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vi Contents
Evidence for the Physical Basis of 8.3 Aneuploidy Is an Increase or Decrease in the
Recombination 174 Number of Individual Chromosomes 222
Predicting the Outcomes of Crosses with Linked Types of Aneuploidy 222
Genes 175 Effects of Aneuploidy 223
Testing for Independent Assortment 176 Aneuploidy in Humans 224
Gene Mapping with Recombination Uniparental Disomy 227
Frequencies 178 Mosaicism 228
Constructing a Genetic Map with the Use of 8.4 Polyploidy Is the Presence of More than Two
Two-Point Testcrosses 179 Sets of Chromosomes 228
Autopolyploidy 228
7.3 A Three-Point Testcross Can Be Used to Map
Allopolyploidy 229
Three Linked Genes 180
The Significance of Polyploidy 232
Constructing a Genetic Map with the Three-Point
Testcross 181 Chapter 9 Bacterial and Viral
CONNECTING CONCEPTS Stepping Through the Three-Point Genetic Systems 241
Cross 186
Life in a Bacterial World 241
Effect of Multiple Crossovers 188
9.1 Genetic Analysis of Bacteria Requires Special
Mapping Human Genes 189 Methods 242
Mapping with Molecular Markers 190 Bacterial Diversity 242
Genes Can Be Located with Genomewide Techniques for the Study of Bacteria 243
Association Studies 191 The Bacterial Genome 244
7.4 Physical-Mapping Methods Are Used to Plasmids 245
Determine the Physical Positions of Genes on 9.2 Bacteria Exchange Genes Through
Particular Chromosomes 192 Conjugation, Transformation, and
Somatic-Cell Hybridization 192 Transduction 247
Conjugation 247
Deletions Mapping 194
Natural Gene Transfer and Antibiotic
Physical Chromosome Mapping Through
Resistance 254
Molecular Analysis 195
Transformation in Bacteria 254
7.5 Recombination Rates Exhibit Extensive Bacterial Genome Sequences 256
Variation 195 Horizontal Gene Transfer 256
v
viii Contents
9.3 Viruses Are Simple Replicating Systems 11.2 Eukaryotic Chromosomes Possess
Amenable to Genetic Analysis 257 Centromeres and Telomeres 306
Techniques for the Study of Bacteriophages 257 Centromere Structure 306
Transduction: Using Phages to Map Bacterial Telomere Structure 307
Genes 258
11.3 Eukaryotic DNA Contains Several Classes of
CONNECTING CONCEPTS Three Methods for Mapping
Sequence Variation 308
Bacterial Genes 261
The Denaturation and Renaturation
of DNA 308
Gene Mapping in Phages 261 Types of DNA Sequences in Eukaryotes 308
Fine-Structure Analysis of Bacteriophage
11.4 Organelle DNA Has Unique
Genes 263
Characteristics 309
RNA Viruses 265
Mitochondrion and Chloroplast Structure 309
Human Immunodeficiency Virus and AIDS 267
The Endosymbiotic Theory 310
Influenza 268
Uniparental Inheritance of Organelle-Encoded
Traits 311
Chapter 10 DNA: The Chemical The Mitochondrial Genome 314
Nature of the Gene 277 The Evolution of Mitochondrial DNA 316
Damage to Mitochondrial DNA Is Associated with
Arctic Treks and Ancient DNA 277
Aging 316
10.1 Genetic Material Possesses Several Key The Chloroplast Genome 317
Characteristics 278 Through Evolutionary Time, Genetic Information
10.2 All Genetic Information Is Encoded in the Has Moved Between Nuclear, Mitochondrial, and
Structure of DNA or RNA 278 Chloroplast Genomes 318
Early Studies of DNA 278
DNA As the Source of Genetic Information 280 Chapter 12 DNA Replication and
Watson and Crick’s Discovery of the Three- Recombination 325
Dimensional Structure of DNA 283
RNA As Genetic Material 285
Topoisomerase, Replication, and
Cancer 325
10.3 DNA Consists of Two Complementary and
Antiparallel Nucleotide Strands That Form a 12.1 Genetic Information Must Be Accurately
Double Helix 286 Copied Every Time a Cell Divides 326
The Primary Structure of DNA 286 12.2 All DNA Replication Takes Place in a
Secondary Structures of DNA 288 Semiconservative Manner 326
Meselson and Stahl’s Experiment 327
CONNECTING CONCEPTS Genetic Implications of DNA
Modes of Replication 329
Structure 290
Requirements of Replication 332
10.4 Special Structures Can Form in DNA and Direction of Replication 332
RNA 291 CONNECTING CONCEPTS The Direction of Synthesis in
Different Models of Replication 334
Chapter 11 Chromosome Structure
and Organelle DNA 299 12.3 Bacterial Replication Requires a Large
Number of Enzymes and Proteins 334
Telomeres and Childhood Adversity 299 Initiation 334
11.1 Large Amounts of DNA Are Packed into a Unwinding 334
Cell 300 Elongation 336
Supercoiling 300 Termination 339
The Bacterial Chromosome 301 The Fidelity of DNA Replication 339
Eukaryotic Chromosomes 302
CONNECTING CONCEPTS The Basic Rules of Replication 340
Changes in Chromatin Structure 304
Contents ix
Chapter 15 The Genetic Code and Negative and Positive Control: Inducible and
Translation 411 Repressible Operons 448
The lac Operon of E. coli 450
Hutterites, Ribosomes, and lac Mutations 453
Bowen–Conradi Syndrome 411 Positive Control and Catabolite Repression 457
15.1 Many Genes Encode Proteins 412 The trp Operon of E. coli 458
The One Gene, One Enzyme Hypothesis 412 Bacterial Enhancers 460
The Structure and Function of Proteins 415 16.3 Some Operons Regulate Transcription
15.2 The Genetic Code Determines How the Through Attenuation, the Premature
Nucleotide Sequence Specifies the Amino Termination of Transcription 460
Acid Sequence of a Protein 417 Attenuation in the trp Operon of E. coli 460
Breaking the Genetic Code 418 Why Does Attenuation Take Place in the trp
The Degeneracy of the Code 420 Operon? 464
The Reading Frame and Initiation Codons 421 16.4 RNA Molecules Control the Expression of
Termination Codons 422 Some Bacterial Genes 464
The Universality of the Code 422 Antisense RNA 464
Riboswitches 464
CONNECTING CONCEPTS Characteristics of the Genetic
RNA-Mediated Repression Through
Code 422
Ribozymes 466
15.3 Amino Acids Are Assembled into a Protein
Through Translation 422 Chapter 17 Control of Gene
The Binding of Amino Acids to Transfer RNAs 423 Expression in Eukaryotes 473
The Initiation of Translation 424
Genetic Differences That Make Us
Elongation 426
Termination 427
Human 473
17.1 Eukaryotic Cells and Bacteria Have Many
CONNECTING CONCEPTS A Comparison of Bacterial and
Features of Gene Regulation in Common, but
Eukaryotic Translation 430 They Differ in Several Important Ways 474
15.4 Additional Properties of RNA and Ribosomes 17.2 Changes in Chromatin Structure Affect the
Affect Protein Synthesis 430 Expression of Genes 474
The Three-Dimensional Structure of the DNase I Hypersensitivity 474
Ribosome 430 Chromatin Remodeling 475
Polyribosomes 431 Histone Modification 475
Messenger RNA Surveillance 431 DNA Methylation 478
Folding and Posttranslational Modifications of 17.3 The Initiation of Transcription Is Regulated
Proteins 433 by Transcription Factors and Transcriptional
Translation and Antibiotics 433 Regulator Proteins 479
Transcriptional Activators and Coactivators 479
Chapter 16 Control of Gene Transcriptional Repressors 481
Enhancers and Insulators 481
Expression in Bacteria 443
Regulation of Transcriptional Stalling and
Operons and the Noisy Cell 443 Elongation 482
Coordinated Gene Regulation 482
16.1 The Regulation of Gene Expression Is Critical
for All Organisms 444 17.4 Some Genes Are Regulated by RNA
Genes and Regulatory Elements 445 Processing and Degradation 483
Levels of Gene Regulation 445 Gene Regulation Through RNA Splicing 483
DNA-Binding Proteins 446 The Degradation of RNA 484
16.2 Operons Control Transcription in Bacterial 17.5 RNA Interference Is an Important Mechanism
Cells 447 of Gene Regulation 485
Operon Structure 447 Small Interfering RNAs and MicroRNAs 485
Contents xi
Mechanisms of Gene Regulation by RNA CONNECTING CONCEPTS The Basic Pathway of DNA
Interference 486 Repair 524
The Control of Development by RNA
Interference 487 Repair of Double-Strand Breaks 524
Translesion DNA Polymerases 524
17.6 Some Genes Are Regulated by Processes That
Genetic Diseases and Faulty DNA Repair 525
Affect Translation or by Modifications of
Proteins 487
CONNECTING CONCEPTS A Comparison of Bacterial and
Chapter 19 Molecular Genetic
Eukaryotic Gene Control 488 Analysis and Biotechnology 535
Helping the Blind to See 535
Chapter 18 Gene Mutations and 19.1 Techniques of Molecular Genetics Have
DNA Repair 493 Revolutionized Biology 536
The Molecular Genetics Revolution 536
A Fly Without a Heart 493
Working at the Molecular Level 536
18.1 Mutations Are Inherited Alterations in the
DNA Sequence 494 19.2 Molecular Techniques Are Used to Isolate,
Recombine, and Amplify Genes 537
The Importance of Mutations 494
Cutting and Joining DNA Fragments 537
Categories of Mutations 494
Viewing DNA Fragments 539
Types of Gene Mutations 495
Locating DNA Fragments with Southern Blotting and
Phenotypic Effects of Mutations 497
Probes 540
Suppressor Mutations 498
Cloning Genes 541
Mutation Rates 502
Application: The Genetic Engineering of Plants with
18.2 Mutations Are Potentially Caused by a Pesticides 545
Number of Different Factors 503 Amplifying DNA Fragments with the Polymerase
Spontaneous Replication Errors 503 Chain Reaction 546
Spontaneous Chemical Changes 504
Chemically Induced Mutations 506 19.3 Molecular Techniques Can Be Used to Find
Genes of Interest 549
Radiation 508
Gene Libraries 549
18.3 Mutations Are the Focus of Intense Study by In Situ Hybridization 552
Geneticists 509 Positional Cloning 552
Detecting Mutations with the Ames Test 509 Application: Isolating the Gene for Cystic Fibrosis 553
Radiation Exposure in Humans 510
19.4 DNA Sequences Can Be Determined and
18.4 Transposable Elements Cause Mutations 511 Analyzed 555
General Characteristics of Transposable Elements 511 Restriction Fragment Length Polymorphisms 555
Transposition 512 DNA Sequencing 556
The Mutagenic Effects of Transposition 513 Next-Generation Sequencing Technologies 559
Transposable Elements in Bacteria 514 DNA Fingerprinting 561
Transposable Elements in Eukaryotes 515 Application: Identifying People Who Died in the
Collapse of the World Trade Center 562
CONNECTING CONCEPTS Types of Transposable
Elements 519 19.5 Molecular Techniques Are Increasingly Used
to Analyze Gene Function 563
Transposable Elements Have Played an
Forward and Reverse Genetics 563
Important Role in Genome Evolution 520
Creating Random Mutations 564
18.5 A Number of Pathways Repair Changes in Site-Directed Mutagenesis 564
DNA 520 Transgenic Animals 565
Mismatch Repair 520 Knockout Mice 566
Direct Repair 522 Silencing Genes with RNAi 567
Base-Excision Repair 522 Application: Using RNAi to Treat Human
Nucleotide-Excision Repair 523 Disease 568
xii Contents
19.6 Biotechnology Harnesses the Power of 21.2 Several Molecular Processes Lead to
Molecular Genetics 569 Epigenetic Changes 615
Pharmaceutical Products 569 DNA Methylation 615
Specialized Bacteria 569 Histone Modifications 617
Agricultural Products 569 Epigenetic Effects by RNA Molecules 618
Genetic Testing 570
21.3 Epigenetic Processes Produce a Diverse Set of
Gene Therapy 571 Effects 619
Paramutation 619
Chapter 20 Genomics and
Behavioral Epigenetics 621
Proteomics 579 Epigenetic Effects of Environmental Chemicals 623
Decoding the Waggle Dance: The Genome Transgenerational Epigenetic Effects on
of the Honeybee 579 Metabolism 623
Epigenetic Effects in Monozygotic Twins 623
20.1 Structural Genomics Determines the DNA
X Inactivation 623
Sequences of Entire Genomes 580
Epigenetic Changes Associated with Cell
Genetic Maps 580
Differentiation 625
Physical Maps 581
Genomic Imprinting 626
Sequencing an Entire Genome 582
The Human Genome Project 583 21.4 The Epigenome 627
Single-Nucleotide Polymorphisms 587
Copy-Number Variations 588 Chapter 22 Developmental
Sequence-Tagged Sites and Expressed-Sequence Genetics and Immunogenetics 633
Tags 589
Bioinformatics 589 The Origin of Spineless Sticklebacks 633
Metagenomics 590 22.1 Development Takes Place Through Cell
Synthetic Biology 591 Determination 634
20.2 Functional Genomics Determines the Cloning Experiments on Plants 635
Function of Genes by Using Genomic-Based Cloning Experiments on Animals 635
Approaches 591 22.2 Pattern Formation in Drosophila Serves
Predicting Function from Sequence 591 As a Model for the Genetic Control of
Gene Expression and Microarrays 592 Development 636
Gene Expression and Reporter Sequences 595 The Development of the Fruit Fly 636
Genome-Wide Mutagenesis 595 Egg-Polarity Genes 637
20.3 Comparative Genomics Studies How Segmentation Genes 640
Genomes Evolve 596 Homeotic Genes in Drosophila 640
Prokaryotic Genomes 596 Homeobox Genes in Other Organisms 642
Eukaryotic Genomes 598
CONNECTING CONCEPTS The Control Of Development 643
Comparative Drosophila Genomics 601
The Human Genome 601 Epigenetic Changes in Development 643
20.4 Proteomics Analyzes the Complete Set of 22.3 Genes Control the Development of Flowers in
Proteins Found in a Cell 603 Plants 644
Determination of Cellular Proteins 603 Flower Anatomy 644
Affinity Capture 604 Genetic Control of Flower Development 644
Protein Microarrays 604
CONNECTING CONCEPTS Comparison of Development in
Structural Proteomics 605
Drosophila and Flowers 646
Chapter 21 Epigenetics 613
22.4 Programmed Cell Death Is an Integral Part of
How Your Grandfather’s Diet Could Affect Development 646
Your Health 613
22.5 The Study of Development Reveals Patterns
21.1 What Is Epigenetics? 614 and Processes of Evolution 647
Contents xiii
25.4 Several Evolutionary Forces Potentially Cause 26.4 The Evolutionary History of a Group of
Changes in Allelic Frequencies 726 Organisms Can Be Reconstructed by Studying
Mutation 726 Changes in Homologous Characteristics 756
Migration 727 The Alignment of Homologous Sequences 757
Genetic Drift 728 The Construction of Phylogenetic Trees 758
Natural Selection 731 26.5 Patterns of Evolution Are Revealed by
CONNECTING CONCEPTS The General Effects of Forces That
Molecular Changes 758
Change Allelic Frequencies 736 Rates of Molecular Evolution 759
The Molecular Clock 760
Evolution Through Changes in Gene Regulation 761
Chapter 26 Evolutionary Genome Evolution 762
Genetics 743
Taster Genes in Spitting Apes 743 Reference Guide to Model Genetic
26.1 Evolution Occurs Through Genetic Change Organisms A1
Within Populations 744 The Fruit Fly Drosophilia melanogaster A2
26.2 Many Natural Populations Contain High The Bacterium Escherichia coli A4
Levels of Genetic Variation 745 The Nematode Worm Caenorhabditis elegans A6
Molecular Variation 745 The Plant Arabidopsis thaliana A8
Protein Variation 746 The Mouse Mus musculus A10
DNA Sequence Variation 747 The Yeast Saccharomyces cerevisiae A12
26.3 New Species Arise Through the Evolution of
Reproductive Isolation 749
Glossary B1
The Biological Species Concept 749
Reproductive Isolating Mechanisms 750 Answers to Selected Questions
Modes of Speciation 751 and Problems C1
Genetic Differentiation Associated with
Speciation 755 Index D1
Letter from the Author
I still remember the excitement I felt when I took my first genetics course. I was
intrigued by the principles of heredity,
which allow one to predict what offspring will look like even before they are
born. I was fascinated to learn that these principles have their foundation in
the chemistry of an elegant molecule called DNA. And I was captivated to find
that genetics underlies the process of evolution, which is responsible for the
endless diversity and beauty of life. These elements of genetics still impress and
excite me today. One of the great things about teaching genetics is the chance to
convey that excitement to students.
Twenty-three years ago, I set out to write a new genetics textbook. My vision
was to create a book that conveys the excitement of genetics, that motivates
students, and that focuses on concepts and problem solving. Those were the
original objectives of the first edition of Genetics: A Conceptual Approach and
they remain the core features of this new fifth edition of the book.
In this book, I’ve tried to share some of what I’ve learned in my 33 years of
teaching genetics. I provide advice and encouragement at places where students
often have difficulty, and I tell stories of the people, places, and experiments of
genetics—past and present—to keep the subject relevant, interesting, and alive.
My goal is to help you learn the necessary details, concepts, and problem-
solving skills, while encouraging you to see the elegance and beauty of the
larger landscape.
At Southwestern University my office door is always open, and my students often drop
by to share their own approaches to learning, things that they have read about genetics,
and their experiences, concerns, and triumphs. I learn as much from my students as they
learn from me, and I would love to learn from you by email ([email protected]),
by telephone (512-863-1974), or in person (Southwestern University, Georgetown, Texas).
Ben Pierce
PROFESSOR OF BIOLOGY AND
HOLDER OF THE LILLIAN NELSON PRATT CHAIR
SOUTHWESTERN UNIVERSITY
xv
Preface
The main goals of Genetics: A Conceptual Approach have always been to help students un-
cover and make connections between the major concepts of genetics. Throughout the four
preceding editions of this book the accessible writing style, simple and instructive illustra-
tions, and useful pedagogical features throughout the book have helped students develop a
fuller understanding of genetics.
Hallmark Features
■ Key Concepts and Connections Throughout the book, I’ve included features to help
students focus on the major concepts of each topic.
■ Concept boxes throughout each chapter summarize the key
blood-type O is 1/2, what is the probability of being either blood-type been a favorite feature for both students and instructors. In addition
A or blood-type O? to carefully walking students through each major concept of genetics,
a. 5/8 c. 1/10 I invite them into the topic with an introductory story. These stories
b. 1/2 d. 1/16 include relevant examples of disease or other biological phenomena to
give students a sample of what they’ll be learning in a chapter. More
than a third of the introductory stories in this edition are new.
■ Clear, Simple Illustration Program The attractive and instructive figures have
proved to be an effective learning tool for students throughout the past four editions
and continue to be signature feature of the new edition. Each figure was carefully
rendered to highlight main points and to step the reader through experiments
and processes. Most figures include text that walks students through the graphical
presentation. Illustrations of experiments reinforce the scientific method by first
proposing a hypothesis, then pointing out the methods and results, and ending with a
conclusion that reinforces concepts explained in the text.
■ Emphasis on Problem Solving One of the things that I’ve learned in my 33
In choosing 0.05 as the cutoff value, scientists have
agreed to assume that chance is responsible for the devia- years of teaching is that students learn genetics best through problem solving.
tions between observed and expected values unless there is Working through an example, equation, or experiment helps students see
strong evidence to the contrary. Bear in mind that, even if concepts in action and reinforces the ideas explained in the text. In the book,
we obtain a probability of, say, 0.01, there is still a 1% prob-
ability that the deviation between the observed and the ex- I help students develop problem-solving skills in a number of ways. Newly
pected numbers is due to nothing more than chance. Calcu- reformatted Worked Problems (see New and Reorganized Content on p. xix)
lation of the chi-square value is illustrated in Figure 3.14. walk students through each step of a difficult concept. Problem Links spread
TRY PROBLEM 38
throughout each chapter point to end-of-chapter problems that students can
work to test their understanding of the material they have just read, all with
xvi
Preface xvii
answers in the back of the book so students can check their results. I provide a
wide range of end-of-chapter problems, organized by chapter section and split into
Comprehension Questions, Application Questions and Problems, and Challenge
Questions. Some of these questions draw on examples from published papers and are
marked by a data analysis icon.
TEXT MEDIA
Reading Quizzes
Students Read
Chapter / View Nature Genetics
Branched Tutorial Animation Activity
article with assessment
LaunchPad Text Icon
simulations now include assessment questions at the end, to help students evaluate
whether they understood the concept or technique they viewed.
■ Comprehensively Revised Assessment has undergone extensive rewriting,
reviewing, and accuracy checking.
■ Online End-of-Chapter Problems include nearly all of the end-of-chapter
problems from the text converted into self-graded multiple-choice and drop-down
menu format for quizzing, homework, or student practice.
■ LearningCurve allows students to test their comprehension of the chapter
chapter.
■ Nature Genetics Articles with Assessment engage students with primary research
and encourage critical thinking. Specifically selected for both alignment with text
coverage and exploration of identified difficult topics, the Nature Genetics articles
include assessment questions that can be automatically graded. Some of the open-ended
(non-multiple choice) questions are also suitable for use in flipped classrooms and
active learning discussions either in class or online.
All of the new media resources for Genetics: A Conceptual Approach will be available in our
LaunchPad system.
LaunchPad is a dynamic, fully integrated learning environment that brings together all
of our teaching and learning resources in one place. It also contains the fully interactive
e-Book and other newly updated resources for students and instructors, including the
following:
■ Newly Updated Clicker Questions allow instructors to integrate active learning in
the classroom and to assess students’ understanding of key concepts during lectures.
Available in Microsoft Word and PowerPoint, numerous questions are based on the
Concept Check questions featured in the textbook.
■ New Lecture PPTs have been developed to minimize preparation time for new
users of the book. These files offer suggested lectures including key illustrations and
summaries that instructors can adapt to their teaching styles.
■ Layered PPTs deconstruct key concepts, sequences, and processes from the
has been fully optimized to increase type sizes and adjust color saturation. These
images have been tested in a large lecture hall to ensure maximum clarity and
visibility.
■ The Solutions and Problem-Solving Manual (written by Jung H. Choi, Georgia
a Solution Strategy and What information is required in your answer to the because gray is recessive. Thus the cross is:
Solution Steps for each problem?
Gg Ž gg
The probability that in a litter of six kittens, three will be
worked problem. Hint black and three will be gray.
Black female Gray male
887n
Acknowledgments
I am indebted to many people for help with this and previous editions of Genetics: A Con-
ceptual Approach. I learned much from my genetics teachers: Ray Canham, who first ex-
posed me to genetics and instilled in me a lifelong love for the subject; and Jeff Mitton, who
taught me the art of genetic research. I’ve learned from the thousands of genetics students
who have filled my classes over the past 33 years, first at Connecticut College, then at Baylor
University, and now at Southwestern University. Their intelligence, enthusiasm, curiosity,
and humor have been a source of motivation and pleasure throughout my professional life.
I have also learned from students worldwide who have used earlier editions of this book and
kindly shared with me—through emails and phone calls—their thoughts about the book
and how it could be improved.
I am grateful for the wonderful colleagues who surround me daily at Southwestern Uni-
versity and whose friendship, advice, and good humor sustain my work. The small classes,
close interaction of students and faculty, and integration of teaching and research have
made working at Southwestern University personally and professionally rewarding. I thank
James Hunt, Provost of Southwestern University and Dean of the Brown College, for help-
ing to create this supportive academic environment and for his continued friendship and
collegiality.
Writing a modern science textbook requires a team effort and I have been blessed with an
outstanding team at W. H. Freeman and Company. Life Sciences Publisher Susan Winslow
has been a champion of the book for a number of years; I value her creativity, insight, and
support. Lauren Schultz, Senior Acquisitions Editor for the Life Sciences team was a won-
derful shepherd of the project, providing ideas, encouragement, and support throughout
the project. It was a pleasure working with Developmental Editor Anna Bristow, my daily
partner in crafting this edition of Genetics. Anna is a superb editor with outstanding orga-
nizational skills and an eye for detail; she kept the project on track and contributed to this
edition in innumerable ways.
Senior Project Editor Georgia Lee Hadler at W. H. Freeman expertly managed the pro-
duction of this fifth edition, as well as all preceding editions. Her dedication to excellence in
all phases of the production process has been a major factor in making the book a success.
Jeanine Furino served as copy editor and provided valuable editorial suggestions. I thank
Dragonfly Media Group for creating and revising the book’s illustrations and Matthew Mc-
Adams for coordinating the illustration program. Thanks to Paul Rohloff at W. H. Freeman
and Assunta Petrone at codeMantra for coordinating the composition and manufacturing
Preface xxi
phases of production. Diana Blume developed the book’s design and the cover for this edi-
tion. I thank Christine Buese and Jacqui Wong for photo research. Allison Michael, Elaine
Palucki, Alexi Garrett, Adam Feil, and Chris Efstratiou developed the excellent media and
supplements that accompany the book. I am grateful to Jung Choi and Mark McCallum for
writing solutions to new end-of-chapter problems. Joseph Ahlander, Ellen France, Robert
Fowler, Brian Kreiser, Joshua Loomis, Amy McMillan, Marcie Moehnke, Douglas Thrower,
and Daniel Williams developed and reviewed assessment questions.
I am grateful to the W. H. Freeman sales representatives, regional managers, and region-
al sales specialists, who introduce my book to genetics instructors throughout world. I have
greatly enjoyed working with this sales staff; their expertise, hard work, and good service
are responsible for the success of Freeman books.
A number of colleagues served as reviewers of this book, kindly lending me their tech-
nical expertise and teaching experience. Their assistance is gratefully acknowledged. Any
remaining errors are entirely my own.
Marlene Tyrrell—my spouse and best friend for 33 years—and our children and their
spouses—Sarah, Matt, Michael, and Amber—provide love, support, and inspiration for
everything that I do.
My gratitude goes to the reviewers of this new edition of Genetics: A Conceptual Approach.
Amy Abdulovic-Cui Spencer Benson Sarah Crawford Laura Frost
Augusta State University University of Maryland Southern Connecticut State Point Park University
Joseph Ahlander Edward Berger University J. Yvette Gardner
Northeastern State University Dartmouth College Marilyn Cruz-Alvarez Clayton State University
David Aiello Laura Bermingham Florida Gulf Coast University William Gilliland
Austin College University of Vermont Sandra Davis DePaul University
Preston Aldrich University of Indianapolis
Aimee Bernard Elliott S. Goldstein
Benedictine University University of Colorado Denver Sarwan Dhir Arizona State University
Kirk Anders Fort Valley State University
Indrani Bose Steven W. Gorsich
Gonzaga University Western Carolina University David Donnell Central Michigan University
Jeanne Andreoli The Citadel
John Braverman Anjali Gray
Marygrove College Saint Joseph’s University Chuanguang Du Lourdes University
Anthony Arment Montclair State University
David Buchanan Bradley Hersh
Central State University Cheryld L. Emmons
North Dakota State University Allegheny College
Minoo Askari Alfred University
Gerald L. Buldak Debra Hinson
Pellissippi State Community William Ettinger
Loyola University Chicago Dallas Baptist University
College Gonzaga University
Alyssa C. Bumbaugh Peter Hoffman
Andrea Bailey Sarah Evans
Penn State University Notre Dame of Maryland
Brookhaven Community Friends University
Dan Caprioglio University
College Victor Fet
Colorado State University Marshall University Margaret Hollingsworth
Paul W. Bates
Maria V. Cattell University at Buffalo
University of Minnesota Ted Fickel
Duluth University of Colorado American Jewish University Carina Endres Howell
Richard Duhrkopf Lock Haven University
Keith Barlow Robert G. Fowler
Guilford Technical Community Baylor University San Jose State University Li Huang
College Joel Chandlee Montana State University
Thomas Fowler
Phillip Barnes University of Rhode Island Southern Illinois University Mary Huff
Connecticut College Henry Chang Edwardsville Bellarmine University
Christine Beatty Purdue University Edison Fowlks Colin Hughes
Benedictine University Cynthia Church Hampton University Florida Atlantic University
John Belote Metropolitan State University Dennis Frisby Jeffrey Hughes
Syracuse University of Denver Cameron University Millikin University
xxii Preface
C
ARRIAGES by the score were waiting at the fine Corinthian
entrance to the Legitimate, when Harold and Archie reached
the theatre in their hansom. The façade of the Legitimate
Theatre is so severely Corinthian that foreign visitors invariably ask
what church it is.
It was probably the classical columns supporting the pediment of
the entrance that caused Archie to abate his frivolous conversation
with his friend in the hansom—Archie had been expressing the
opinion that it was exhilarating—only exhilarating was not the word
he used—to swear at a man who had once been a clergyman and
who still wore the dress of a cleric. “A chap feels that his turn has
come,” he had said. “No matter how wrong they are you can’t swear
at them and tell them to come down out of that, when they’re in
their own pulpits—they’d have you up for brawling. That’s why I like
to take it out of old Playdell. He tells me, however, that there’s no
dean in the Church that gathers in the decimals as he does in my
shop. But, bless you! he saves me his screw three times over.”
But now that the classical front of the Legitimate came in view,
Archie became solemn.
He possibly appreciated the feelings of a conscientious clergyman
when about to enter his Church.
Shakespeare was a great responsibility.
So was Mrs. Mowbray.
The performance was not quite over; but before Archie had paid
the hansomeer, the audience was streaming out from every door.
“Stand here and listen to what the people are saying.” whispered
Archie. “I often do it. It is only in this way that you can learn how
much appreciation for Shakespeare still remains in England.”
He took up his position with Harold at the foot of the splendid
staircase of the theatre, where the people chatted together while
waiting for their carriages.
With scarcely an exception, the remarks had a hearing upon the
performance of “Cymbeline.” Only two ladies confined their criticisms
to their respective medical advisers.
Of the others, one man said that Mrs. Mowbray bore a striking
resemblance to her photographs.
A second said that she was the most beautiful woman in England.
A third said that she knocked sparks out of Polly Floss in the same
line of business. (Polly Floss was the leading exponent of burlesque).
One woman said that Mrs. Mowbray was most picturesquely
dressed.
A second said that she was most picturesquely undressed.
A third wondered if Liberty had got the exact tint of the robe that
Mrs. Mowbray had worn in the second act.
“And yet some people say that there’s no appreciation of
Shakespeare in England!” said Archie, as he led Harold round the
stalls, over which the attendants were spreading covers, and on to
Mrs. Mowbray’s private rooms.
“From the crowds that went out by every door, I judge that the
theatre is making money, at any rate; and I suppose that’s the most
practical test of appreciation,” said Harold.
“Oh, they don’t all pay,” said Archie. “That’s a feature of theatrical
management that it takes an outsider some time to understand. Mrs.
Mowbray should understand it pretty well by this time, so should her
business manager. I’m just getting to understand it.”
“You mean to say that the people are allowed to come in without
paying?”
“It amounts to that in the long run—literally the long run—of the
piece, I believe. Upon my soul, there are some people who fancy
that a chap runs a show as a sort of free entertainment for the
public. The dramatic critics seem to fancy that a chap produces a
play, simply in order to give them an opportunity of showing off their
own cleverness in slating it. It seems that a writer-chap can’t show
his cleverness in praising a piece, but only in slanging it.”
“I think that I’d try and make people pay for their seats.”
“I used always to pay for mine in the old days—but then, I was
always squandering my money.”
“I have always paid for mine.”
“The manager says that if you asked people to pay, they’d be
mortally offended and never enter the theatre again, and where
would you be then?”
“Where, indeed?” said Harold. “I expect your manager must know
his business thoroughly.”
“He does. It requires tact to get people to come to see
Shakespeare,” said Archie. “But a chap can’t build a monument for
himself without paying for it.”
“It would be ridiculous to expect it,” said Harold.
Pushing aside a magnificent piece of heavy drapery, Archie
brought his friend into a passage illuminated by the electric light;
and knocking at a door at the farther end, he was admitted by Mrs.
Mowbray’s maid, into a prettily-furnished sitting-room and into the
presence of Mrs. Mowbray, who was sitting robed in something very
exquisite and cloud-like—not exactly a peignoir but something that
suggested a peignoir.
She was like a picture by Romney. If one could imagine all the
charm of all the pictures of Emma Hamilton (née Lyon) which
Romney painted, meeting harmoniously in another creature, one
would come within reasonable distance of seeing Mrs. Mowbray, as
Harold saw her when he entered the room.
Even with the disadvantage of the exaggerated colour and the
over-emphasized eye-lashes necessary for the searching illumination
of the footlights, she was very lovely, Harold acknowledged.
But all the loveliness of Mrs. Mowbray produced but a trifling
effect compared to that produced by her charm of manner. She was
the most natural woman ever known.
The position of the natural man has been defined by an eminent
authority. But who shall define the position of the natural woman?
It was Mrs. Mowbray’s perfect simplicity, especially when talking to
men—as a matter of fact she preferred talking to men rather than to
women—that made her seem so lovely—nay, that made a man feel
that it was good for him to be in her presence. She was devoid of
the smallest trace of affectation. She seemed the embodiment of
truth. She never smiled for the sake of conventionality. But when she
did smile, just as Harold entered the room, her head turning round
so that her face was looking over her shoulder, she had all the
spiritual beauty of the loveliest picture ever painted by Greuze,
consequently the loveliest picture ever painted by the hand of man.
And yet she was so very human.
An Algy and an Eddy were already in the room—the first was a
Marquis, the second was the eldest son of a duke. Both were
handsome lads, of quiet manners, and both were in the Household
Cavalry. Mrs. Mowbray liked to be surrounded by the youngest of
men.
Harold had been acquainted with her long before she had become
an actress. He had not had an opportunity of meeting her since; but
he found that she remembered him very well.
She had heard of his father, she said, looking at him in a way that
did not in the least suggest a picture by Greuze.
When people referred to his father they did not usually assume a
look of innocence. Most of them would have had difficulty in
assuming such a look under any circumstances.
“My father is frequently heard of,” said Harold.
“And your father’s son also,” said Mrs. Mowbray. “What a freak of
Lady Innisfail’s! She lured you all across to Ireland. I heard so much.
And what came of it, after all?”
“Acute admiration for the allurements of Lady Innisfail in my case,
and a touch of acute rheumatism in my father’s case,” said Harold.
“Neither will be fatal to the sufferers,” said Mrs. Mowbray—“or to
Lady Innisfail, for that matter,” she added.
“I should say not,” remarked Algy. “We all admire Lady Innisfail.”
“Few cases of acute admiration of Lady Innisfail have proved fatal,
so far as I can hear, Lord Brackenthorpe,” said Mrs. Mowbray. “Young
mem have suffered from it and have become exemplary husbands
and parents.”
“And if they don’t live happy, that we may,” said Archie.
“That’s the end of the whole matter,” said. Harold.
“That’s the end of the orthodox fairy tale,” said Mrs. Mowbray.
“Was your visit to Ireland a fairy story, Mr. Wynne?”
Harold wondered how much this woman knew of the details of his
visit to Ireland. Before he-could think of an answer in the same
strain, Mrs. Mowbray had risen from the little gilt sofa, and had
taken a step or two toward her dressing-room. The look she tossed
to Harold, when she turned round with her fingers on the handle of
the door, was a marvellous one.
Had it been attempted by any other woman, it would have
provoked derision on the part of the average man—certainly on the
part of Harold Wynne. But, coming from Mrs. Mowbray, it conveyed
—well, all that she meant it to convey. It was not merely fascinating,
it was fascination itself.
It was such a look as this, he felt—but nearly a year had passed
before he had thought of the parallel—that Venus had cast at Paris
upon a momentous occasion. It was the glance of Venus Victrix. It
made a man think—a year or so afterwards—of Ahola and Aholibah,
of Ashtoreth, of Cleopatra, of Faustina, of Iseult, of Rosamond.
And yet the momentary expression of her features was as simple
and as natural as that worn by one of Greuze’s girls.
“She’ll not be more than ten minutes,” said
Archie. “I don’t know how she manages to dress herself in the
time.”
He did not exaggerate. Mrs. Mowbray returned in ten minutes,
with no trace of paint upon her face, wearing a robe that seemed to
surround her with fleecy clouds. The garment was not much more
than an atmosphere—it was a good deal less substantial than the
atmosphere of London in December or that of Sheffield in June.
“We shall have the pleasantest of suppers,” she said, “and the
pleasantest of chats. I understand that Mr. Wynne has solved the
Irish problem.”
“And what is the solution, Mrs. Mowbray?” said Lord
Brackenthorpe.
“The solution—ah—‘a gray eye or so’,” said Mrs. Mowbray.
The little Mercutio swagger with which she gave point to the
words, was better than anything she had done on the stage.
“And now, Mr. Wynne, you must lead the way with me to our little
supper-room,” said she, before the laugh, in which everyone joined,
at the pretty bit of comedy, had ceased.
Harold gave her his arm.
When at the point of entering the room—it was daintily furnished
with old English oak and old English silver—Mrs. Mowbray said, in
the most casual way possible, “I hope you will tell me all that may
be told about that charming White Lady of the Cave. How amusing it
must have been to watch the chagrin of Lord Fotheringay, when Mr.
Airey gave him to understand that he meant to make love to that
young person with the wonderful eyes.”
“It was intensely amusing, indeed,” said Harold, who had become
prepared for anything that Mrs. Mowbray might say.
“Yes, you must have been amused; for, of course, you knew that
Mr. Airey was not in earnest—that he had simply been told off by
Miss Craven to amuse himself with the young person, in order to
induce her to take her beautiful eyes off—off—someone else, and to
turn them admiringly upon Mr. Airey.”
“That was the most amusing part of the comedy, of course,” said
Harold.
“What fools some girls are!” laughed Mrs. Mowbray. It was well
known that she disliked the society of women.
“It’s a wise provision of nature that the fools should be the girls.”
“Oh, I have known a fool or two among men,” said Mrs. Mowbray,
with another laugh.
“Have known—did you say have known?” said Harold.
“Any girl who has lived in this world of ours for a quarter of a
century, should have seen enough to make her aware of the fact
that the best way to set about increasing the passion of, let us say,
the average man—”
“No, the average man is passionless.”
“Well, the passion of whatever man you please—for a young
woman whom he loves, or fancies he loves—it’s all the same in the
end—is to induce him to believe that several other men are also in
love with her.”
“That is one of the rudiments of a science of which you are the
leading exponent,” said Harold.
“And yet Miss Craven was foolish enough to fancy that the man of
whom she was thinking, would give himself up to think of her so
soon as he believed that Mr. Airey was in love with her rival! Ah,
here are our lentils and pulse. How good it is of you to imperil your
digestions by taking supper with me, when only a few hours can
have passed since you dined.”
“Digestion is not an immortal soul,” said Harold, “and I believe
that immortal souls have been imperilled before now, for the sake of
taking supper with the most beautiful woman in the world.”
“Have you ever heard a woman say that I am beautiful?” she
asked.
“Never,” said Harold. “That is the one sin which a woman never
pardons in another.”
“You do not know women—” with a little pitying smile. “A woman
will forgive a woman for being more beautiful than herself—for being
less virtuous than herself, but never for being better-dressed than
herself.”
“For how many of the three sins do you ask forgiveness of woman
—two or three?” said Harold, gently.
But instead of making an answer, Mrs. Mowbray said something
about the necessity of cherishing a digestion. It was disgraceful, she
said, that bread-and-butter and arithmetic should be forced upon a
school boy—that such magnificent powers of digestion as he
possessed should not be utilized ta the uttermost.
Lord Brackenthorpe said he knew a clever artist chap, who had
drawn a sketch of about a thousand people crowding over one
another, in an American hotel, in order to see a boy, who had been
overheard asking his mother what was the meaning of the word
dyspepsia.
Mrs. Mowbray wondered if the melancholy of Hamlet was due to a
weak digestion.
Harold said he thought it should rather be accepted as evidence
that there was a Schleswig-Holstein question even in Hamlet’s day.
Meantime, the pheasants and sparkling red Burgundy were
affording compensation for the absence of any brilliant talk.
Then the young men lit their cigarettes. Mrs. Mowbray had never
been known to risk her reputation (for femininity) by letting a
cigarette between her lips; but her femininity was in no way
jeopardized—rather was it accentuated—by her liking to be in the
neighbourhood of where cigarettes were being smoked—that is,
when the cigarettes were good and when the smokers were pleasant
young men with titles, or even unpleasant young men with
thousands.
After the lapse of an hour, a message came regarding Mrs.
Mowbray’s brougham. Her guests rose and she looked about for her
wrap.
While Harold Wynne was laying it on her lovely shoulders, she
kept her eyes fixed upon his. Hers were full of intelligence. When he
had carefully fastened the gold clasp just beneath the hollow of her
throat—it required very careful handling—she poised her head to the
extent of perhaps a quarter of an inch to one side, and laughed;
then she moved away from him, but turned her head so that her
face was once more over her shoulder, like the face of the Greuze
girl from whom she had learnt the trick.
He knew that she wanted him to ask her from whom she had
heard the stories regarding Castle Innisfail and its guests.
He also knew that the reason she wanted him to ask her this
question, was in order that she might have the delight of refusing to
answer him, while keeping him in the expectancy of receiving an
answer.
Such a delight would, of course, be a malicious one. But he knew
that it would be a thoroughly womanly one, and he knew that Mrs.
Mowbray was a thorough woman.
Therefore he laughed back at her and did not ask her anything—
not even to take his arm out to her brougham.
Archie Brown did, and she took his arm, still looking over her
shoulder at Harold.
It only needed that the lovely, wicked look should vanish in a
sentence.
And it did.
The full lips parted, and the poise of the head was increased by
perhaps the eighth part of an inch.
“‘A gray eye or so,’” she murmured.
Her laughter rang down the corridor.
“And the best of it all is, that no one can say a word against her
character,” said Archie.
This was the conclusion of his rhapsody in the hansom, in which
he and Harold were driving down Piccadilly—a rhapsody upon the
beauty, the genius, and the expensiveness of Mrs. Mowbray.
Harold was silent. The truth was that he was thinking about
something far apart from Mrs. Mowbray, her beauty, her doubtful
genius, and her undoubted power of spending money.
“What do you say?” said Archie. “Great Godfrey! you don’t mean
to say that you’ve heard a word breathed against her character?”
“On the contrary,” said Harold, “I’ve always heard it asserted that
Mrs. Mowbray is the best dressed woman in London.”
“Give me your hand, old chap; I knew that I could trust you to do
her justice,” cried Archie.
CHAPTER XXXIII.—ON BLESSING
OR DOOM.
E
VEN before he slept, Harold Wynne found that he had a good
many matters to think about, in addition to the exquisitely
natural poises of Mrs. Mowbray’s shapely head.
It was apparent to him that Mrs. Mowbray had somehow obtained
a circumstantial account of the appearance of Beatrice Avon at the
Irish Castle, and of the effect that had been produced, in more than
one direction, by her appearance.
But the most important information that he had derived from Mrs.
Mowbray was that which had reference to the attitude of Edmund
Airey toward Beatrice.
Undoubtedly, Mrs. Mowbray had, by some means, come to be
possessed of the truth regarding the apparent fascination which
Beatrice had for Edmund Airey. It was a trick—it was the result of a
conspiracy between Helen Craven and Edmund, in order that he,
Harold, should be prevented from even telling Beatrice that he loved
her. Helen had felt certain that Beatrice, when she fancied—poor
girl!—that she had produced so extraordinary an impression upon
the wealthy and distinguished man, would be likely to treat the poor
and undistinguished man, whose name was Harold Wynne, in such a
way as would prevent him from ever telling her that he loved her!
And Edmund had not hesitated to play the part which Helen had
assigned to him! For more than a moment did Harold feel that his
friend had behaved in a grossly dishonourable way. But he knew that
his friend, if taxed with behaving dishonourably, would be ready to
prove—if he thought it necessary—that, so far from acting
dishonourably, he had shown himself to be Harold’s best friend, by
doing his best to prevent Harold from asking a penniless girl to be
his wife. Oh, yes, Mr. Edmund Airey would have no trouble in
showing, to the satisfaction of a considerable number of people—
perhaps, even to his own satisfaction—that he was acting the part of
a truly conscientious; and, perhaps, a self-sacrificing friend, by
adopting Helen Craven’s suggestion.
Harold felt very bitter toward his friend Edmund Airey; though it
was unreasonable for him to do so; for had not he come to precisely
the same conclusion as his friend in respect of Beatrice, this
conclusion being, of course, that nothing but unhappiness could be
the result of his loving Beatrice, and of his asking Beatrice to love
him?
If Edmund Airey had succeeded in preventing him from carrying
out his designs, Harold would be saved from the necessity of having
with Beatrice that melancholy interview to which he was looking
forward; therefore it was unreasonable for him to entertain any
feeling of bitterness toward Edmund.
But for all that, he felt very bitterly toward Edmund—a fact which
shows that, in some men as well as in all women, logic is
subordinate to feeling.
It was also far from logical on his part to begin to think, only after
he had accused his friend of dishonourable conduct, of the source
whence the evidence upon which he had founded his accusation,
was derived.
How had Mrs. Mowbray come to hear how Edmund Airey had
plotted with Helen Craven, he asked himself. He began to wonder
how she could have heard about the gray eyes of Beatrice, to which
she had alluded more than once, with such excellent effect from the
standpoint of art. From whom could she have heard so much?
She certainly did not hear it from Mr. Durdan, even if she was
acquainted with him, which was doubtful; for Mr. Durdan was
discreet. Besides, Mr. Durdan was rarely eloquent on any social
subject. He was the sort of man who makes a tour on the Continent
and returns to tell you of nothing except a flea at Bellaggio.
Was it possible that some of the fishing men had been taking
notes unknown to any of their fellow guests, for the benefit of Mrs.
Mowbray?
Harold did not think so.
After some time he ceased to trouble himself with these vain
speculations. The fact—he believed it to be a fact—remained the
same: someone who had been at Castle Innisfail had given Mrs.
Mowbray a highly circumstantial account of certain occurrences in
the neighbourhood of the Castle; and if Mrs. Mowbray had received
such an account, why might not anyone else be equally favoured?
Thus it was that he strayed into new regions of speculation, where
he could not possibly find any profit. What did it matter to him if
everyone in London knew that Edmund Airey had plotted with Helen
Craven, to prevent an impecunious man from marrying a penniless
girl? All that remained for him to do was to go to the girl, and tell
her that he had made a mistake—that he would be asking her to
make too great a sacrifice, were he to hold her to her promise to
love him and him only.
It was somewhat curious that his resolution in this matter should
be strengthened by the fact of his having learned that Edmund Airey
had not been in earnest, in what was generally regarded at Castle
Innisfail as an attitude of serious, and not merely autumn, love-
making, in respect of Beatrice.
He did not feel at all annoyed to learn that, if he were to withdraw
from the side of Beatrice, his place would not be taken by that
wealthy and distinguished man, Edmund Airey. When he had at first
made up his mind to go to Beatrice and ask her to forget that he
had ever told her that he loved her, he had had an uneasy feeling
that his friend might show even a greater interest than he had done
on the evening of the tableaux at the Castle, in the future
movements of Beatrice.
At that time his resolution had not been overwhelming in its force.
But now that Mrs. Mowbray had made that strange communication—
it almost amounted to a revelation—to him, he felt almost impatient
at the delay that he knew there must be before he could see the girl
and make his confession to her.
He had two more days to think over his resolution, in addition to
his sleepless night after receiving Mrs. Mowbray’s confidences; and
the result of keeping his thoughts in the one direction was, that at
last he had almost convinced himself that he was glad that the
opportunity had arrived for him to present himself to the girl, in
order to tell her that he would no longer stand in the way of her
loving someone else.
When he found himself in her presence, however, his convictions
on this particular point were scarcely so strong as they might have
been.
She was sitting in front of the fire in the great drawing-room that
retained all the original decorations of the Brothers Adam, and she
was wearing something beautifully simple—something creamy, with
old lace. The furniture of the room also belonged to the period of
the Adams, and on the walls were a number of coloured engravings
by Bartolozzi after Cipriani and Angelica Kauffmann.
She was in his arms in a moment. She gave herself to him as
naturally and as artlessly as though she were a child; and he held
her close to him, looking down upon her face without uttering a
word—kissing her mouth conscientiously, her shell-pink cheeks
earnestly, her forehead scrupulously, and her chin playfully.
This was how he opened the interview which he had arranged to
part them for ever.
Then they both drew a long breath simultaneously, and both
laughed in unison.
Then he held her away from him for a few seconds, looking upon
her exquisite face. Again he kissed her—but this time solemnly and
with something of the father about the action.
“At last—at last,” he said.
“At last,” she murmured in reply.
“It seems to me that I have never seen you before,” said he. “You
seem to be a different person altogether. I do not remember
anything of your face, except your eyes—no, by heavens! your eyes
are different also.”
“It was dark as midnight in the depths of that seal-cave,” she
whispered.
“You mean that—ah, yes, my beloved! If I could have seen your
eyes at that moment I know I should have found them full of the
light that I now see in their depths. You remember what I said to
you on the morning after your arrival at the Castle? Your eyes meant
everything to me then—I knew it—beatitude or doom.”
“And you know now what they meant?”
He looked at her earnestly and passionately for some moments.
Then his hands dropped suddenly as though they were the hands of
a man who had died in a moment—his hands dropped, he turned
away his face.
“God knows, God knows,” he said, with what seemed like a moan.
“Yes,” she said; “God knows, and you know as well as God that in
my heart there is nothing that does not mean love for you. Does
love mean blessing or doom?”
“God knows,” said he again. “Your love should mean to me the
most blessed thing on earth.”
“And your love makes me most blessed among women,” said she.
This exchange of thought could scarcely be said to make easier
the task which he had set himself to do before nightfall.
He seemed to become aware of this, for he went to the high
mantelpiece, and stood with his hands upon it, earnestly examining
the carved marble frieze, cream-tinted with age, which was on a
level with his face.
She knew, however, that he was not examining the carving from
the standpoint of a critic; and she waited silently for whatever was
coming.
It came when he ceased his scrutiny of the classical figures in high
relief, that appeared upon the marble slab.
“Beatrice, my beloved,” said he, and her face brightened. Nothing
that commenced with the assumption that she was his beloved could
be very bad. “I have been in great trouble—I am in great trouble
still.”
She was by his side in a moment, and had taken one of his hands
in hers. She held it, looking up to his face with her eyes full of
sympathy and concern.
“My dearest,” he said, “you are all that is good and gracious. We
must part, and for ever.”
She laughed, still looking at his face. There really was something
laughable in the sequence of his words. But her laugh did not make
his task any easier.
“When I told you that I loved you, Beatrice, I told you the truth,”
said he. “If I were to tell you anything else now it would be a
falsehood. But I had no right ever to speak to you of love. I am
absolutely penniless.”
“That is no confession,” said she. “I knew all along that you were
dependent upon your father for everything. I felt for you—so did Mr.
Airey.”
“Mr. Airey?” said he. “Mr. Airey mentioned to you that I was a
beggar?”
“Oh, he didn’t say that. He only said—what did he say?—
something about the affairs of the world being very badly arranged,
otherwise you should have thousands—oh, he said he felt for you
with all his heart.”
“‘With all his appreciation of the value of an opportunity,’ he
should have said. Never mind Edmund Airey. You, yourself, can see,
Beatrice, how impossible it would be for any man with the least
sense of honour, situated as I am, to ask you to wait—to wait for
something indefinite.”
“You did not ask me to wait for anything. You did not ask me to
wait for your love—you gave it to me at once. There is nothing
indefinite in love.”
“My Beatrice, you cannot think that I would ask you for your love
without hoping to marry you?”
“Then let us be married to-morrow.”
She did not laugh, speaking the words. He could see that she
would not hesitate to marry him at any moment.
“Would to heaven that we could be, my dearest! But could there
be anything more cruel than for a penniless man, such as I am, to
ask a girl, such as you are, to marry him?”
“I cannot see where the cruelty would be. People have been very
happy together before now, though they have had very little money
between them.”
“My dear Beatrice, you were not meant to pass your life in squalid
lodgings, with none of the refinements of life around you; and I—
well, I have known what roughing it means; I would face the worst
alone; but I am not selfish enough to seek to drag you down to my
level—to ask you to face hardship for my sake.”
“But I——”
“Do not say anything, darling: anything that you may say will only
make it the harder to part. I can do it, Beatrice; I am strong enough
to say good-bye.”
“Then say it, Harold.”
She stood facing him, with her wonderful eyes looking steadily
into his. The message that they conveyed to him was such as he
could not fail to read aright. He knew that if he had said goodbye,
he would never have a chance of looking into those eyes again.
And yet he made the attempt to speak—to say the word that she
had challenged him to utter. His lips were parted for more than a
moment. He suddenly dropped her hand—he had been holding it all
the time—and turned away from her with a passionate gesture.
“I cannot say it—God help me! I cannot say good-bye,” he cried.
He had flung himself into a sofa and had buried his face in his
hands.
For a short time he had actually felt that he was desirous to part
from her. For some minutes he had been quite sincere. The force of
the words he had made use of to show Beatrice how absolutely
necessary it was that they should part, had not been felt by her;
those words had, however, affected him. He had felt—for the first
time, in spite of his previous self-communing—that he must say
good-bye to her, but he found that he was too weak to say it.
He felt a hand upon his shoulder. He could feel her gracious
presence near to him, before her voice came.
“Harold,” she said, “if you had said it, I should never have had an
hour’s happiness in my life. I would never have seen you again. I felt
that all the happiness of my life was dependent upon your refraining
from speaking those words. Cannot you see, my love, that the
matter has passed out of our hands—that it is out of our power to
part now? Harold, cannot you see that, let it be for good or evil—for
heaven or doom—we must be together? Whatever is before us, we
are not two but one—our lives are joined beyond the power of
separation. I am yours; you are mine.”
He sprang to his feet. He saw that tears were in her eyes. “Let it
be so,” he cried. “In God’s name let it be so. Whatever may happen,
no suggestion of parting shall come from me. We stand together,
and for ever, Beatrice.”
“For ever and ever,” she said.
That was how their interview came to a close.
Did he know when he had set out for her home that this would be
the close of their interview—this clasping of the hands—this meeting
of the lips?
Perhaps he did not. But one thing is certain: if it had not had this
ending, he would have been greatly mortified.
His vanity would have received a great blow.
CHAPTER XXXIV.—ON THE
MESSAGE OF THE LILY.
W
ALKING Westward to his rooms, he enjoyed once again the
same feeling of exultation, which had been his on the
evening of the return from the seal-hunt. He felt that she
was wholly his.
He had done all that was in his power to show her how very much
better it would be for her to part from him and never to see him
again—how much better it would be for her to marry the wealthy
and distinguished man who had, out of the goodness of his heart,
expressed to her a deep sympathy for his, Harold’s, unfortunate
condition of dependence upon a wicked father. But he had not been
able to convince her that it would be to her advantage to adopt this
course.
Yet, instead of feeling deeply humiliated by reason of the failure of
his arguments, he felt exultant.
“She is mine—she is mine!” he cried, when he found himself alone
in his room in St. James’s. “There is none like her, and she is mine!”
He reflected for a long time upon her beauty. He thought of Mrs.
Mowbray, and he smiled, knowing that Beatrice was far lovelier,
though her loveliness was not of the same impressive type. One did
not seem to breathe near Beatrice that atmosphere heavy with the
scent of roses, which Mrs. Mowbray carried with her for the
intoxication of the nations. Still, the beauty of Beatrice was not a
tame thing. It had stirred him, and it had stirred other men.
Yes, it had stirred Edmund Airey—he felt certain of it, although he
did not doubt the truth of Mrs. Mowbray’s communication on this
subject.
Even though Edmund Airey had been in a plot with Helen, still
Harold felt that he had been stirred by the beauty of Beatrice.
He thought over this point for some time, and the conclusion that
he came to was, that he could easily find out if Edmund meant to
play no more important a rôle than that of partner in Helen Craven’s
plot. It was perfectly clear, that if Edmund had merely acted as he
had done at the suggestion of Helen, he would cease to take any
further interest in Beatrice, unless it was his intention to devote his
life to carrying out the plot.
In the course of some weeks, he would learn all that could be
known on this point; but meantime his condition was a peculiar one.
He would have felt mortified had he been certain that Edmund
Airey had not really been stirred by the charm of Beatrice; but he
would have been somewhat uneasy had he felt certain that Edmund
was deeply in love with her. He trusted her implicitly—he felt certain
of himself in this respect. Had he not a right to trust her, after the
way in which she had spoken to him—the way in which she had
given herself up to him? But then he felt that he had made use of
such definite arguments to her, in pointing out the advisability of
their parting, as caused it to be quite possible that she might begin
to perceive—after a year or two of waiting—that there was some
value in those arguments of his, after all.
By the time that he had dined with some people, who had sent
him a card on his return to London, and had subjected himself to the
mortifying influence of some unfamiliar entrées, and a conversation
with a woman who was the survivor of the wreck of spiritualism in
London, he was no longer in the exultant mood of the afternoon.
“A Fool’s Paradise—a Fool’s Paradise!” he murmured, as he sat in
an easy-chair, and gazed into his flickering fire.
It had been very glorious to think that he was beloved by that
exquisite girl—to think of the kisses of her mouth; but whither was
the love leading him?
His father’s words could not be forgotten—those words which he
had spoken from beneath the eiderdown of his bed at Castle
Innisfail; and Harold knew that, should he marry Beatrice, his father
would certainly carry out his threat of cutting off his allowance.
Thus it was that he sat in his chair feeling that, even though
Beatrice had refused to be separated from him, still they were as
completely parted by circumstances as if she had immediately
acknowledged the force of his arguments, and had accepted, his
invitation to say good-bye for ever.
Thus it was that he cried, “A Fool’s Paradise—a Fool’s Paradise!” as
he thought over the whole matter.
What were the exact elements of the Paradise in which his
exclamation suggested that he was living, he might have had some
difficulty in defining.
But then the site of the original Paradise is still a matter of
speculation.
The next day he went to take lunch with her and her father—he
had promised to do so before he had left her, when they had had
their interview.
It so happened, however, that he only partook of lunch with
Beatrice; for Mr. Avon had, he learned, been compelled to go to
Dublin for some days, to satisfy himself regarding a document which
was in a library in that city.
Harold did not grumble at the prospect of a long afternoon by her
side; only he could not help feeling that the ménage of the Avon
family was one of the most remarkable that he had ever known. The
historical investigations of Mr. Avon did not seem to induce him to
take a conventional view of his obligations, as the father of an
extremely handsome girl—assuming that he was aware of the fact of
her beauty—or a pessimistic view of modern society. He seemed to
allow Beatrice to be in every way her own mistress—to receive
whatever visitors she pleased; and to lay no narrow-minded
prohibition upon such an incident as lunching tête-à-tête with a
young man, or perhaps—but Harold had no knowledge of such a
case—an old man.
He wondered if the historian had ever been remonstrated with on
this subject, by such persons as had not had the advantage of
scrutinizing humanity through the medium of state papers.
Harold thought that, on the whole, he had no reason to take
exception to the liberality of Mr. Avon’s system. He reflected that it
was to this system he was indebted for what promised to be an
extremely agreeable afternoon.
What he did not reflect upon, however, was, that he was indebted
to Mr. Avon’s peculiarities—some people would undoubtedly call the
system a peculiar one—for a charmingly irresponsible relationship
toward the historian’s daughter. He did not reflect upon the fact, that
if the girl had had the Average Father, or the Vigilant Mother, to say
nothing of the Athletic Brother, he would not have been able,
without some explanation, to visit her, and, on the strength of
promising to love her, to kiss her, as he had now repeatedly done, on
the mouth—or even on the forehead, which is somewhat less
satisfying. Everyone knows that the Vigilant Mother would, by the
application of a maternal thumb-screw which she always carries
attached to her bunch of keys, have extorted from Beatrice a full
confession as to the incidents of the seal-hunt—all except the
hunting of the seals—and that this confession would have led to a
visit to the study of the Average Father, in one corner of which
reposes the rack, in working order, for the reception of the suitor.
Everyone knows so much, and also that the alternative of the
paternal rack, is the fist of the Athletic Brother.
But Mr. Harold Wynne did not seem to reflect upon these points,
when he heard the lightly uttered excuses of Beatrice for her father’s
absence, as they seated themselves at the table in the large dining-
room.
His practised eye made him aware of the fact that Beatrice
understood what he considered to be the essentials of a recherché
lunch: a lunch appeals to the eye; but wine appeals to other senses
than the sense of seeing; and the result of his judgment was to
convince him that, if Mr. Avon was as careless in the affairs of the
cellar as he was in the affairs of the drawing-room, he was to be
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