problems. You will see these ten strategies over and over
again as you progress through the textbook:
1. Define key terms. In some cases, a question may be difficult
to understand because you don’t know the meaning of one or
more key terms in the question. If so, you will need to begin
your problem-solving by defining such terms, ­either by look-
ing them up in the glossary or by using the index to find the
location in the text where the key terms are explained.
2. Make a drawing. Genetic problems are oftentimes difficult
to solve in your head. Making a drawing may make a big
difference in your ability to see the solution.
3. Predict the outcome. Geneticists may want to predict the out-
come of an experiment. For example, in Chapters 3 through 6,
you will learn about different ways to predict the outcome of
genetic crosses. Becoming familiar with these methods will
help you to predict the outcomes of particular experiments.
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La più gran parte delle pitture antiche erano parietarie, quelle che in
oggi, dai diversi procedimenti adottati nel pingere, chiameremmo
affreschi. Nondimeno anche allora non è escluso che si potesse da
quegli artisti dipingere sull’intonaco recente. Anzi il dottissimo cav.
Minervini ha constatato non dubbie differenze nei diversi sistemi
onde sono condotti gli intonachi che ricoprono le pareti pompejane.
Ne ha distinti di più fini, per i quali, a suo credere, gli antichi
dipingevano a fresco le composizioni meglio accurate, i paesaggi e
le figure; mentre che le semplici decorazioni erano dipinte a secco
da pittori inferiori.
Il più spesso rilevasi evidentemente come suolessero dipingere a
guazzo od all’acquerello con colori preparati a gomma, ovvero con
altro genere di glutine, e se ne adducon le prove in gran numero di
dipinti pompejani esistenti nel Museo di Napoli, i colori de’ quali si
ponno separare e staccare dall’intonaco, talvolta ben anco
vedendosi ricomparire, nel cadere lo strato superiore, la sottoposta
tinta, ciò che non può accadere di pitture frescate. Quando poi si
ponga mente che fra i colori adoperati allora eranvi quelli che
chiamavano purpurissum, indicum, cæruleum, melinum,
auripigmentum, appianum e cerussa, e i quali non resistono alla
calce [325], si mette in sodo che il processo non poteva essere a
fresco.
Uno dei metodi più usitati in quel tempo era del pingere all’encausto,
e se ne valeva assai per le pitture sulle pareti. Più sopra nella
taberna del mercante de’ colori, abbiam trovato molti ingredienti che
servivano a tale scopo. Era l’encaustica l’arte di dipingere con colori
mescolati con cera e di poi induriti coll’azione stessa del fuoco. Di
qual modo si praticasse in antico, di presente più non si sa,
essendosene smarrito il procedimento. Vero è che il conte Caylus
pretese averlo rivendicato e ne scrisse al proposito un trattato; ma
per general sentimento non credasi che sia riuscito ad ottenerne i
medesimi risultamenti. Pare, scrive Rych [326], ch’essi seguissero
diversi metodi e conducessero l’operazione in differentissime guise:
o con colori mescolati con cera, distesi con una spazzola asciutta e
poi bruciati con un cauterio (cauterium), ovvero marcando i contorni
con un ferro rovente (cestrum) si incideva sopra l’avorio; nel qual
processo non pare che la cera entrasse punto, o infine liquefacendo
la cera con cui i colori erano mescolati, cosicchè la spazzola era
intinta nel liquido composto e il colore disteso in istato di fluidità,
come si fa all’acquerello, ma di poi rammorbidito e fuso coll’azion del
calorico.
Contrariamente a quanto afferma Bréton, che l’encausto fosse bensì
in uso allora, ma indubbiamente solo pei quadri e non per le pitture
di semplice decorazione [327], vogliono altri che il metodo
dell’encausto fosse quello adottato in tutte le pitture pompejane,
come in quelle scoperte ad Ercolano. Mengs per altro portava
opinione che esse fossero invece condotte a fresco [328].
Ho poi detto che la più gran parte dei dipinti fossero parietarii, ciò
che significa che non tutti lo fossero; ben sapendosi come alcuna
volta si dipingesse sulle tavole di legno, ch’erano di larice, preparate
prima con uno strato di creta, specie di pietra tenera, friabile e
bianca, sì che tabula equivalesse, senz’altro, a quadro, come a mo’
d’esempio leggasi in Cicerone: Epicuri imaginem non modo in
tabulis, sed etiam in poculis et in anulis habere [329]; e altrove:
Tabulas bene pictas collocare in bono lumine [330], come
istessamente oggidì ogni artista domanda di collocare sotto buon
punto di luce le sue opere, per conseguirne i migliori effetti. Plinio poi
aggiunge che si pingesse pure sovra pelli o membrane [331] ed
anche sulla tela. Su tela, per la prima volta venne dipinta quella
grande demenza che fu la colossale immagine di Nerone, alta
centoventi piedi, che fu consunta dal fulmine, come disse lo stesso
Plinio, nei giardini di Maja: Nero princeps jusserat colosseum se
pingi CXX pedum in linteo incognitum ad hoc tempus. Ea pictura
cum peracta esset in Majanis hortis, accensa fulmine cum optima
hortorum parte conflagravit [332]. Nè fu sola codesta prova del
pingere sulla tela allora. Un liberto dello stesso Nerone, dando al
popolo in Azio uno spettacolo di gladiatori, coprì i publici portici di
tappezzerie dipinte su cui rappresentavansi al naturale i gladiatori e i
loro inservienti.
Del resto anche le pitture pompejane offrirono dati, di cui giova tener
conto, in conferma di quest’uso di dipingere a pittura sulla tela,
avendosene una su parete, la quale rappresenta una dama che sta
dinanzi al cavalletto intenta a dipingere un quadro o piuttosto a
copiare un Bacco che le sta davanti; un’altra in cui si raffigura un
pigmeo, che dipinge nella medesima guisa, ed una terza, nella quale
è un’altra donna che pur così dipinge, la quale, scoperta nel 1846, gli
Archeologi opinarono potesse essere la famosa Jaia di Cizica
conosciuta da Varrone e celebrata da Plinio [333].
Recenti scavi fatti a Pompei (1872) rivelarono come si dipingesse
anche sul marmo. Su di una tavola appunto marmorea, rinvenuta
non ha guari, si ammirò una buona pittura a più figure, che l’illustre
Fiorelli interpretò per la scena tragica dell’infelice Niobe; importante
scoperta codesta che viene ad aggiungere una notizia di più alla
storia della pittura.
Le Pitture di Pompei si possono classificare, oltre quelle
architettoniche e decorative, di cui dissi più sopra, in istoriche —
intendendo comprendersi in esse tutti i soggetti mitologici od eroici
— e giovan moltissimo, oltre che a rischiarare quanto noi sappiamo
di mitologia e della storia dei secoli favolosi ed eroici, a fornire altresì
nozioni circa la scienza del costume, l’arte drammatica e l’antichità
propriamente detta; in quelle delle figure isolate e di genere; di
paesaggi e animali e fiori; ommettendo per ora qui d’intrattenermi
particolarmente di quelle delle insegne delle tabernæ, perchè ne ho
già detto abbastanza nel Capitolo antecedente, per quel che ne
importava di sapere; molto più che a rigore implicitamente io ridirò di
esse, favellando della pittura di genere a cui appartenevan le dipinte
insegne.
Ho più sopra, sulla fede di Plinio, ricordato alla sfuggita i nomi di
alcuni pittori conosciuti in Roma: or ne dirò il giudizio che di loro
opere fu lasciato ai posteri. Per Fabio, l’arte non doveva essere che
un divertimento, perchè, come patrizio, non aveva d’uopo
d’esercitarla, ne diffatti riuscì grande la riputazion sua come pittore.
D’altronde così ragionava Cicerone: «Crederemmo noi che ove si
fosse fatto titolo di gloria a Fabio l’inclinazione che mostrava per la
pittura, non fossero stati anche fra noi dei Polignoti e dei Parrasii?
L’onore alimenta le arti: ciascuno è spronato dall’amor della gloria a
dedicarsi ai lavori che possono procurargliela: ma languono gli
ingegni ovunque sieno tenuti in non cale.» Pacuvio, poeta e pittore,
quantunque arricchisse de’ proprii dipinti il Tempio di Ercole nel Foro
Boario, sembra nondimeno che queste sue opere d’arte non
oscurassero del loro bagliore la maggior fama acquistata da lui colla
tragedia di Oreste, ancorchè la drammatica si trovasse in quel tempo
nella sua infanzia. Arellio, vissuto qualche tempo appena prima
d’Augusto, si rese celebre, ma fu rimproverato d’aver corrotto l’onore
dell’arte sua con una insigne turpitudine e quando toglieva a pingere
una qualche Iddia, le prestava i tratti e le sembianze di qualche
sciupata, della qual fosse innamorato; suppergiù come fanno tanti
dei pittori odierni col dipinger sante e madonne, onde la pittura sacra
cessò di produrre que’ capilavori, che ispirava ne’ tempi addietro la
fede. Amulio invece fu severo e si narra che la sua Minerva
riguardasse lo spettatore da qualunque parte le si fosse rivolto; Accio
Prisco si loda per accostarsi meglio alla maniera antica, e
quest’ultimo viveva ai tempi di Vespasiano, quindi presso agli ultimi
giorni di Pompei.
In questa città, più ancora che in Roma, risentono le pitture del far
greco: certo molti artefici greci quivi furono e lavorarono. Voler tener
conto, oltre quelli che già rammentai lungo il corso dell’opera, di tutti
i dipinti di storia e di mitologia, non mi trarrei sì presto di briga: d’altra
parte ho accennato in capo di questo articolo che tratta delle Arti
belle, le opere che ne discorsero in proposito partitamente e ne
riprodussero i disegni. Tuttavia, come non dire del quadro di Tindaro
e di Leda, meritamente considerato come uno dei più preziosi avanzi
della pittura degli antichi? Vivacità di colorito, armonia di tinte e
leggiadria di composizioni fecero lamentare che i suoi colori oggi
sieno di tanto smarriti. Così la bella Danae scoperta nella casa di
Pansa; l’Adige ferito nelle braccia di Venere, che diede il nome alla
casa in cui fu trovato; il Meleagro, onde fu detta la casa in cui era,
nella quale pur si trovarono Achille e Deidamia; Teti che riceve da
Vulcano le armi d’Achille; Dejanira su d’un carro che presenta Ilio
figliuol suo ad Ercole, mentre il Centauro Nesso offre di tragittarla sul
fiume Eveno, e Meleagro vincitore del cinghiale Calidonio; ambe
rinvenute nella casa detta del Centauro; Castore e Polluce, il Satiro
ed Ermafrodito, Apollo, Saturno, Achille fanciullo immerso nello
Stige, Marte e Venere, Endimione e Diana, Eco e Narciso, Giove, La
Fortuna, Bacco, pitture tutte scoperte nella preziosissima casa che
fu detta di Castore e Polluce od anche del Questore; Venere che
pesca all’amo; Ercole ed Onfale, grande pittura scoperta nella Casa
di Sirico e i diversi subbietti spiccati all’Iliade, che si giudicarono fra i
dipinti più egregi per composizione ed esecuzione finora tratti in
luce, che si videro nella Casa d’Omero, detta altrimenti del Poeta; fra
cui pure fu ammirata la bellissima Venere, che Gell non esitò punto a
paragonare per la forma a quella sì celebrata de’ Medici, e pel colore
a quella non men famosa di Tiziano.
Fu nel peristilio della medesima casa di Sirico che si trovò il
Sacrificio di Ifigenia, copia del rinomato quadro che fu il capolavoro
di Timanto.
Certo che tutte le opere di pittura in Pompei non hanno egual merito,
nè vantar si ponno di molta correzione; ma in ricambio quasi sempre
vi è l’evidenza, l’espressione, la verità, vivacità di colorito e
intonazione ad un tempo e la luce ben diffusa e tale spesso da
rischiarare di sè la piccola camera. «Et quel mouvement dans toutes
ces figures, sclama Marc Monnier, che amo citare, perchè autorità di
straniero, quelle souplesse et quelle vérité!! Rien ne se tord; mais
rien ne pose. Ariane dort réellement, Hercule ivre s’affaisse, la
Danseuse flotte dans l’air comme dans son élément, le centaure
galope sans effort; c’est la réalité simple (tout le contraire du
Réalisme), la nature belle qu’elle est belle, dans la pleine effusion de
sa grâce, marchant en reine parce quelle est reine et qu’elle ne
saurait marcher autrement. Enfin ces peintres subalternes, vils
barbouilleurs de parois, avaient à defaut de science et de correction,
le genie perdu, l’instinct de l’art, la spontaneité, la liberté, la vie [334].
La quale splendida testimonianza che ho recata colle parole del
francese scrittore, applicar non si vogliono — intendiamoci bene — a
quelle principali composizioni storiche o mitologiche che ho più
sopra annoverate, ma sì a quelle altre sole che superiormente ho
distinte nella classe delle Figure isolate. Sentimento di quel critico,
diviso pure da ben altri e da me, è che alle più importanti opere di
pittura, alle storie onde si decoravano le pareti più ampie delle case
si ponessero artisti i meglio riputati, e che invece a quelle isolate, si
applicassero altri di minor levatura.
Queste figure isolate servivano il più spesso a decorare le pareti
minori, quando pure le grandi non venissero disposte a piccoli
quadri, a decorar lacunari ed agli effetti architettonici: erano il più
spesso Ninfe, Danzatrici, Baccanti, Centauri, Sacerdotesse,
Canefore e Cernofore, Genj alati, Amorini, Fame, Vittorie e Fauni,
Muse e Iddii. Ma non furon sempre opere di poco momento o di
merito secondario; perocchè talune, che si ebbero conservate,
fossero altrettanti capolavori. Le raccolte infatti che si publicarono
de’ pompeiani dipinti, fra le tante figure isolate, recarono i disegni di
veri piccoli capolavori. Tale a mo’ d’esempio è la imponente Cerere
rinvenuta nella Casa detta di Castore e Polluce o del Questore, in
cui sembrarono essere di proposito state accumulate le più leggiadre
opere di arte; il Giove che stava nella casa detta del Naviglio di
fronte ad uno dei lati del tempio della Fortuna, e il quale ha tutta la
maestà del padre degli Dei. Ma forse la figura isolata più pregevole,
è per generale avviso la Meditazione seduta su d’una sedia dorata, i
lineamenti della quale sono eseguiti con tanta cura e dotati di tanta
espressione, direbbesi individuale, da essere indotti a credere che
dovesse essere un ritratto. E poichè sono a dire di questo genere di
pittura, che è de’ ritratti, il lettore ricorderà i due ritratti di Paquio
Proculo e di sua moglie, de’ quali m’ebbi ad intrattenere nel passato
capitolo: se non sono essi da collocarsi fra le migliori opere,
attestano nondimeno della costumanza che si aveva fin d’allora di
farsi ritrarre, oggi divenuta omai una mania, atteso il buon mercato
della fotografia.
Non vogliono poi essere passati sotto silenzio l’Apollo Musagete, o
duce delle Muse e le nove Muse, isolatamente dipinti in tanti quadri,
che si rinvennero negli scavi nell’anno 1785 nella vicina Civita,
avente ognuna figura i proprj emblemi, e più che questi, la propria
peculiare espressione.
In quanto alla pittura di genere, non voglio dire si possano vantare
tante meraviglie quante se ne ammirarono in quella istorica, e quel
che ne vado a dire ne fornirà le ragioni. In quella vece vuolsene
segnalare un certo pregio in ciò che a me, come agli scrittori delle
cose pompejane, giovarono alla interpretazione di importanti cose
attinenti l’arti e i mestieri.
Fin da quando le scuole greche piegarono a decadenza, per la
smania del nuovo, molti artisti si buttarono ad una pittura casalinga e
di particolari, che i moderni chiamarono di genere. Là, dove l’arte si
ispirava sempre al grande ed al meraviglioso ed era aliena per
conseguenza dal realismo, l’avvenimento non fece che sollevare il
dispregio e questi artisti si designarono come pittori di cenci, e noi
diremmo da boccali. Tuttavia sulla mente del popolo que’ dipinti che
si esponevano ad insegna di bottega facevano impressione: erano
richiami influenti e que’ dipinti si ricercarono a furia da merciai e
venditori. Così Pireico ottenne fama, sapendo dipingere insegne da
barbiere e da sarto; Antifilo, pingendo uno schiavo soffiante nel
fuoco ad una fabbrica di lana; Filisco un’officina da pittore e Simo il
laboratorio d’un follone [335]. Orazio ci ha poi ricordato ne’ versi, che
in altro Capitolo (Anfiteatro) ho citati, un’insegna gladiatoria e nel
capitolo delle Tabernæ ne ho pur toccato: or ne dirò, poichè mi cade
in taglio, qualche cosuccia ancora.
L’Accademia d’Ercolano publicò ne’ suoi atti una serie di quadri
scoperti fin dal 25 maggio 1755 in cui sono appunto espressi
mercanti e lavoratori in pieno mercato, forse nel Foro di Pompei, dal
quale appena differenziano i capitelli delle colonne, che in quel di
Pompei son dorici, mentre nella architettura de’ quadretti
summentovati appajono corintii; perocchè nel resto tutto vi sia
fedelmente riprodotto.
In uno si veggono due mercanti di drappi di lana, che trattan di loro
merce con due avventrici: in altro è un calzolajo che in ginocchio
prova a calzare delle scarpe a un suo cliente e nel fondo appiccate
veggonsi al muro diverse calzature.
Un altro calzolajo è raffigurato in un altro quadro e colla bacchetta
alla mano con cui misura il piede, porge un calzaretto a quattro
pompejane sedute, di cui l’una ha in grembo un fanciullo. Buona la
composizione, pessima ne è la esecuzione. Evvi un’altra pittura, in
cui sono due genietti pure calzolai: l’uno ha nelle mani una forma,
l’altro ha il cuojo che si dovrà adattare su di essa, onde foggiarlo a
calzatura. In un armadietto a due battenti aperti veggonsi scarpe
diverse e presso un vasetto e un bacino contenenti il colore per dare
il lucido alle pelli. Evvi anche il commerciante di forbici, fibule,
spilloni; il vasajo e il panattiere seduto alla turca sul suo banco.
Parlando della Fullonica, già dissi del dipinto che vi si trovò nel 1826
e che reca tutto il progresso di quell’arte de’ gualcherai, di
esecuzione assai migliore delle altre, nè mi vi arresterò di più: invece
nelle pitture di decorazione veggonsi fabbri, genietti alati con arnesi
fabbrili, altri recanti nelle mani quelli del pistrino e del torchio.
Del pregio de’ paesaggi, che formavano soventissime volte il fondo
dei dipinti storici più importanti, dissi più sopra, e noterò meglio ora,
che pur frequenti volte si trovino nei fregi e sotto altri dipinti di figura,
riprodotti animali, come buoi, lupi, pantere, capre ed uccelli, talvolta
condotti con non dubbia abilità.
Ora una breve parola della pittura de’ fiori. Nulla, scrive Beulé, nulla
è più grazioso che le pitture rappresentanti piccoli genii che figurano
il commercio de’ fiori. La gran tavola coperta di foglie e di fiori, i
panieri che essi portano, che vuotano e riempiono, le ghirlande
sospese, tutto ci richiama Glicera, la bella fioraia di Sicione che il
pittor Pausia amava, e della quale egli copiava i bei mazzi, tanto
sapeva comporli, disponendovi i colori ed ogni loro gradazione. Le
più leggiadre rappresentazioni in questo genere, trovansi in quattro
comparti decorativi d’una camera sepolcrale che Santo Bartoli ha
disegnati. V’hanno fanciulli che colgono de’ fiori, ne riempiono i
cestelli, ne caricano le spalle loro, attaccati in equilibrio su d’un
bastone. Il picciolo mercante che va tutto nudo recando questa serie
di ghirlande, è veramente degno di osservazione.

Scultura.

Veniamo ora alla parte statuaria, seguendo anche in questo

argomento l’egual sistema di trattar delle sue condizioni generali nel
mondo romano, particolareggiando poi intorno alle opere
pompejane.
Io intendo trattare in proposito non solo delle statue in marmo, ma
ben anco di quelle in bronzo e de’ bassorilievi; perocchè sieno tutte
opere attinenti la scultura; solo omettendo di quelle ne’ più preziosi
metalli, per averne già toccato alcunchè parlando degli orefici e delle
orificerie in Pompei.
Anche nella statuaria l’Etruria precedette ogni altra parte d’Italia. Se
le sue prime produzioni presentarono un genere presso che eguale
alle produzioni de’ popoli incolti, presto per altro assunsero una
propria maniera: anzi dai molti saggi recati alla luce dagli scavi in più
località praticati, e di cui sono arricchiti i nostri musei, si può asserire
che due fossero le maniere, o stili affatto distinti di disegno. Del
primo, dice Winkelmann, esistono ancora alcune figure e somigliano
le statue egiziane, in quanto che hanno esse pure le braccia pendule
ed aderenti alle anche, ed i piedi posti parallelamente l’un presso
l’altro: il secondo è caratterizzato dalle esagerazioni dei movimenti e
di una ruvidezza di espressione che gli artisti si sforzarono di
imprimere ai loro personaggi.
Da questa seconda maniera si fa passaggio alla maniera de’ Greci,
di cui gli Etruschi, dopo l’incendio di Corinto e il saccheggio d’Atene
datovi da Silla, che spinse gli artisti di Grecia in Italia, divennero
discepoli e collaboratori.
Tuttavia i primi Etruschi potevano a buon dritto vantarsi d’aver con
successo trattato scultura e pittura fino dai tempi in cui i Greci non
avevano che assai scarsa cognizione delle arti che dipendono dal
disegno: argomento pur questo che avvalora la credenza dal
Mazzoldi espressa nelle sue Origini Italiche, non abbastanza
apprezzate come si dovrebbe, che la civiltà fosse prima dall’Italia
importata alla Grecia, i Pelasghi appunto e gli Atalanti procedendo
dalla terra nostra.
Plinio ne fa sapere infatti che in Italia venisse eseguita una statua
innanzi che Evandro, mezzo secolo prima della guerra di Troja,
giungendo dalla nativa Arcadia, sostasse sulle sponde del Tevere e
vi fondasse Pallantea, che ho più sopra ricordato; e ne trasmise pur
la notizia che nel suo tempo si vedevano ancora a Ceri, una delle
dodici principali città dell’Etruria, affreschi d’una data anteriore alla
fondazione di Roma. Egualmente le pitture del tempio di Giunone,
condotte da Ludio Elota, prima che Roma esistesse, in Ardea, città
del Lazio, capitale dei Rutuli e soggiorno del re Turno, come
raccogliesi dal Lib. VII dell’Eneide, della freschezza delle quali
rimaneva quell’enciclopedico scrittore maravigliato, non meno che
dell’Atalanta e dell’Elena di Lanuvio, la prima ignuda, la seconda
invece decentemente palliata e spirante timidezza e candore.
Fondata Roma, Romolo e Numa ricorsero, come mezzo di
civilizzazione, a imporre il culto a quelle immagini di numi che
Evandro ed Enea avevano recato in Italia ne’ più remoti tempi,
quantunque poi per crescere loro autorità, li avessero a circondar di
mistero tenendoli gelosamente ascosi agli sguardi profani. Le prime
opere di scultura presso i Romani si vuole essere state statue di
legno o di argilla, raffiguranti divinità. D’altri personaggi si citano le
statue: di Romolo, cioè, di Giano Gemino, consacrate da Numa, la
prima anzi delle quali coronata dalla Vittoria, fu poscia collocata
sopra una quadriga di bronzo tolta dalla città di Camerino. Più
innanzi il simulacro dell’augure Accio Nevio e delle due Sibille e le
statue equestri di Orazio Coclite e di Clelia trovasi scritto che
venissero fuse in bronzo in epoca assai prossima a quella in cui
siffatti personaggi avevano vissuto. Anche nel Foro Boario venne
collocata in quei primi tempi una statua d’Ercole trionfatore; come
più tardi sulla piazza del Comizio venne posta quella di un legista di
Efeso, appellato Ermodoro. Un Turriano, scultore di Fregela lavorò in
rosso una statua di Giove e fu cinque secoli avanti l’Era Volgare. Un
Apollo di smisurata grandezza fu da Spurio Carvilio, dopo la sua
vittoria contro i Sanniti, fatta fondere in bronzo, usando all’uopo delle
spoglie tolte a’ vinti nemici.
Non è che dugentottanta anni prima di Cristo, dopo la sconfitta di
Pirro, che a Roma venne offerto lo spettacolo di una esposizione di
molti e pregevoli oggetti d’arte, quadri e statue che dall’esercito
Romano erano stati predati in un con ingente quantità d’oro e
d’argento, ed altre preziosità negli accampamenti del Re epirota, che
alla sua volta erano stati da lui derubati nella Sicilia ai Locri, nel
saccheggio dato al tesoro del loro tempio sacro a Proserpina.
I monumenti che la nazione consacrava ad eternare i fasti più
gloriosi consistevano d’ordinario in semplici colonne, rado a’ più
valorosi capitani concedevasi l’onoranza di una statua, limitata
l’altezza a tre piedi soltanto, onde designate venivano codeste statue
col nome di tripedaneæ; mentre poi sigillæ dicevansi quelle più
piccole d’oro, d’argento, di bronzo od avorio, le quali erano per solito
d’accuratissimo lavoro.
Pel contrario nelle case private, i patrizj collocavano in apposite
nicchie le immagini dei loro illustri maggiori, e se n’era anzi costituito
un diritto, solo concesso a coloro che avessero sostenuto alcuna
carica curule, come il tribunato e la questura, ed erano la più parte
lavorate in cera; talvolta per altro in legno, in pietra od in metallo. Chi
di tal diritto fruiva, poteva processionalmente siffatte immagini
portare nelle pompe funerali, lo che ognuno si recava a sommo di
onore.
La presa di Siracusa per opera di Marcello e le conquiste della
Macedonia, della Grecia e dell’Asia, per quella di Paolo Emilio, di
Metello e de’ Scipioni, dovevano rendere più acuto il desiderio di
possedere oggetti d’arte, comunque non fosse giunto, come dissi più
sopra, a farsene il virtuale apprezzamento neppur a’ tempi della
maggiore coltura, come furono quelli di Cesare e di Cicerone.
Perocchè i trionfatori tutte le meraviglie dell’arte, di cui i Re
macedoni avevano impreziosito le loro principali città, mandassero a
centinaja di carra a Roma, in un con tante altre ricchezze; per modo
che per centoventitre anni di seguito non s’avesse bisogno di
prelevare imposte sui cittadini.
Come già narrai, si travasarono dalla Grecia in Roma, coi capolavori
dell’arte, anche i loro artefici, e poterono così stabilirvi le loro scuole
e sistemi e farli prevalere, che il Senato con apposito decreto mandò
multarsi gli scultori che si fossero allora allontanati dalle dottrine di
essi, la eccellenza delle quali veniva tanto splendidamente attestata
da sì egregie ed immortali opere.
Le statue, che dapprincipio avevano giovato soltanto ad illustrare i
trionfi, vennero pertanto adoperate ben presto a decorar piazze e
monumenti e quindi a crescere il fasto delle abitazioni private. Silla,
Lucullo, Ortensio e il cliente di lui Cajo Verre, diedero il primo
esempio: dietro di essi vennero tutti i più facoltosi degli ultimi tempi
della Republica, seguiti pure da tutti quelli delle colonie, come
rimangono ad attestarcelo tanti capolavori, de’ quali più avanti dirò,
rinvenuti in Ercolano e Pompei.
Di tal guisa gli artisti greci venuti a Roma vi trovarono molto lavoro e
fortuna. Si citano fra essi un Pasitele, che eseguì una statua nel
tempio di Giove e n’ebbe il diritto di cittadinanza; i suoi allievi Colote
e Stefanio; Arcesilao, che scolpì la Venere genitrice per Cesare, il
qual si vantava da lei discendere; Apollonio e Glicone, autore
dell’Ercole Farnese, condotti a Roma da Pompeo; Alcamene e
Cleomene, Poside e Menelao, Decio e Damasippo, e sotto Augusto
Menofante e Lisia, Nicolao e Critone e l’ateniese Diogene, che
condusse le statue che si collocarono sul frontone del Pantheon di
Agrippa. Augusto stesso fece erigere sotto il portico del suo Foro le
statue degli oratori e de’ trionfatori più illustri, e in Campidoglio si
vede tuttavia una statua di questo imperatore con un rostro di nave
a’ suoi piedi in memoria di sua vittoria sovra Sesto Pompeo.
Ma ho ricordato testè Cajo Verre. Or come si può, anche
sommariamente, come faccio io qui stretto dall’economia dell’opera,
ritessere la storia dell’arti, o piuttosto compendiarla, senza ricordare
le gesta infami di quest’uomo, contro cui si scagliarono le folgori
della eloquenza del sommo Oratore?
Cajo Licinio Verre, senatore, era stato proquestore dapprima in
Cilicia, poscia pretore per tre anni in Sicilia, dove, non pago
dell’ordinaria rapacità de’ suoi colleghi, trattò la Sicilia più che paese
di conquista. Abusando del suo potere, fece man bassa su tutte le
preziosità publiche e private. Quadri e statue de’ più celebri artisti
attiravano specialmente la sua cupidigia, nè ristava davanti alcuna
legge divina ed umana per venirne in possesso. Cicerone,
difendendo contro lui le ragioni de’ Siciliani, aveva potuto senza
iperboli esclamare: «Dove sono le ricchezze strappate a forza a tanti
popoli soggetti e ridotti omai alla miseria? Potete voi chiederlo, o
Romani, quando vedete Atene, Pergamo, Mileto, l’Asia, la Grecia
inghiottite nei palagi di alcuni spogliatori impuniti?» Minacciato per
ciò d’accusa capitale, egli aveva prese le sue cautele, nè restava
punto dal dichiararle svergognatamente a’ suoi amici, dicendo: io
feci del prodotto del triennio che durò la mia pretura tre parti: una per
il mio difensore, una per i miei giudici e la più grossa per me.
Il suo difensore fu Ortensio, che oltre essere rinomatissimo oratore,
era appassionatissimo delle opere di arte, vantandosi fra gli altri di
possedere il dipinto degli Argonauti di Cidia, che aveva costato
centoquarantaquattro mila sesterzi, che sarebbero lire ventottomila
delle nostre. Cicerone dice di Verre: che in lui la cupidigia d’artistiche
cose era un furore, un delirio, una malattia, dacchè in tutta la Sicilia
non esistesse un vaso d’argento o di bronzo, fosse di Delo o di
Corinto, non una pietra incisa, non un lavoro di oro, d’avorio, di
marmo, non un quadro prezioso, non un arazzo, che quell’avido
governatore non volesse vedere co’ proprj occhi, per trattenere poi
quanto gli sembrava opportuno ad arricchire la di lui raccolta. Basti
per tutti il seguente aneddoto. Antioco, figlio del re di Siria, volendo
sollecitare l’amicizia del Senato romano, viaggiava alla volta della
grande città, seco recando un ricchissimo candelabro, per donarlo al
tempio di Giove Capitolino. Transitando per la Sicilia, ricambiando
Verre d’una cena, questi vedute le mille preziosità che aveva, sotto
pretesto di mostrarle a’ suoi orefici, se le faceva col candelabro
portar a casa. Vi consentiva Antioco; ma quando si trattò della
restituzione, il ladro pretore tanto insistè per aver tutto in dono, che
alla perfine, a lasciargli ogni cosa si decise, purchè almeno gli
rendesse il candelabro destinato al popolo romano. Verre ricorse
dapprima a pretesti per ricusargli anche questo; ma poi impose
termine a ogni contesa, intimando recisamente al re che
sgombrasse avanti notte dalla provincia.
Chiederà il lettore se tanto ladro venisse poi condannato.
Risponderò che non fossero questi soltanto i gravami contro lui
formulati: altri furti e depredamenti d’ogni genere da lui commessi in
Acaja, in Cilicia, in Panfilia, in Asia ed in Roma; la venduta giustizia,
le violenze, gli stupri, l’oltraggiata cittadinanza; sì che li potesse così
Cicerone riassumere nella Verrina De Frumento: Omnia vitia, quæ
possunt in homine perdito nefarioque esse, reprehendo, nullum esse
dico indicium libidinis, sceleris, audaciæ, quod non in unius istius vita
perspicere possitis [336].
Ebbene, Cicerone appena ne potè assumere l’accusa protetto da
Pompeo; ma il fece con tanto vigore fin dalla prima arringa, che
spaventatone Ortensio, disperando dell’esito del suo patrocinio, ne
dimise tosto il pensiero, e Verre stesso persuaso, andò
spontaneamente in esiglio, dove passò la trista vecchiezza e morì
proscritto da’ Triumviri, come ce lo lasciarono ricordato Seneca e
Lattanzio [337], condannato però a restituire a’ Siciliani
quarantacinque milioni di sesterzi, dei cento che essi avevano
addomandato e che forse non rappresentavano tutto il danno della
sua rapina.
Le altre orazioni che contro Verre si hanno di Cicerone, non furono
recitate, attesa la contumacia dell’accusato; ma corsero allora
manoscritte per le mani di tutti e a’ posteri rimasero monumento,
come di eloquenza, così della condotta de’ magistrati romani, che
nella propria Verre compendiava.
Toccai più sopra di Augusto, come l’arti favorisse e gli artisti, e da
Tito Livio infatti egli venne chiamato per antonomasia riedificatore
dei templi, ed a buon diritto però potè morendo dire di sè: Trovai
Roma fabbricata di mattoni e la lascio fabbricata di marmo.
Fu sotto di lui che Dioscoride portò la glittica, od incisione in pietre
dure, al più perfetto suo grado e si levano a cielo il suo Perseo, la Io,
il Mercurio Coriforo, ossia portante un capro, e il ritratto dell’oratore
Demostene inciso su d’un’ametista. Nello stesso genere furono pure
lodatissimi Eutichio figliuolo di Dioscoride, Solone, Aulo e chi sa
quanti altri, se i Musei tutti serbano i più squisiti cammei di quel
tempo e se dagli scavi di Pompei se ne tolsero di preziosissimi, tali
da formare la meraviglia di chi li vede adesso nel Museo Nazionale
di Napoli.
E poichè m’avvenne di ricordare la glittica o glittografia, come altri la
chiama, nel difetto di trattati antichi intorno a quest’arte, — perocchè
non se n’abbiano che pochi cenni nelle opere di Plinio, — parmi
potersi ritenere che avessero a un dipresso in antico i metodi che si
han pur di presente in Italia, dove a preferenza fu sempre coltivata e
portata a perfezione. Ad intaccar la pietra, si usava dagli artefici
romani una girella che chiamavano ferrum obtusum, e la cannella da
forare appellata da Plinio terebra, giovandosi del naxium, specie di
grès di Levante, poi dello schisto d’Armenia, a cui sostituirono da
ultimo lo smirris, che noi diremmo smeriglio. Credesi nota agli antichi
la punta di diamante come strumento attissimo all’incisione. Delle
pietre dure incise valevansi principalmente a farne anelli e sigilli;
onde, litoglifi dicevansi gli artisti intagliatori di pietre, e dattilioglifi gli
intagliatori di anelli, usandosi anche nello stesso significato le parole
più prettamente latine sculptores e cavatores.
E qui chiudo la parentesi, che parvemi necessaria a dare una
qualunque idea di questa particolare arte del disegno e ritorno al
primo subbietto.
Del tempo di Tiberio, ha poco a registrare la storia dell’arte, se non è
la statua di quel Cesare che fu ritrovata in Carpi e una effigie
colossale ricordata da Flegone, liberto d’Adriano, nel suo Trattato
delle cose meravigliose. Nè di meglio del tempo di Caligola e di
Claudio; comunque del primo si sappia ch’ei chiedesse agli artisti
invenzioni straordinarie e prodigiose: di quello di Nerone varrebbe
invece intrattenerci, se di lui stesso ebbe a scrivere Svetonio:
pingendi fingendique, non mediocre habuit studium [338]; ciò che
inoltre gioverebbe a provare che la professione dell’Arti Belle, non
fosse più, come per lo addietro, cotanto disprezzata.
A lui è imputato d’aver dato l’incendio a parecchi quartieri della città;
ma lo si pretende escusare, dicendo averne avuto il pensiero per
distruggere tante indecenti catapecchie, onde in luogo di esse
sorgessero palagi, si ampliassero le vie, e potesse meglio allinearne
le mura e imporre alla città il proprio nome, appellandola Neropoli. E
vi fabbricò infatti la Casa di Oro, che dissi più sopra quant’area
occupasse e vi fe’ portare statue e quadri de’ più famosi autori,
arredi d’oro, d’argento, d’avorio e madreperla, e collocarvi quel
colosso, di cui pure più sopra accennai, eseguitovi da Zenodoro, alto
cento piedi, nel cui lavoro impiegò l’artefice ben dieci anni e il cui
valore fu di quaranta milioni di sesterzi, a dir come nove milioni di lire
italiane.
Sotto gli imperatori di casa Flavia, Vespasiano e Tito, dai quali
veduto abbiamo eretto l’Anfiteatro, detto il Colosseo, non si sa che
importanti opere scultorie venissero in Italia condotte, dove si
eccettuino una bella statua rappresentante l’imperatore Tito ed una
bella testa colossale del medesimo imperatore, che Winkelmann
ricorda.
È anche a quest’epoca appartenente di certo, secondo l’opinione
d’Ennio Quirino Visconti, l’incomparabile gruppo del Laocoonte, degli
artisti rodii Agesandro, Atenodoro e Polidoro, che Plinio non esitò a
dichiarare Opus omnibus et picturæ et statuariæ præponendum [339].
Viste così le principali cose di scultura pertinenti all’epoca imperiale
infino ai giorni del cataclisma vesuviano, mi conviene ora
completarne il discorso colle opere scoperte in Ercolano e in
Pompei.
Le più grandi si rinvennero in Ercolano: tali la statua equestre in
marmo di Nonio Balbo e quella del figliuol suo, che ne
fiancheggiavano la basilica e la importanza delle quali opere lasciò
indovinare l’importanza altresì de’ personaggi che rappresentavano;
onde ne dolse che nè la storia, nè gli scavi abbiano finora portato
alcun lume su di essi; tali il magnifico cavallo in bronzo, rinvenuto nel
teatro e già spettante ad una quadriga, che nei primi scavi colà
praticati venne messa sventuratamente a pezzi, e ricomposta poi,
forma oggidì una delle più interessanti e preziose opere del Museo
di Napoli in un con alcune figure del bassorilievo del carro, con un
Bacco, otto statue consolari, quelle di Nerone, Claudio Druso e sua
moglie Antonia, un ministro di sagrifici in bronzo e due teste di
cavallo, e una statua di Vespasiano, e due di bronzo rappresentanti
Augusto e Claudio Druso. Nello stesso Museo di Napoli, sono pure
le statue di sei celebri danzatrici, trovate nella casa detta dei
Papiri [340], il magnifico Fauno ebbro, in bronzo, ch’era a capo della
piscina nello xisto di essa casa, i busti di Claudio Marcello, di Saffo,
di Speusippo, Archita, Epicuro, Platone, Eraclito, Democrito,
Scipione l’Africano, Silla, Lepido, Augusto, Livia, Cajo e Lucio
Cesare, Agrippina, Caligola, Seneca, Tolomeo Filadelfo, Tolomeo
Filometore, Tolomeo IX, Tolomeo Apione, e Tolomeo Sotero I; di due
Berenici e di due altri personaggi sconosciuti. Una statua e cinque
busti in bronzo si trovarono nella stessa casa, su l’un de’ quali si
lesse il nome dell’artefice: Apollonio figlio di Archia ateniese.
Circa alle opere di statuaria rinvenute in Pompei, esse sono in
numero minore che non ad Ercolano; ma non sono meno pregevoli.
Io ne ricorderò taluna e saranno le principali.
Nel tempio d’Iside fu rinvenuta una bella statua in marmo
rappresentante Bacco, che fu detto, forse dal luogo in cui fu trovato,
Isiaco. Essa appare coronata di pampini la testa, e colla destra
alzata, cui tien rivolto affettuosamente lo sguardo, ci fa supporre che
stringesse un grappolo. Al piede sta una pantera, e sulla base
leggesi la iscrizione che ho già riferita parlando di questo
tempio [341], dove ho pur ricordata la statua di Venere Anadiomene
coll’ombilico dorato. Nel Pantheon, o piuttosto, come più rettamente
fu giudicato, tempio d’Augusto, nel 1821 si trovarono quelle due
statue che già ricordai, le quali si ritennero rappresentare l’una Livia
sacerdotessa d’Augusto, l’altra Druso: la prima sopratutto è una
delle opere di scultura più rimarchevoli che vi si scoprissero. Ha
essa una tale maestà che a chi la riguarda incute reverenza.
Ma siccome a suo luogo ho già ogni volta ricordate le opere di
plastica; così de’ marmi, altro non ricorderò qui che il bassorilievo in
marmo di Luni, o come direbbesi oggi di Carrara, raffigurante una
biga, su cui sta un africano, e alla quale sono attelati due corridori
preceduti da un araldo, come si usava comparire in publico da’
magistrati. La purezza dei disegno e della esecuzione rivelano
l’artefice greco.
Dirò meglio de’ bronzi.
Leggiadra è la statuetta trovata in una nicchia innalzata nel mezzo di
una stanza, rappresentante Apollo, e sì ben conservata, che
neppure appajano danneggiate le corde argentee della lira. Taluni,
all’acconciamento alquanto muliebre de’ capelli, alla delicatezza de’
lineamenti del volto, ed all’espansione del bacino, vollero invece
ravvisarvi un Ermafrodito.
Una Diana vendicatrice, sarebbe la più armonica figura, se non vi
disdicessero certe alacce mal attaccate: più lodevole e prezioso
lavoro è il gruppo di Bacco ed Ampelo, dove gli occhi sono intarsiati
in argento.
Ma tre statuette vi sono che vengono considerate come altrettanti
capolavori e i migliori che furono scavati in Pompei: l’una
rappresentante il Fauno danzante; l’altra Narciso; la terza un Sileno.
Il loro merito si costituisce principalmente dalla maravigliosa e
caratteristica espressione, dalla perfetta concordanza di tutte le parti,
dalla irreprensibile finezza d’ogni particolare, da una esecuzione
insomma completa del bello ideale.
Il Fauno ritrovato nella casa, cui per la propria eccellenza fu imposto
il suo nome ed era nel mezzo dell’impluvium, ha il capo incoronato di
foglie di pino, le braccia alzate, le spalle alquanto rigettate
all’indietro, ogni muscolo in movimento e il corpo tutto in atto di chi
sta per muovere alla danza. Non può essere ammirato il minuto
lavoro del metallo, se non che vedendolo: l’epidermide è così resa
morbidamente, da vedervi sotto la vita: opera certo codesta della più
perfetta fusione, che non ebbe d’uopo per bave o altre scabrosità, di
lima o di cesello dell’artefice.
Fa riscontro al Fauno, il Narciso, statuetta trovata in una povera
casuccia, ed è d’una grazia tutta particolare e nell’atteggiamento
scorgesi come stia in ascolto della Ninfa Eco. La sua testa piega da
una parte, come appunto farebbe chi sta ascoltando voce che
giunga da lontano, teso ha l’orecchio, e il dito rivolto verso dove la
voce muove. L’espressione non poteva essere più felicemente côlta.
Il Sileno finalmente, più recentemente trovato dal comm. Fioretti, è
ancora più perfetto, se è possibile; quantunque pel soggetto si mostri
rattrappito e curvo. Era pur desso decoro d’una fonte: ciò che ne fa
ragionevolmente supporre che dunque nell’interno delle camere vi
fossero ancora maggiori preziosità di quelle ritrovate, se così
all’aperto si tenevano siffatte maraviglie.
Molte opere peraltro della statuaria pompejana accusano la
decadenza dell’arte, come infatti al tempo della catastrofe della città
l’arte romana se ne andava degenerando nel barocco. E fu in questo
tempo che alle statue prevalsero i busti, l’abbondanza de’ quali
segna appunto il nuovo periodo della decadenza cui si veggono
spesso aggiunte le spalle, parte del torace e talvolta le mani e
qualche panneggiamento. Peccano questi assai sovente
d’esagerazione, ma in ricambio conservano l’individualità. Sempre a’
giorni del decadimento si fecero busti di più marmi e colori, l’una
parte nell’altra innestando, massime gli occhi e le vesti.
 Marco Bruto. Vol. II. Cap. XVIII. Belle Arti.

Per completare quanto è attinente alla scultura nel suo più esteso
significato, dovrei dire delle gemme. Qualche cenno ne ho fatto non
ha guari più sopra, parlando della glittica, e per non entrare in
maggiori particolari, ai quali assai si presterebbe il Museo Nazionale
di Napoli, per quanti oggetti si raccolsero negli scavi d’Ercolano e
Pompei, mi basti riassumerne in concetto generale il discorso; che
cioè anche in questo ramo dell’arte i Romani furono dapprima
imitatori de’ Greci, adottandone i soggetti e desumendoli da fatti