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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

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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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Louisiana
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University–Corpus Christi
State University
 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
Another random document with
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pieces far more evident than it is in the ‘Well-Tempered Clavichord’
or even the ‘Inventions’ of Bach. He undoubtedly offers the player
enormous opportunity to exercise his arms and his fingers in the
production of brilliant, astonishing effects.

Of these effects two will always be associated with his name: the
one obtained by the crossing of the hands, the other by the rapid
repetition of one note. Both devices will be found freely used in the
works of his father, and it is absurd to suppose that the son invented
them. Yet it is hardly an exaggeration to say that he made more use
of them than any man down to the time of Liszt. The crossing of the
hands is not employed to interweave two qualities of sound, as it
oftenest is in music for the organ or for the German and French
harpsichords which have two or more manuals that work
independently of each other. The Italian harpsichords had but one
bank of keys, and Scarlatti’s crossing of the hands, if it be not
intended merely for display, succeeds in making notes wide apart
sound relatively simultaneous, and thus produces qualities of
resonance which hitherto had rested silent in the instrument.

It has been suggested that the device of repeated notes was

borrowed from the mandolin, on which, as is well known, a cantabile
is approximated by rapid repetition of the notes of the melody.
Scarlatti, however, rarely employs it to sustain the various notes of
his tune. In his sonatas it is usually, if not intentionally, effective
rhythmically; as it is, unfailingly, in more modern pianoforte music.
On the harpsichord, moreover, as on the pianoforte, it can make a
string twang with a sort of barbaric sound that still has the power to
stir us as shrieking pipes and whistles stirred our savage ancestors.

Still another mannerism of his technique or style is the wide leap of

many of his figures. A plunge from high to low notes was much
practised in contemporary violin music and was considered very
effective, and probably suggested a similar effect upon the
harpsichord. Into this matter again Scarlatti may well have been
initiated by his father, by whom it was not left untried. In the son’s
sonatas it succeeds in extending the range of sonority of the
harpsichord, and thus points unmistakably to developments in the
true pianoforte style.

It is, in fact, by this extension of figures, by sudden leaps, by

crossing of hands, that Scarlatti frees harpsichord music from all
trace of slavery to the conjunct style of organ music; and he may
therefore be judged the founder of the brilliant free style which
reached its extreme development in the music of Liszt. Though we
may not fail to mention occasionally his indebtedness to his father
and to instrumental music of his time, we cannot deny that he is a
great inventor, the creator of a new art. He was admitted by
composers of his day to have not only wonderful hands, but a
wonderful fecundity of invention.

What guided him was chiefly instinct. He had, no doubt, considerable

strict training in the science of counterpoint and composition. He
wrote, as we know, not only harpsichord pieces, but operas and
sacred music as well. In the sonatas there is a great deal of neat
two-part writing, and an occasional flash of skill in imitations; but
musical science is almost the last thing we should think of in
connection with them. Rules are not exemplified therein. Burney
relates, through L’Augier, that Scarlatti knew he had broken
established rules of composition, but reasoned that ‘there was
scarce any other rule worth the attention of a man of genius than that
of not displeasing the only sense of which music is the object.’ And,
further, that he complained of the music of Alberti and other ‘modern’
composers because it did not in execution demand a harpsichord,
but might be equally well or perhaps better expressed by other
instruments. But, ‘as Nature had given him [Scarlatti] ten fingers,
and, as his instrument had employment for them all, he saw no
reason why he should not use them.’ He might have included his two
arms among his natural gifts. Certainly the free use he made of them
in most of his sonatas marks a new and extraordinary advance in the
history of keyboard music.

In the matter of form Scarlatti is not so strikingly an innovator as he

is in that of style. He is in the main content to cast his pieces in the
binary mold common to most short instrumental pieces of his day.
Yet, as has already been suggested, the harmonic freedom which he
enjoys within these relatively narrow limits is significant in the
development of the sonata form; and even more significant is his
distribution of musical material within them.

The binary form, such as we find it in the suites of Froberger and

even in those of J. S. Bach, is essentially a harmonic structure. The
balance and contrast which is the effect of any serviceable shape of
music is here one of harmony, principally of tonic and dominant and
dominant and tonic, with only a few measures of modulation for
variety. There is, in addition, some contrast between that musical
material which is presented first in the tonic key and that which
appears later in the dominant. But, while we may speak of these
materials as first and second themes or subjects, their individuality is
hardly distinct and is, in effect, obliterated by the regularity and
smoothness of style in which these short pieces are conventionally
written. The composer makes no attempt to set them off clearly, one
against the other. The entrance into the dominant key is almost
never devised in such a way as to prepare the listener for a new
musical thought, quite separate and different from that which he has
already heard. The transitional passage from tonic to dominant
emerges from the one and merges into the other, without break or
distinctions.

In the matter of setting his themes in their frame, Scarlatti hardly

differs from his contemporaries. His style, though free and varied, is
in constant motion. But his genius was especially fertile in clean-cut
figures; and when, as he often does, he combines two or three
distinct types of these in one short piece, the music is full of thematic
variety and sparkles with an animation which at times is almost
dramatic.

Scarlatti is, indeed, hovering close to the sonata form in a great

many of his pieces, and in one actually strikes it.[14] We shall,
however, postpone a more detailed discussion of Scarlatti’s pieces in
relation to the sonata form to the next chapter. The distribution of his
musical material is quite whimsical and irregular, always more
instinctive than experimental. It is chiefly by the quality of this
material that he stands apart from his contemporaries, and as the
founder of the free and brilliant pianoforte style.

There remains little to be said of the æsthetic worth of his music.

During the years of his most vigorous manhood he was almost
invariably a virtuoso. Sheer delight in tonal effects rather than more
sober need of self-expression stimulated him. The prevalence of
trumpet figures such as those which constitute the opening phrases
of the eleventh and fourteenth sonatas in the Breitkopf and Härtel
edition already referred to, suggests that he took a good deal of
material ready-made from the operas of the day. Burney says there
are many passages in which he imitated the melody of tunes sung
by carriers, muleteers, and common people. But what he added to
these was his own. A number of pieces are conspicuous by
especially free modulation and expansion of form; and in these,
technical effects are not predominant, but rather a more serious
interest in composition. It has therefore been suggested that these
pieces are the work of later years.[15] Though it is said that while in
Spain he grew too fat to cross his hands at the harpsichord as was
his wont in his youth, this physical restriction is not alone responsible
for the mellowness and warmth of such pieces as the so-called
Pastoral in D minor, familiar to audiences in Tausig’s elaborated
transcription. A great number of his pieces are rich in pure musical
beauty; and the freshness which exhales from all true musical
utterance is and probably always will be theirs.

None of his contemporaries in Italy approached him in the peculiar

skill which has made him conspicuous in the history of pianoforte
music. Francesco Durante (1684-1755) and Nicolo Porpora (1686-
1767), the great singing master, both wrote pieces for the
harpsichord; the one, ‘sonatas’ in several movements, the other
fugues; but their music lacks charm and can hardly be considered at
all influential in the development of the art of writing for keyboard
instruments. Domenico Alberti and P. D. Paradies will be considered
in the following chapter.
 II
The art of Couperin is flawless, the charm of his music not to be
described. It has that quality of perfection with which Nature marks
her smallest flowers. It is the miniature counterpart in music of a
perfected system of living, of the court life of France under Louis XIV.

Scarlatti was a rover. He tried his fortune in Italy, in England, in

Portugal and Spain. He won it by the exhibition of his extraordinary
and startling powers. He was on the alert to startle, his tribute the
bravas and mad applause of his excited hearers. He was the
virtuoso in an old sense of the word, the man with his powers
consciously developed to the uttermost. Bach, on the other hand,
was an introspective, mighty man, immeasurably greater than his
surroundings, fathomless, personal, suggestive. Between them
stands Couperin, for the greater part of his life in the intimate service
of the most brilliant court the world has ever seen, delicate in health,
perfect in etiquette, wise and tender.

Of his life little need be said. He was born in Paris on November 10,
1668, the son of Charles Couperin, himself a musician and brother to
Louis and François Couperin, disciples of the great Chambonnières.
The father died about a year after his son was born, and the musical
education of the young François seems to have been undertaken by
his uncle, François, and later by Jacques Thomelin, organist in the
king’s private chapel in Versailles. Practically nothing is known of his
youth, and, though it is certain that he was for many years organist
at the church of St. Gervais in Paris, as his uncle and even his
grandfather had been before him, the time at which he took up his
duties there has not been exactly determined. There is on record,
however, the account of a meeting held on the twenty-sixth of
December, 1693, at Versailles, at which Louis XIV heard Couperin
play and chose him from other competitors to succeed Thomelin as
his private organist. Thenceforth he passed his life in service of the
king and later of the regent. He died in Paris in 1733, after several
years of ill health.
The great François was, no doubt, an unusually skillful organist, but
his fame rests upon his work for the clavecin, the French
harpsichord, and his book of instruction for that instrument. His
duties at court were various. He says himself that for twenty years he
had the honor to be with the king, and to teach, almost at the same
time, Monseigneur le Dauphin, the Duke of Burgundy, and six
princes or princesses of the royal house.

In his preface to the Concerts royaux he informs us that chamber

concerts were given in the king’s presence on Sunday afternoons at
Versailles, and that he was commanded to write music for them and
that he himself played the clavecin at them. His book on the art of
playing the clavecin, written in 1716, was dedicated to the king. By
all accounts he was a beloved and highly prized teacher and
performer. And neither his pupils nor his fame were confined solely
to the court.

There is no doubt that he was a public favorite and that he published

his pieces for the clavecin to satisfy a general demand. Also in a
measure to safeguard his music. For at that time instrumental pieces
were not often published, but were circulated in manuscript copies in
which gross errors grew rapidly as weeds; and which, moreover,
were common booty to piratical publishers, especially in the
Netherlands. So Couperin took minute care in preparing his music
for his public. Each set of pieces was furnished with a long preface,
nothing in the engraving was left to chance, the books were
beautifully bound so that all might be in keeping with the dainty and
exquisite art of the music itself. Since his day his pieces were never
published again until Madame Farrenc included the four great sets in
her famous Trésor des Pianistes (1861-72). This edition was,
according to Chrysander,[16] very carelessly prepared and is full of
inaccuracies. Chrysander planned a new, accurate and complete
edition, to be edited by Brahms, of which unhappily only one volume,
containing Couperin’s first two books, ever came to print.

The original editions being now rare and priceless, and hardly
serviceable to the average student on account of the confusing
obsolete clef signs, it is to be hoped that before long Chrysander’s
plan will be carried out and the almost forgotten treasures of
Couperin’s clavecin music be revealed in their great beauty to the
lover of music.

Couperin published in all five books of pièces de clavecin. Of these

the first appeared early in the century and is not commonly reckoned
among his best works. The other four sets appeared respectively in
1713, 1716, 1722, and 1730.

Each book contains several sets of pieces grouped together in

ordres, according to key.[17] The canon of the suite is wholly
disregarded and there is very little of the spirit of it. The first ordre, it
is true, has as the first six pieces an allemande, two courantes, a
sarabande, a gavotte, and a gigue; but there are twelve pieces in
addition, of which only three are named dances. The second ordre,
too, has an allemande, two courantes, and a sarabande at the
beginning; but there follow eighteen more pieces of which only four
are strictly dances. The fourth ordre is without true dance forms; so
are the sixth, the seventh, the tenth, and others. Even the orthodox
dances are given secondary titles, or the dance name is itself
secondary. In fact, not only by including within one ordre many more
pieces than ever found place within the suite, but by the very
character of the pieces themselves, Couperin is dissociated from the
suite writers.

He wrote in the preface to his first book of pieces,[18] that in

composing he always had a particular subject before his eyes. This
accounts for the titles affixed to most of his pieces. We have already
referred to ‘battle’ pieces of earlier composers, and to Kuhnau’s
narratives in music. Couperin’s music is not of the same sort. The
majority of his titled pieces are pure music, admirable and charming
in themselves. They are seldom copies. They make their appeal, or
they are intelligible, not by what they delineate, but by what they
express or suggest. The piece as a whole gives an impression, not
the special figures or traits of which it is composed.
Let us consider a few of many types. Take what have been often
called the portraits of court ladies. In these we cannot by any effort of
the imagination find likenesses. It would be ludicrous to try. As ladies
may differ in temperament from each other, so do these little pieces
differ. There is the allemande L’Auguste, which is a dignified,
somewhat austere dance piece in G minor; another, La Laborieuse,
in a complicated contrapuntal style unusual with him. There are three
sarabandes called La Majesteuse, La Prude, and La Lugubre,
impressive, meagre, and profound in turn. These pieces are hardly
personal, nor have they peculiar characteristics apart from the spirit
which is clear in each of them.

Another type of portrait fits its title a little more tangibly. There is La
Mylordine, in the style of an English jig; La Diane, which is built up
on the fanfare figure always associated with the hunt; La Diligente,
full of bustling finger work. Les Nonnettes are blonde and dark, the
blondes, oddly enough, in minor, the dark in major.

Many others are so purely music, delicate and tender, that the titles
seem more to be a gallant tribute to so and so, rather than the
names of prototypes in the flesh. La Manon, La Babet, La fleurie, ou
la tendre Nanette, L’Enchanteresse, La tendre Fanchon, and many
others are in no way program music; nor can they ever be
interpreted as such, since no man can say what charming girl, two
centuries dead, may have suggested their illusive features.

It is these ‘portraits’ particularly which are Couperin’s own new

contribution to the art of music. So individual is the musical life in
each one, so special and complete its character, so full of sentiment
and poetry, that, small as it is, it may stand alone as a perfect and
enduring work of art. It has nothing to do with the suite or with any of
the cyclic forms. Here are the first flowers from that branch of music
from which later were to grow the nocturnes of Field, the Moments
musicals of Schubert, the preludes of Chopin.

Between these and the few pieces which are frankly almost wholly
dependent upon a program are a great number of others lightly
suggestive of their titles. Sometimes it is only in general character.
Les vendangeuses and Les moissoneurs do not seem so particularly
related to wine-gathering or harvesting that the titles might not be
interchanged; but both have something of a peasant character. In
Les abeilles and in Le moucheron the characterization is finer. The
pleasant humming of the bees is reproduced in one, the monotonous
whirring of the gnat in the other. Les bergeries is simply pastoral, Les
matelots Provençales is a lively march, followed by a horn-pipe. Les
papillons is not unlike the little piece so named in the Schumann
Carnaval, though here it means but butterflies. There are some
imitative pieces which are in themselves charming music, such as
Les petits moulins à vent, Le réveille-matin, Le carillon de Cythère,
and Les ondes, with its undulating figures and fluid ornamentation.

Finally the program music is in various degrees programmistic. A

little group of pieces called Les Pèlerines (Pilgrims) begins with a
march, to be played gaily. Then comes a little movement to
represent the spirit of alms-giving, in a minor key, to be played
tenderly; and this is followed by a cheerful little movement of thanks,
to which is added a lively coda. The whole is rather an expression of
moods than a picture of actions. Les petits ages is in some respects
more literal. The first movement, La muse naissante, is written in a
syncopated style, the right hand always following the left, which may
well express weakness and hesitation. L’adolescente, the third
movement, is a lively rondo in vigorous gavotte rhythm.

Two sets are entirely program music. One of these, Les

Bacchanales, has a march (pésament, sans lenteur) of the gray-clad
ones; then three movements expressive of the delights of wine, the
tenderness to which it warms and the madness to which it enflames.
The music is not of itself interesting. More remarkable, though
devoid of musical worth save a good bit of the comical, is Les fastes
de la grande et ancienne Mxnxstrxndxsx. These records or tales are
divided into five acts, which represent the notables and judges of the
kingdom, the old men and the beggars (over a drone bass), the
jugglers, tumblers and mountebanks, with their bears and monkeys,
the cripples (those with one arm or leg played by the right hand,
those who limp played by the left), and, finally, the confusion and
flight of all, brought about by the drunkards and the bears and
monkeys.

III
The last of these compositions are in no way representative of
Couperin the artist. They might have been written by any one who
had a love for nonsense, and they are not meant to be taken
seriously. The quality of Couperin’s contribution to music must be
tested in such pieces as Le bavolet-flottant, La fleurie, Les
moissoneurs, Le carillon de Cythère, and La lugubre. His harmony is
delicate, suggesting that of Mozart and even Chopin, to whom he is
in many ways akin. He does not, like Scarlatti, wander far in the
harmonic field; but in a relatively small compass glides about by
semi-tones. There is, of course, a great deal of tonic and dominant,
such as will always be associated with a certain clear-cut style of
French dance music; but the grace of his melody and his style is too
subtle to permit monotony. The harmonies of the sarabande La
lugubre are profound.

In form he is precise. His use of the rondo deserves special

attention. In this form he cast many of his loveliest pieces, and it is
one which never found a place in the suite. It is very simple, yet in
his hands full of charm. The groundwork of one main theme
recurring regularly after several episodes or contrasting themes was
analyzed in the previous chapter. Couperin called his episodes
couplets, and his rondos are usually composed of the principal
theme and three couplets. He does not invariably repeat the whole
theme after each couplet, but sometimes, as in Les bergeries, only a
characteristic phrase of it. The couplets are generally closely related
to the main theme, from which they differ not in nature, but chiefly in
ornamentation and harmony. Much of the charm of his music is due
to the neat proportions of this hitherto neglected form. It was native
to him as a son of France, where, from the early days of the singers
of Provence, the song in stanzas with its dancing refrain had been
beloved of the people. Through him it found a place in the great
instrumental music of the world.

Couperin’s style is too delicate to be caught in words. To call it the

style galant merely catalogues it as a free style, highly adorned with
agrémens. The freedom is of course the freedom from all trace of
polyphony in the old sense, of strict leading of voices from beginning
to end. Couperin adds notes to his harmonic background when and
where he will; so that it is impossible to say whether a piece is in
two, or three, or four parts, because it is in no fixed number of parts
at all.

The countless agrémens are more than an external feature of his

music, and of other music of his time. The analogies which have
often been drawn between them and the formal superficialities of
court life under the great Louis are in the main false. Both Couperin
and Emanuel Bach, a man of perhaps less sensitive, certainly of less
elegant, taste, regarded them as of vital importance. Even the
learned Kuhnau, who can hardly be called a stylist at all, considered
them the sugar of his fruit. It would seem as if only by means of
these flourishes harpsichord music could take on some grace of line
and warmth of color. Whatever subtlety of expression the dry-toned
instrument was capable of found life only in the agrémens. We
cannot judge of the need of them nor of their peculiar beauties by the
sound of them on the modern pianoforte, even under the lightest
fingers. It is open to question whether any but a few of them should
be retained in the performance of Couperin’s works, now that the
instrument, the shortcomings of which they were intended to
supplement, has been banished in general from the concert stage.

This is not only because the peculiarities of the pianoforte call for a
different kind of ornamentation, but also because the playing of
harpsichord flourishes is practically a lost art. Couperin and Emanuel
Bach left minute directions and explanations in regard to them; but in
their treatises we have only the letter of the law, not the spirit which
inspired it. Even in their day, in spite of all laws, the agrémens were
subject to the caprice of the player; and they remained so down to
the time of Chopin.

Neither the freedom from polyphonic strictness nor the profusion of

ornaments are the special peculiarities of Couperin’s style. They
were more or less common to a great deal of the harpsichord music
of his day. But he had a way, all his own at that time, of
accompanying his melodies with a sort of singing bass or a
melodious inner voice that moved with the melody in thirds or sixths,
or in smooth contrary motion. This may be studied in such pieces as
La fleurie, Le bavolet-flottant, Les moissoneurs, Les abeilles, and
many others. It has little to do with polyphony. The accompanying
voices are only suggested. They never claim attention by their own
movement. They seem a sort of spirit or tinted shadow of the
melody, hardly more than whispering.

This accounts in part for what we may call the tenderness of

Couperin’s music, a quality which makes itself felt no matter how
elusive it may be. He marked most of his pieces to be played with a
special expressiveness, and frequently used the word tendrement.
This, he admitted in one of his prefaces, was likely to surprise those
who were aware of the limitations of the clavecin. He knew that the
‘clavecin was perfect as regards scope and brilliance, but that one
could not increase or diminish the tone on it.’ His thanks would be
forthcoming to one who through taste and skill would be able to
improve its expression in this respect. He was not above all else a
virtuoso. 'J’ayme beaucoup mieux ce qui me touche que ce qui me
surprend,’ he wrote in 1713. There is no doubt that he desired the
greatest refinement of touch and shading in the expression of his
music, and that he suffered under the limitations of the instrument for
which he wrote. For the texture of his music is soft and delicate, its
loveliness has a secret quality, hardly more than suggested by the
shadowy inner voices. We cannot but be reminded of Chopin, in
whose music alone the spirits of music whispered again so softly
together.
Among the contemporaries of Couperin, Marchand, Claude Daquin
(1694-1772), and J. P. Rameau (1683-1764) are best known, at least
by name, today. Marchand is remembered chiefly by reason of the
episode with Bach in Dresden. Daquin enjoyed a brilliant reputation
as an organist in his day. One of his pieces for clavecin—‘The
Cuckoo’—is still heard today. J. B. Weckerlin quoted an amusing
bird-story[19] about Daquin, the burden of which is that one
Christmas eve Daquin imitated the song of a nightingale so perfectly
on the organ in church that the treasurer of the parish dispatched
beadles throughout the edifice in search of a live songster.

Rameau is a greater figure in the general history of music than

Couperin himself; yet, though his harpsichord pieces are, perhaps
therefore, better known than those of the somewhat earlier man,
they lack the most unusual charm and perfection of Couperin’s.
There are fifty-three of these in all. Ten were published in 1706, of
which a gavotte in rondo form in A minor is best known. A second
set of twenty-one pieces appeared in 1724, containing the still
famous Rappel des oiseaux, the Tambourin, Les niais de Sologne,
La poule, the Gavotte with variations, in A minor, and many others.
Sixteen more followed, written between 1727 and 1731. In 1747 a
single piece—La Dauphine—was published. Besides these, all
written originally for harpsichord, he published five arrangements of
his Pièces de concert, written in the first place for a group of three or
more instruments.

Rameau’s style is less delicate than Couperin’s. It is not only that

there are fewer agrémens. The workmanship is more vigorous, more
dramatic; the music itself less intimate. The first gavotte in A minor,
the doubles in the Rigaudon and in Les niais de Sologne, the
variations in the second gavotte in A minor, and La Dauphine, all
speak of a technical enlargement. Yet a certain fineness is lacking. It
will be noticed that he showed hardly more allegiance to the canon
of the suite than Couperin had shown; and there is a large portion of
titles such as Les tendres plaintes, Les soupirs, L’entretien des
muses, and there are also many portraits: La joyeuse, La
triomphante, L’Egyptienne, L’agaçante, and others.
In the preface to the new edition of his works published under the
supervision of Camille Saint-Saëns, there is the following quotation
from Amadée Mereaux’s Les clavecinistes de 1637-1790, which
summarizes his position in the history of harpsichord music. ‘If there
is lacking in his melodies the smoothness of Couperin, the
distinction, the delicacy, the purity of style which give to the music of
that clavecin composer to Louis XIV its so precious quality of charm,
Rameau has at least a boldness of spirit, an animation, a power of
harmony and a richness of modulation. The reflection of his operatic
style, lively, expressive, always precise and strongly rhythmical, is to
be found in his instrumental style. In treatment of the keyboard
Rameau went far ahead of his predecessors. His technical forms, his
instrumental designs, his variety and brilliance in executive
resources, and his new runs and figures are all conquests which he
won to the domain of the harpsichord.’ Rameau is primarily a
dramatic composer. It may be added that several of his harpsichord
pieces later found a place in his operas, usually as ballet music.

IV
A glance over the many pieces of Scarlatti and Couperin discovers a
vast field of unfamiliar music. If one looks deep enough to perceive
the charm, the beauty, the perfection of these forgotten
masterpieces, one cannot but wonder what more than a trick of time
has condemned them to oblivion. For no astonished enthusiasm of
student or amateur whose eye can hear, renders back glory to music
that lies year after year silent on dusty shelves. The general ear has
not heard it. The general eye cannot hear it as it can scan the
ancient picture, the drama, the poetry of a time a thousand or two
thousand years ago. Music that is silent is music quite forgotten if not
dead.

And, what is more, the few pieces of Couperin which are still heard
seem almost to live on sufferance, as if the life they have were not of
their own, but lent them by the listener disposed to imagine a
courtier’s life long ago washed out in blood. ‘Sweet and delicate,’
one hears of the music of Couperin, as one hears of some bit of old
lace or old brocade, that has lain long in a chest of lavender. Yet the
music of Couperin is far more than a matter of fashion. It is by all
tokens great art. The lack is in the race of musicians and of men who
have lost the art of playing it and the simplicity of attentive listening.

To a certain extent the music of Sebastian Bach suffers from the

same lack. On the other hand, the spirit of his music is perennial and
it holds a rank in the modern ear far above that held by any other
harpsichord music. Apart from indefinable reasons of æsthetic worth
there are other reasons why Bach’s music, at any rate a
considerable part of it, is still with us.

In the first place, the style of its texture is solid. Instead of being
crushed, as Couperin’s music is, by the heavy, rich tone of the
modern pianoforte, it seems to grow stronger by speaking through
the stronger instrument. Bach’s style is nearly always an organ style,
whether he is writing for clavichord, for chorus, for bands or strings.
It is very possible that a certain mystical, intimate sentiment which is
innate in most of his clavichord music cannot find expression through
the heavy strings of the pianoforte. This may be far dearer than the
added depth and richness which the pianoforte has, as it were,
hauled up from the great reservoirs of music he has left us. But it is
none the less true that the high-tensioned heavy strings on their
gaunt frame of cast iron need not call in vain on the music of Bach to
set the heart of them vibrating.

In the second place, the two-and three-part ‘Inventions,’ and the

preludes and fugues in the ‘Well-Tempered Clavichord’ have proved
themselves to be, as Bach himself hoped, the very best of teaching
or practice pieces. It is not that your conventional Mr. Dry-as-dust
teacher has power to inflict Bach upon every tender, rebellious
generation. It is rather that the pieces themselves cannot be excelled
as exercises, not only for the fingers but for the brain. One need not
delve here into the matter of their musical beauty, but one must
pause in amazement before their sturdiness, which can stand up,
still resilient, under the ceaseless hammering of ten million sets of
fingers. Clementi and Czerny are being pounded into insensibility;
Cramer, despite the recommendations of Beethoven, is breathing his
last; Moscheles, Dohler, Kalkbrenner, and a host of others are laid to
rest. But here comes Bach bobbing up in our midst seeming to say:
‘Hit me! Hit me as hard as you like and still I’ll sing. And when you
know me as well as I know you, you’ll know how to play the piano.’
So Bach has been, is, and will be introduced to young people. He
inspires love, or hate, or fear—a triple claim to remembrance.

In the third place, there is an intellectual complexity in his music

which, as a triumph of human skill over the masses of sound,
deserves and has won an altar with perpetual flame. And the marvel
is that this skill is rarely used as an end in itself, but as a means of
expressing very genuine and frank emotion. Here we come upon
perhaps the great reason of Bach’s immortality—the warmth of his
music. It is almost uniquely personal and subjective. In it he poured
forth his whole soul with a lack of self-consciousness and a complete
concentration. His was a powerful soul, always afire with
enthusiasm; and his emotion seems to have clarified and crystallized
his music as heat and pressure have made diamonds out of carbon.

Bach was a lovable man, but a stern and somewhat bellicose one as
well. He was shrewd enough to respect social rank quite in the
manner of his day, as the dedication of the Brandenburg concertos
plainly shows; but the records of his various quarrels with the
municipal authorities of Leipzig prove how quick he was to
unrestrained wrath whenever his rights either as man or artist were
infringed upon. A great deal of independence marked him. The same
can hardly be said either of Scarlatti or of Couperin, the one of whom
was lazy and good-natured, the other gently romantic and extremely
polite. Scarlatti rather enjoyed his indifference to accepted rules of
composition; and there was nothing either of self-abasement or of
self-depreciation in Couperin; but both lacked the stalwart vigor of
Bach. Scarlatti aimed, confessedly, to startle and to amuse by his
harpsichord pieces. He cautioned his friends not to look for anything
particularly serious in them. It is hard to dissociate an ideal of pure
and only faintly colored beauty from Couperin. But in the music of
Bach one seldom misses the ring of a strong and even an impetuous
need of self-expression. In the mighty organ works, and in the vocal
works, one may believe with him that he sang his soul out to the
glory of his Maker; but in the smaller keyboard pieces sheer delight
in expressing himself is unmistakable.

It is this that makes Bach a romanticist, while Couperin, with all his
fanciful titles, is classic. It is this that made Bach write in nearly the
same style for all instruments, drawing upon his personal inspiration
without consideration of the instrument for which he wrote; while
Couperin, exquisitely sensitive to all external impressions, forced his
fine art to conformity with the special and limited qualities of the
instrument for which he wrote the great part of his music. And, finally,
it is this which produced utterance of so many varied moods and
emotions in the music of Bach; while in the music of Couperin we
find all moods and emotions tempered to one distinctly normal cast
of thought.

Bach has been the subject of so much profound and special study
that there is little to be added to the explanation of his character or of
his works. In considering him as a composer for the harpsichord or
clavichord, one has to bear two facts in mind: that he was a great
player and a great teacher.

There is much evidence from his son and from prominent musicians
who knew him, that the technical dexterity of his fingers was
amazing. He played with great spirit and, when the music called for
it, at a great speed. Perhaps the oft-repeated story of his triumph
over the famous French player, Marchand, who, it will be
remembered, defaulted at the appointed hour of contest, has been
given undue significance. As we have had occasion to remark, in
speaking of the contest between Handel and D. Scarlatti, such
tourneys at the harpsichord were tests of wits, not of fingers. Bach
was first of all an organist and it may be suggested, with no
disloyalty to the great man among musicians, that he played the
harpsichord with more warmth than glitter. We find little evidence in
his harpsichord music of the sort of virtuosity which makes D.
Scarlatti’s music astonish even today; or, it may be added, of the
special flexible charm which gives Couperin’s its inimitable grace.

Bach is overwhelming as a virtuoso in his organ music, especially in

passages for the pedals. In his harpsichord music he achieves a
rushing, vigorous style. It must not be overlooked that Bach wrote
also for the clavichord, quite explicitly, too. Most of the Forty-eight
Preludes and Fugues are distinctly clavichord, not harpsichord,
music. That is to say, they require a fine shading which is impossible
on the harpsichord. When he wrote for the harpsichord he had other
effects in mind. The prelude of the English suite in G minor or the
last movement of the Italian concerto may be taken as
representative of his most vigorous and effective harpsichord style.
They are different not only in range and breadth, but in spirit as well,
from practically all of the ‘Well-Tempered Clavichord.’ Nevertheless,
though these may be taken fairly as examples of his harpsichord
style at its best and strongest, they are not especially effective as
virtuoso music. There is sheer virtuosity only in the Goldberg
Variations.

To Bach as a teacher we owe the Inventions and the ‘Well-Tempered

Clavichord,’ both written expressly for the use and practice of young
people who wished to learn about music and to acquire a taste for
the best music. Volumes might well be filled with praise of them. It
will suffice us only to note, however, that to master the technical
difficulties of the keyboard was always for Bach only a step toward
the art of playing, which is the art of expressing emotion in music.
These two sets of pieces are all-powerful evidence of this—his creed
—in accordance with which he always nobly lived and worked. They
have but one parallel in pianoforte music: the Études of Chopin. The
‘Well-Tempered Clavichord’ is, and always will be, essentially a study
in expression.

His system of tempering or tuning the clavichord, by reason of which

he has often been granted a historical immortality, was the relatively
simple one of dividing the octave into twelve equal intervals. Only the
octave itself was strictly in tune, but the imperfections of the other
intervals were so slight as to escape detection by the most practised
ear. By paying the nominal toll of theoretical inaccuracy, Bach
opened the roads of harmonic modulation on every hand. It must not
be forgotten, however, that most of the pieces of Couperin or
Scarlatti, not to mention many an outlandish chromatic tour de force
in the works of the early English composers, would have been
intolerable on a harpsichord strictly in tune. Other men than Bach
had their systems of temperament. We may take Bach’s only to be
the simplest.

Furthermore, that he created a new development of pianoforte

technique by certain innovations in the manner of fingering
passages, is open to question. It is well known that up to the
beginning of the eighteenth century the use of the thumb on the
keyboard was generally discountenanced. Bach himself had seen
organists play who avoided using the thumb even in playing wide
stretches. Scales were regularly played by the fingers, which, without
the complement of the thumb, passed sideways over each other in a
crawling motion which is said to have been inherited from the
lutenists. Couperin advocated the use of the thumb in scales, but
over, and not under, the fingers. Bach seems the first to have openly
advised and practised passing the thumb under the fingers in the
manner of today. Yet even he did not give up entirely the older
method of gliding the fingers over each other in passages up and
down the keyboard.

His system passed on through the facile hands of his son Emanuel,
the greatest teacher of the next generation; and if it is not the crest of
the wave of new styles of playing which was to break over Europe
and flood a new and special pianoforte literature, is at any rate a
considerable part of its force. Yet it must be borne in mind that
Scarlatti founded by his own peculiar gifts a tradition of playing the
piano and composing for it, in which Clementi was to grow up; and
that, influential as Emanuel Bach was, Clementi was the teacher of
the great virtuosi who paved the way to Chopin, the composer for the
piano par excellence.
The foundation of all Bach’s music is the organ. Even in his works for
violin alone, or in those for double chorus and instruments, the
conjunct, contrapuntal style of organ music is unmistakable. His
general technique was acquired by study of the organ works of his
great predecessors, Frescobaldi, Sweelinck, Pachelbel, Buxtehude,
Bohm, and others. He was first and always an organist. So it is not
surprising to find by far the greater part of his harpsichord and
clavichord music shaped to a polyphonic ideal; and, what is more,
written in the close, smooth style which is primarily fitting to the
organ.

His intelligence, however, was no less alert than it was acute. There
is evidence in abundance that he not only knew well the work of
most of his contemporaries, but that he appropriated what he found
best in their style. He seems to have found the violin concertos of
Vivaldi particularly worthy of study. He was indebted to him for the
form of his own concertos; and, furthermore, he adapted certain
features of Vivaldi’s technique of writing for the violin to the
harpsichord. Of the influence of Couperin there is far less than was
once supposed. The ‘French Suites’ were not so named by Bach
and are, moreover, far more in his own contrapuntal style than in the
tender style of Couperin. Kuhnau’s Bible sonatas are always cited as
the model for Bach’s little Capriccio on the departure of his brother;
but elsewhere it is hard to find evidence of indebtedness to Kuhnau.

But he even profited by an acquaintance with the trivial though

enormously successful Italian opera of his day, and used the da
capo aria as frankly as A. Scarlatti or J. A. Hasse. Still, whatever he
acquired from his contemporaries was but imposed upon the great
groundwork of his art, his organ technique. He never let himself go
upon the stream of music of his day, but held steadfast to the ideal
he had inherited from a century of great German organists, of whom
he was to be the last and the greatest.

So, for the most part, the forms which had evolved during the
seventeenth century were the forms in which he chose to express
himself. Of these, two will be for ever associated with him, because