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vii
Preface

Connections Forging a link between an essential concept


and everyday life sometimes profits by a more intensive look.
Sprinkled throughout the text are 32 full-page connections written
by the author that create these stronger links. There are four sorts:
Biology and Staying Healthy discusses health issues that impact
each student, such as Working Out, The Paleo Diet (you can see
this one on the right), and Why Don’t Men Get Breast Cancer?;
Today’s Biology examines advances in biology that importantly
affect society, such as the recent discovery that a father’s age criti-
cally affects the risk of mutations in his children—older fathers put
their babies at risk; A Closer Look examines interesting points of
biology in more detail, such as a two-page look at the extinct great
reptiles that fascinated so many of us as children in Dinosaurs, and
an examination of how beer is brewed, of more interest to college
students, in Microbial Bartenders; Author’s Corner takes a more
personal view (the author’s) of how science relates to our everyday
lives, such as Where Are All My Socks Going? These connections
are not intended to provide even more stuff that a student must learn
but rather to build bridges between the concepts treated in the text
and the world of more immediate interest to the student.

Following a Learning Path Students learn best when they are


given a clear idea of what they are supposed to learn. With this in mind,
each chapter is broken into conceptual blocks, each block introduced with
a specific learning objective that pinpoints the concept or process that is
the focus of the block. Listed at the beginning of the chapter to provide an
overview of what is coming, the individual learning objectives are repeated
again within the chapter when that content arises, placing the bull’s-eye
(the learning objective) right on its target (the text to be understood and
learned). At the end of each learning block, the student encounters a
­Putting the Concept to Work question. This question requires the s­ tudent
to draw a conclusion from what he or she has learned—to put the learning
objective to work and in that way reinforce its retention. As the student
moves through the chapter, these blocks of content form a learning path,
a conceptual journey taken one step at a time, each step reinforced as
it is taken.

Inquiry & Analysis One of the most useful things a student


can take away from his or her biology class is the ability to judge sci-
entific claims that they encounter as citizens, long after college is over.
When a new advance is announced on television or trumpeted by a
friend on Facebook, how relevant are the data being advanced to support
this claim? Is the claim justified by these data? Because it is touted on
the media doesn’t make a claim reliable. As an educated citizen who
has completed a biology course, a student should be able to make an
informed ­judgment. As a way of teaching that important after-the-class-
is-over skill, the Inquiry & Analysis features introduced in previous
editions have been expanded. All chapters now end with a full-page
presentation of an actual scientific investigation that requires the student
to analyze the data and reach conclusions. These exercises encourage a
student to think like a scientist does, evaluating alternatives carefully,
assessing the evidence. This is learning that lasts.
About the Author
Dr. George B. Johnson is a researcher, educator,
and author. Born in 1942 in Virginia, he went to college in
New Hampshire (Dartmouth), attended graduate school in
California (Stanford), and is Professor Emeritus of Biol-
ogy at Washington University in St. Louis, where he has
taught freshman biology and genetics to undergraduates for
over 35 years. Also Professor of Genetics at Washington
University’s School of Medicine, Dr. Johnson is a student
of population genetics and evolution, authoring more than
50 scientific journal publications. His laboratory work is
renowned for pioneering the study of previously undis-
closed genetic variability. His field research has centered on
alpine butterflies and flowers, much of it carried out in the
Rocky Mountains of Colorado and Wyoming. Other ecosys-
tems he has explored in recent years include Brazilian and
Costa Rican rainforest, the Florida Everglades, the seacoast
of Maine, coral reefs off Belize, the ice fields and mountains
of Patagonia, and, delightfully, vineyards in Tuscany.
A profilic writer and educator, Dr. Johnson is the
author of seven nationally recognized college texts for
McGraw-Hill, including the hugely successful majors texts
Biology (with botanist Peter Raven) and three nonmajors
texts: Understanding Biology, The Living World, and this
text, Essentials of The Living World. He has also authored
two widely used high school biology textbooks, Holt Biol-
ogy and Biology: Visualizing Life. In the 30 years he has
been authoring biology texts, over 3 million students have
been taught from textbooks Dr. Johnson has written.
Dr. Johnson has been involved in innovative efforts
to incorporate interactive learning and Internet experi-
ences into our nation’s classrooms. He has served on a
National Research Council task force to improve high
school biology teaching and as the founding director of
The Living World, the education center at the St. Louis
Zoo, where he was responsible for developing a broad
range of innovative high-tech exhibits and an array of new
educational programs.
St. Louis students may be familiar with Dr. Johnson as the author of a weekly science column, “On Science,” appearing for many years
in the St. Louis Post Dispatch. Dedicated to educating the general public about today’s science, Dr. Johnson continues to write new columns
regularly on current issues where science plays a key role, issues such as AIDS, the environment, cloning, genetic engineering, and evolution.
The columns, focused on explaining “how” and “why,” are intended to give readers the tools to think about these issues as citizens and voters.
You may follow his columns on Twitter @BiologyWriter.
Dr. Johnson is best known for the clarity of his writing. Most students are very interested in science, he points out, but are put off by the
terminology. When you don’t know what the words mean, its easy to slip into thinking that the matter is difficult, when actually the ideas are
simple, easy to grasp, and fun to consider. It’s the terms that get in the way, that stand as a wall between students and science. In his writing,
Dr. Johnson aims to turn those walls into windows, so that students can peer in and join the fun. Analogies are his tool. In each chapter, he
looks for simple analogies that relate the matter at hand to things we all know. As science, analogies are not exact, trading precision for clarity.
“But if I do my job right,” Johnson explains, “the key idea is not compromised by the analogy I use to explain it, but rather revealed.”
viii
New to This Edition

Chapter 1 The Science of Biology • Potentially-habitable planet orbits distant star (p. 304)
• The Biology of Aging: In 2015, the antiaging protein GDF11 is shown to reverse • New feature: Microbial Bartenders (p. 307)
aging (p. 17) • Total revision of phylogeny of the protists to reflect new DNA genome sequence
• Denisovan: genome of an unsuspected human ancestor (p. 32) analyses (p. 314)
Chapter 2 The Chemistry of Life • Total revision of phylogeny of the fungi to reflect
• How cyanide poisoned Kentucky racehorse foals (p. 35) new DNA genome sequence analysis (p. 318)
Chapter 3 Molecules of Life Chapter 17 Evolution of Plants
• Baseball superstar Alex Rodriguez and anabolic steroids (p. 51) • Account of primary succession on a new volcanic island (p. 323)
• New feature: Prions and Mad Cow Disease (p. 58) Chapter 18 Evolution of Animals
• Chemical structures of sex hormones contrasted (p. 64) • Discovery of two previously unknown human species (p. 341)
Chapter 4 Cells • Our genome contains genes of three human species (p. 372)
• Internal structure of single-celled protist Dileptus (p. 67) Chapter 19 Populations and Communities
• Role of gradients in directional cell-cell interactions (p. 90) • Resource Partitioning Among Darwin’s Finches (p. 391)
Chapter 5 Energy and Life • How Africanized bees outcompete Texas bees (p. 400)
• Vegetarian and vegan diets (p. 93) Chapter 20 Ecosystems
• Analysis of the diet of a cow (p. 102) • Is the ivory-billed woodpecker really extinct? (p. 403)
Chapter 6 Photosynthesis: Acquiring Energy from the Sun Chapter 21 Behavior and the Environment
• Attacking global warming by Fe fertilization of oceans (p. 105) • Human sex pheromones (p. 429)
Chapter 7 How Cells Harvest Energy from Food • Menstrual synchrony and EST in sweat (p. 448)
• New feature: Evaluation of the Paleo Diet (p. 131) Chapter 22 Human Influences on the Living World
• The human body’s “set point” for controlling weight (p. 132) • Polar bears lose their home to global warming (p. 451)
Chapter 8 Mitosis Chapter 23 The Animal Body and How It Moves
• How tanning causes skin cancer (p. 135) • Why osteoporosis is a woman’s problem (p. 473)
• Curing cancer by disabling immune system inhibitors (p. 145) • Shifting the parathyroid hormone–calcitonin balance (p. 486)
• Role of UV in blocking p53 and so kick-starting cancer (p. 146) Chapter 24 Circulation
Chapter 9 Meiosis • Vampires and vampire bats (p. 489)
• Solving the evolutionary puzzle of the origin of sex (p. 149) Chapter 25 Respiration
• Why parthenogenesis is common in nature (p. 158) • How whales live and breathe in the sea (p. 503)
Chapter 10 Foundations of Genetics • Lung cancer statistics updated (p. 511)
• Public policy and childhood intelligence (p. 186) • Myoglobin and hemoglobin contrasted (p. 512)
Chapter 11 DNA: The Genetic Material Chapter 26 The Path of Food Through the Animal Body
• Using DNA in crime scene investigations (p. 189) • Causes of today’s type II diabetes epidemic (p. 515)
• Revising the role of Wilkins in DNA studies (p. 192, 193) • Fecal transplants (p. 528)
• New feature: Father’s Age Affects Risk of Mutation (p. 199) Chapter 27 Maintaining the Internal Environment
• DNA forensics from a single hair (p. 202) • Why kangaroo rats never drink (p. 531)
Chapter 12 How Genes Work • How camels and migrating birds conserve water (p. 540)
• Editing your genes with CRISPR (p. 205) Chapter 28 How the Animal Body Defends Itself
• Role of RNA scaffolding in positioning the catalytic proteins of a • Promising new approach to AIDS vaccine (p. 543)
ribosome (p. 209) • AIDS statistics updated (p. 557)
• New feature: Can CRISPR Eliminate Malaria? (p. 219) • Combining broadly neutralizing HIV antibodies (p. 558)
• Using CRISPR to edit sperm and eggs (p. 220) Chapter 29 The Nervous System
Chapter 13 The New Biology • E-cigarettes and nicotine addiction (p. 561)
• Frankenfoods (p. 223) Chapter 30 Chemical Signaling Within the Animal Body
• Are insect pests developing resistance to GM crops? (p. 237) • The hormonal link between obesity and diabetes (p. 583)
• Epigenetics (p. 239) • Endocrine disruptors, BPA, and breast cancer (p. 593)
Chapter 14 Evolution and Natural Selection Chapter 31 Reproduction and Development
• Your dog’s inner wolf (p. 249) • Gardasil: Should boys get a vaccine for cervical cancer? (p. 597)
• Roles of genomic and mitochondrial DNA in evolution (p. 282) • Genital warts (p. 614)
Chapter 15 Exploring Biological Diversity Chapter 32 Plant Form and Function
• How the platypus sees with its eyes shut (p. 285) • Maple sap as squirrel candy (p. 617)
• Echidnas (p. 298) Chapter 33 Plant Reproduction and Growth
Chapter 16 Evolution of Microbial Life • Analysis of how redwood trees are able to grow so tall (p. 635)
• Ebola epidemic of 2014 in Central Africa (p. 301) • There are two different kinds of great redwood trees (p. 648)
ix
® ® ® ® ®

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Acknowledgments
Every author knows that he or she labors on the
shoulders of many others. The text you see is the
result of hard work by an army of behind-the-
scenes editors, spelling and grammar checkers,
photo researchers, and artists who perform their
magic on my manuscript and an even larger army of
production managers and staff who then transform
this manuscript into a bound book. I thank them all.
The key players: Anne Winch was my devel-
opmental editor, as she has been on many of my
past books; she continues to be a delight to work
with, a strong but cheerful guide, experienced,
patient—and quietly inflexible when I am trying to
do something stupid.
Chris Loewenberg was my editor—what they
now call a “brand manager”—with whom I worked
every day. His background is in marketing, so he is
very sensitive to the audience for whom I am writ-
ing and doesn’t let me forget it. The strong approach
this edition takes to relevance in its Chapter Open-
ers reflects this. In the best of all possible worlds,
editors are supposed to guide authors; while after all these years another Texan. This edition involved the selection of a great many
authoring I don’t steer very easily, Chris has proven to be very good new photos, and Emily made this process a joy, while giving me a
at it, as the book shows. chance to create a stronger visual book.
Publisher Michael Hackett (what they now call Managing The work of my long-time, off-site developmental editors and
Director) solved the many management problems his author inad- right arms, Megan Berdelman and Liz Sievers, can be seen in every
vertently created in his excess of enthusiasm and provided valuable page of this book, years after they have ceased to actively work
advice and support. Mike has a clear vision of what tomorrow’s revising it. Every Connect and SmartBook question associated
digital textbooks will be like and never takes his eyes off of where with this text has Megan’s fingerprints on it. Liz’s intelligence and
he wants to go. In over 30 years of authoring textbooks, I don’t perseverance shape the layout of every chapter. Their creative con-
think I have ever enjoyed working with a publisher as much. tributions continue to play a major role in the quality of this book.
Marty Lange (General Manager) and Kurt Strand (Senior The marketing of this new edition was planned and supervised
Vice President) oversaw all of this with humor and consistent sup- by Marketing Manager Chris Ho, a new but more-than-eager war-
port. When I occasionally explode with irritation over paperwork rior working fist and glove with my editorial team while fighting
issues, Marty and Kurt calm me down—and then solve my prob- long hours in the trenches alongside the many able sales reps who
lem. I suspect there are few publishing companies where upper present my book to instructors. She was joined by Market Develop-
management is so hands-on involved with supporting their authors. ment Manager Jenna Paleski, whose incisive questioning revealed
This is one author who appreciates it. much new information about the needs of students and instructors
Kelly Heinrichs and Vicki Krug spearheaded our production across the country.
team, which for several editions now has made a habit of working Finally, I have authored other texts, and all of my writing
miracles with a tight schedule. Copy editors Emily Nelson (work- efforts have taught me the great value of reviewers in improving my
ing all the way from Texas) and Marilynn Taylor spent many texts. Scientific colleagues from around the country have provided
hours carefully trying to teach me, after all my years of writing, numerous suggestions on how to improve the content, and many
how to use a comma. I don’t easily learn new tricks—but it is instructors and students using previous editions have suggested
sometimes possible to teach me old ones. Thank you, Emily and ways to clarify explanations, improve presentations, and expand on
Marilynn, for your patience. important topics. I have tried to listen carefully to all of you. Every
The photo program was carried out by Lori Hancock, who as one of you has my thanks.
always has done a super job, with photos selected by Emily Tietz, George Johnson
xii
Contents

Preface iv Water: Cradle of Life 44


2.4 Unique Properties of Water 44
0 Studying Biology 2 2.5 Water Ionizes 46

Learning 4 3 Molecules of Life 50


0.1 How to Study 4 Forming Macromolecules 52
0.2 Using Your Textbook 7
3.1 Building Big Molecules 52
0.3 Using Your Textbook's Internet Resources 9
Types of Macromolecules 54
Putting What You Learn to Work 11
3.2 Proteins 54
0.4 Science Is a Way of Thinking 11
3.3 Nucleic Acids 59
0.5 How to Read a Graph 13
3.4 Carbohydrates 61
3.5 Lipids 63

Part 1 4 Cells 66
The Study of Life The World of Cells 68
4.1 Cells 68
1 The Science of Biology 16 Kinds of Cells 71
Biology and the Living World 18 4.2 Prokaryotic Cells 71
1.1 The Diversity of Life 18 4.3 Eukaryotic Cells 72
1.2 Properties of Life 19
1.3 The Organization of Life 20 Tour of a Eukaryotic Cell 74
1.4 Biological Themes 22 4.4  he Plasma Membrane 74
T
4.5 The Nucleus: The Cell’s Control Center 76
The Scientific Process 24 4.6 The Endomembrane System 78
1.5 Stages of a Scientific Investigation 24 4.7 Organelles That Harvest Energy 80
1.6 Theory and Certainty 26 4.8 The Cytoskeleton: Interior Framework of the Cell 82
Core Ideas of Biology 28 Transport Across Plasma Membranes 84
1.7 Four Theories Unify Biology as a Science 28 4.9 Diffusion and Osmosis 84
4.10 Bulk Passage into and out of Cells 86
4.11 Selective Permeability 87
Part 2
The Living Cell 5 Energy and Life 92
Cells and Energy 94
2 The Chemistry of Life 34 5.1 T he Flow of Energy in Living Things 94
5.2 The Laws of Thermodynamics 95
Some Simple Chemistry 36
2.1 Atoms 36 Cell Chemistry 96
2.2 Ions and Isotopes 38 5.3 Chemical Reactions 96
2.3 Molecules 39

xiii
xiv Contents

Enzymes 97
5.4 How Enzymes Work 97
9 Meiosis 148
5.5 How Cells Regulate Enzymes 99 Meiosis 150
9.1 Discovery of Meiosis 150
How Cells Use Energy 100 9.2 The Sexual Life Cycle 151
5.6 ATP: The Energy Currency of the Cell 100 9.3 The Stages of Meiosis 152

6 Photosynthesis: Acquiring Energy


Comparing Meiosis and Mitosis 156
9.4 How Meiosis Differs from Mitosis 156
from the Sun 104
Photosynthesis 106 10 Foundations of Genetics 160
6.1 A n Overview of Photosynthesis 106 Mendel 162
6.2 How Plants Capture Energy from Sunlight 110
10.1  endel and the Garden Pea 162
M
6.3 How Photosystems Convert Light to Chemical
10.2 W hat Mendel Observed 164
Energy 112
10.3 Mendel Proposes a Theory 166
6.4 Building New Molecules 114
10.4 Mendel’s Laws 169
Photorespiration 115
From Genotype to Phenotype 170
6.5 P
 hotorespiration: Putting the Brakes on
10.5 H ow Genes Influence Traits 170
Photosynthesis 115
10.6 W hy Some Traits Don’t Show Mendelian

7 How Cells Harvest Energy from


Inheritance 172

Chromosomes and Heredity 175


Food 118
10.7 C
 hromosomes Are the Vehicles of Mendelian
An Overview of Cellular Respiration 120 Inheritance 175
7.1 Where Is the Energy in Food? 120 10.8 Human Chromosomes 177
Respiration without Oxygen: Glycolysis 122 Human Hereditary Disorders 179
7.2 Using Coupled Reactions to Make ATP 122 10.9 Studying Pedigrees 179
Respiration with Oxygen: The Krebs Cycle 123 10.10 The Role of Mutations
in Human Heredity 181
7.3 Harvesting Electrons from Chemical Bonds 123
10.11 Genetic Counseling and Therapy 185
7.4 Using the Electrons to Make ATP 126

Harvesting Electrons without Oxygen: 11 DNA: The Genetic Material 188


Fermentation 129 Genes Are Made of DNA 190
7.5 Cells Can Metabolize Food without Oxygen 129
11.1 The Griffith Experiment 190
Other Sources of Energy 130 11.2 The Hereditary Material 191
7.6 Glucose Is Not the Only Food Molecule 130 11.3 Discovering the Structure of DNA 192

DNA Replication 194


11.4 How DNA Copies Itself 194
Part 3 Altering the Genetic Message 198
The Continuity of Life 11.5 Mutation 198

8 Mitosis 134 12 How Genes Work 204


Cell Division 136 From Gene to Protein 206
8.1  rokaryotes Have a Simple Cell Cycle 136
P 12.1 The Central Dogma 206
8.2 Eukaryotic Cell Cycle 137 12.2 Transcription 207
8.3 Chromosomes 138 12.3 Translation 208
8.4 Cell Division 140 12.4 Gene Expression 211

Cancer and the Cell Cycle 143 Regulating Gene Expression 214
8.5 What Is Cancer? 143 12.5 T ranscriptional Control in Prokaryotes 214
12.6 Transcriptional Control in Eukaryotes 216
12.7 RNA-Level Control 218
xv
Contents

15.3 Higher Categories 288


13 The New Biology 222 15.4 What Is a Species? 289
Sequencing Entire Genomes 224
Inferring Phylogeny 290
13.1 Genomics 224
15.5 How to Build a Family Tree 290
13.2 The Human Genome 226

Genetic Engineering 228 Kingdoms and Domains 294


15.6 The Kingdoms of Life 294
13.3 A Scientific Revolution 228
15.7 Domain: A Higher Level of Classification 296
13.4 Genetic Engineering and Medicine 231
13.5 Genetic Engineering and Agriculture 235

The Revolution in Cell Technology 238


16 Evolution of Microbial Life 300
Origin of Life 302
13.6 Reproductive Cloning 238
16.1 How Cells Arose 302
13.7 Stem Cell Therapy 240
13.8 Therapeutic Cloning 242 Prokaryotes 305
13.9 Gene Therapy 244 16.2 The Simplest Organisms 305

Viruses 308
16.3 Structure of Viruses 308
Part 4 16.4 How Viruses Infect Organisms 310
The Evolution and Diversity of Life
Protists 312
14 Evolution and Natural 16.5 General Biology of Protists
16.6 Kinds of Protists 314
312

Selection 248
Fungi 316
Evolution 250
16.7 A Fungus Is Not a Plant 316
14.1 D arwin’s Voyage on HMS Beagle 250
16.8 Kinds of Fungi 318
14.2 Darwin’s Evidence 252
14.3 The Theory of Natural Selection 253

Darwin’s Finches: Evolution in Action 255


17 Evolution of Plants 322
Plants 324
14.4 T he Beaks of Darwin’s Finches 255
17.1 Adapting to Terrestrial Living 324
14.5 How Natural Selection Produces Diversity 257
17.2 Plant Evolution 326
The Theory of Evolution 258
Seedless Plants 328
14.6 The Evidence for Evolution 258
17.3 Nonvascular Plants 328
14.7 Evolution’s Critics 262
17.4 The Evolution of Vascular Tissue 329
How Populations Evolve 266 17.5 Seedless Vascular Plants 330
14.8 Genetic Change in Populations 266 The Advent of Seeds 332
14.9 Agents of Evolution 268
17.6 Evolution of Seed Plants 332
Adaptation Within Populations 272 17.7 Gymnosperms 334
14.10 S ickle-Cell Disease 272 The Evolution of Flowers 336
14.11 P  eppered Moths and Industrial Melanism 274
17.8 Rise of the Angiosperms 336
14.12 Selection on Color in Guppies 276

How Species Form 279 18 Evolution of Animals 340


14.13 The Biological Species Concept 279 Introduction to the Animals 342
14.14 Isolating Mechanisms 280
18.1 G eneral Features of Animals 342
18.2 Six Key Transitions in Body Plan 344
15 Exploring Biological Diversity 284 18.3 The Animal Family Tree 346
The Classification of Organisms 286
Evolution of the Animal Phyla 348
15.1 The Invention of the Linnaean System 286
18.4 T he Simplest Animals 348
15.2 Species Names 287
18.5 Advent of Bilateral Symmetry 350
xvi Contents

18.6 C hanges in the Body Cavity 352 20.7 Latitude and Elevation 415
18.7 Redesigning the Embryo 357 20.8 Patterns of Circulation in the Ocean 416

The Parade of Vertebrates 360 Major Kinds of Ecosystems 418


18.8  verview of Vertebrate Evolution 360
O 20.9 Ocean Ecosystems 418
18.9 Fishes Dominate the Sea 362 20.10 Freshwater Ecosystems 420
18.10 Amphibians and Reptiles Invade the Land 364 20.11 Land Ecosystems 422
18.11 Birds Master the Air 368
18.12
18.13
Mammals Adapt to Colder Times 369
Human Evolution 370
21 Behavior and the Environment 428
Some Behavior Is Genetically Determined 430
21.1 A pproaches to the Study of Behavior 430
21.2 Instinctive Behavioral Patterns 431
Part 5 21.3 Genetic Effects on Behavior 432
The Living Environment Behavior Can Also Be Influenced by
Learning 433
19 Populations and 21.4 How Animals Learn 433
Communities 374 21.5 Instinct and Learning Interact to Determine
Ecology 376 Behavior 434
21.6 Animal Cognition 435
19.1 What Is Ecology? 376
Evolutionary Forces Shape Behavior 436
Populations 378
21.7 Behavioral Ecology 436
19.2 Population Range 378
21.8 A Cost-Benefit Analysis of Behavior 437
19.3 Population Distribution 379
21.9 Migratory Behavior 438
19.4 Population Growth 381
21.10 Reproductive Behaviors 440
19.5 The Influence of Population Density 384
19.6 Life History Adaptations 385 Social Behavior 442
19.7 Population Demography 386 21.11  ommunication Within Social Groups 442
C
How Competition Shapes Communities 387 21.12 Altruism and Group Living 444
21.13 Animal Societies 446
19.8 Communities 387
21.14 Human Social Behavior 447
19.9 The Niche and Competition 388

Species Interact in Many Ways 392 22 Human Influences


19.10  oevolution and Symbiosis 392
C on the Living World 450
19.11 Predation 394
19.12 Plant and Animal Defenses 396 Global Change 452
19.13 Mimicry 397 22.1 Pollution 452
22.2 Acid Precipitation 453
Community Stability 399 22.3 Global Warming 454
19.14 Ecological Succession 399 22.4 Loss of Biodiversity 455
22.5 The Ozone Hole 457
20 Ecosystems 402 Saving Our Environment 458
The Energy in Ecosystems 404 22.6 Reducing Pollution 458
20.1 Energy Flows Through Ecosystems 404 22.7 Preserving Nonreplaceable Resources 459
20.2 Ecological Pyramids 408 22.8 Curbing Population Growth 461
Materials Cycle Within Ecosystems 409 Solving Environmental Problems 464
20.3 The Water Cycle 409 22.9 P reserving Endangered Species 464
20.4 The Carbon Cycle 411 22.10 Finding Cleaner Sources of Energy 467
20.5 The Nitrogen and Phosphorus Cycles 412 22.11 Individuals Can Make the Difference 469
How Weather Shapes Ecosystems 414
20.6 The Sun and Atmospheric Circulation 414
xvii
Contents

Part 6 27 Maintaining the Internal


Animal Life Environment 530
Homeostasis 532
23 The Animal Body 27.1 How the Animal Body Maintains Homeostasis 532
and How It Moves 472 Osmoregulation 534
The Animal Body Plan 474 27.2 R egulating the Body’s Water Content 534
23.1 Organization of the Vertebrate Body 474 27.3 The Mammalian Kidney 536
Tissues of the Vertebrate Body 476 27.4 Eliminating Nitrogenous Wastes 539
23.2 E pithelium Is Protective Tissue 476
23.3 Connective Tissue Carries Out Various 28 How the Animal
Functions 477 Body Defends Itself 542
23.4 Muscle Tissue Lets the Body Move 479 Three Lines of Defense 544
23.5 Nerve Tissue Conducts Signals Rapidly 480
28.1 S kin: The First Line of Defense 544
The Skeletal and Muscular Systems 481 28.2 Cellular Counterattack: The Second Line of
23.6 Types of Skeletons 481 Defense 546
23.7 Muscles and How They Work 483 28.3 Specific Immunity: The Third Line of Defense 548

The Immune Response 549


24 Circulation 488 28.4 I nitiating the Immune Response 549
Circulation 490 28.5 T Cells: The Cellular Response 550
24.1 O pen and Closed Circulatory Systems 490 28.6 B Cells: The Humoral Response 551
24.2 Architecture of the Vertebrate Circulatory 28.7 Active Immunity Through Clonal Selection 553
System 492 28.8 Vaccination 554
24.3 Blood 495 28.9 Antibodies in Medical Diagnosis 555
24.4 Human Circulatory System 497
Defeat of the Immune System 556

25 Respiration 502
28.10 O veractive Immune System 556
28.11 AIDS: Immune System Collapse 557
Respiration 504
25.1  ypes of Respiratory Systems 504
T 29 The Nervous System 560
25.2 Respiration in Aquatic Vertebrates 505
Neurons and How They Work 562
25.3 The Mammalian Respiratory System 506
29.1 T he Animal Nervous System 562
25.4 How Respiration Works: Gas Exchange 508
29.2 Neurons and Nerve Impulses 563
Lung Cancer and Smoking 510 29.3 The Synapse 565
25.5 The Nature of Lung Cancer 510
The Central Nervous System 567

26 The Path of Food Through the


29.4 How the Brain Works 567
29.5 The Spinal Cord 570
Animal Body 514
The Peripheral Nervous System 571
Food Energy and Essential Nutrients 516
29.6 T
 he Voluntary and Autonomic
26.1 Food for Energy and Growth 516 Nervous Systems 571
Digestion 518 The Sensory Nervous System 573
26.2  ypes of Digestive Systems 518
T 29.7  ensing the Internal Environment 573
S
26.3 Vertebrate Digestive Systems 519 29.8 Sensing Gravity and Motion 574
26.4 The Mouth and Teeth 520 29.9 Sensing Chemicals: Taste and Smell 575
26.5 The Esophagus and Stomach 522 29.10 Sensing Sounds: Hearing 576
26.6 The Small and Large Intestines 524 29.11 Sensing Light: Vision 577
26.7 Accessory Digestive Organs 526
xviii Contents

30 Chemical Signaling
The Plant Body 622
32.3 Roots 622
Within the Animal Body 582 32.4 Stems 624
The Neuroendocrine System 584 32.5 Leaves 626
30.1 Hormones 584
Plant Transport and Nutrition 628
30.2 How Hormones Target Cells 586
32.6 Water Movement 628
The Major Endocrine Glands 588 32.7 Carbohydrate Transport 631
30.3 T he Hypothalamus and the Pituitary 588
30.4 The Pancreas 590 33 Plant Reproduction and Growth 634
30.5 The Thyroid, Parathyroid, and Adrenal Glands 591
Flowering Plant Reproduction 636

31 Reproduction and Development 596


33.1
33.2
 ngiosperm Reproduction
A
Seeds 639
636

Modes of Reproduction 598 33.3 Fruit 640


31.1 Asexual and Sexual Reproduction 598 33.4 Germination 641

The Human Reproductive System 600 Regulating Plant Growth 642


31.2 Males 600 33.5 Plant Hormones 642
31.3 Females 602 33.6 Auxin 644
31.4 Hormones Coordinate the Reproductive Cycle 604
Plant Responses to Environmental Stimuli 646
The Course of Development 606 33.7 Photoperiodism and Dormancy 646
31.5 E mbryonic Development 606 33.8 Tropisms 647
31.6 Fetal Development 610

Birth Control and Sexually Transmitted


Glossary 650
Diseases 612
Credits 664
31.7 C
 ontraception and Sexually Transmitted
Subject Index 667
Diseases 612

Application Index 685

Part 7
Plant Life

32 Plant Form and


Function 616
Structure and Function of Plant Tissues 618
32.1 Organization of a Vascular Plant 618
32.2 Plant Tissue Types 619
0
Studying
Biology

HER SUCCESS WILL depend not only on how much she


studies, but also on when.
Pulling an All-Nighter LE A R N I N G PATH ▼

A t some point in the next months you will face that scary rite, the first exam in this
course. Many students face the challenge of exams by cramming. They live and die
by the all-nighter, black coffee their closest friend during exam week and sleep a
luxury they can’t afford. Trying to cram enough in to meet any possible question, they feel they
can’t waste time sleeping.
Learning
1. How to Study
0.1.1 Explain why it is important to
If you take this approach, you won’t have much luck. Why doesn’t the hard work of cramming recopy your lecture notes promptly.
give good grades? Because of how humans learn. Researchers have demonstrated that memory of 0.1.2 Name two things you can do to
newly learned information improves only after hours of sleeping. If you wanted to do well on an exam, slow down the forgetting process.
you could not have chosen a poorer way to prepare than an all-nighter. 0.1.3 List three general means of
Learning is, in its most basic sense, a matter of forming memories. Research shows that a person rehearsal.
trying to learn something does not improve his or her knowledge until after they have had more than 0.1.4 Describe three strategies to
six hours of sleep (preferably eight). It seems the brain needs time to file new information away in the improve studying efficiency.
proper slots so they can be retrieved later. Without enough sleep to do this filing, new information
2. Using Your Textbook
does not get properly encoded into the brain’s memory circuits.
To sort out the role of sleep in learning, Harvard Medical School researchers used undergrads as
0.2.1 Describe how you can use your
guinea pigs. The undergraduates were trained to look for particular visual targets on a computer screen text to reinforce and clarify what you
and to push a button as soon as they were sure they had seen one. At first, responses were relatively learn in lecture.
sluggish—it typically took 400 milliseconds for a target to reach a student’s conscious awareness. With 0.2.2 Review the assessment tools
an hour’s training, however, many students were hitting the button correctly in 75 milliseconds. that your text provides to help you
How well had they learned? When retested from 3 to 12 hours later on the same day, there was master the material.
no further improvement past a student’s best time in the training session. If the researchers let a 3. Using Your Textbook’s
student get a little sleep, but less than six hours, then retested the next day, the student still showed Internet Resources
no improvement in performing the target identification.
For students who slept more than six hours,
0.3.1 Describe the five kinds of
the story was very different. Sleep greatly improved
interactive questions encountered
performance. Students who achieved 75 millisec- in Connect.
onds in the training session would reliably perform 0.3.2 Describe how LearnSmart and
the target identification in 62 milliseconds after a SmartBook test how well you have
good night’s sleep! After several nights of ample learned.
sleep, they often got even more proficient.
Why six or eight hours, and not four or five? Putting What You Learn to Work
The sort of sleeping you do at the beginning of a 4. Science Is a Way of Thinking
night’s sleep and the sort you do at the end are 0.4.1 Analyze how biological
different, and both, it appears, are required for scientists have come to a conclusion
efficient learning.
when confronted with problems of
The first two hours of sleeping are spent in deep sleep, what psychiatrists call slow-wave sleep.
major public importance.
During this time, certain brain chemicals become used up, which allows information that has been
gathered during the day to flow out of the memory center of the brain, the hippocampus, and into the
5. How to Read a Graph
cortex, the outer covering of the brain where long-term memories are stored. Like moving information 0.5.1 Explain why correlation of
in a computer from active memory to the hard drive, this process preserves experience for future dependent variables does not prove
reference. Without it, long-term learning cannot occur. causation.
Over the next hours, the cortex sorts through the information it has received, distributing it to 0.5.2 Discriminate between arithmetic
various locations and networks. Particular connections between nerve cells become strengthened and logarithmic scales.
as memories are preserved, a process that is thought to require the time-consuming manufacturing 0.5.3 Explain how a regression line
of new proteins. If you halt this process before it is complete, the day’s memories do not get fully
is drawn.
“transcribed,” and you don’t remember all that you would have, had you allowed the process to
0.5.4 List and discuss the four distinct
continue to completion. A few hours are just not enough time to get the job done. Four hours, the
Harvard researchers estimate, is a minimum requirement.
steps scientists use to analyze a graph.
The last two hours of a night’s uninterrupted sleep are spent in rapid-eye-movement (rem) sleep.
This is when dreams occur. The brain shuts down the connection to the hippocampus and runs through
the data it has stored over the previous hours. This process is also important to learning, as it reinforces
and strengthens the many connections between nerve cells that make up the new memory. Like a child
repeating a refrain to memorize it, the brain goes over what it has learned, until practice makes perfect.
That’s why getting by on three or four hours of sleep during exam week and crashing for 12 hours
on weekends doesn’t work. After a few days, all of the facts memorized during “all-nighters” fade away,

never given a chance to integrate properly into memory circuits. 3
4 Chapter 0 Studying Biology

Learning
Take
0.1 How to Study
Get a good night’s sleep before the exam exam
Taking Notes
Revisit notes & text where indicated
LEARNING OBJECTIVE 0.1.1 Explain why it is important to
Quiz yourself
Review revised notes recopy your lecture notes promptly.
Learning

for exams

Recopy notes Listening to lectures and reading the text are only the first steps in
Link notes to text
soon after lecture learning enough to do well in a biology course. The key to mastering
Attend Take comprehensive notes
the mountain of information and concepts you are about to encounter is
lecture to take careful notes. Studying from poor-quality notes that are sparse,
disorganized, and barely intelligible is not a productive way to approach
Read assigned
text before lecture preparing for an exam.
There are three simple ways to improve the quality of your notes:
Time
1. Take many notes. Always attempt to take the most complete notes
Figure 0.1 A learning timeline.
possible during class. If you miss class, take notes yourself from a tape
of the lecture, if at all possible. It is the process of taking notes that
promotes learning. Using someone else’s notes is but a poor substitute.
When someone else takes the notes, that person tends to do most of the
learning as well.
2. Take paraphrased notes. Develop a legible style of abbreviated
note taking. Obviously, there are some things that cannot be easily
paraphrased (referred to in a simpler way), but using abbreviations
and paraphrasing will permit more comprehensive notes. Attempting
to write complete organized sentences in note taking is frustrating and
too time-consuming—people just talk too fast!
3. Revise your notes. As soon as possible after lecture, you should deci-
pher and revise your notes. Nothing else in the learning process is more
important, because this is where most of your learning will take place.
By revising your notes, you meld the information together and put it into
a context that is understandable to you. As you revise your notes, orga-
nize the material into major blocks of information with simple “heads”
to identify each block. Add ideas from your reading of the text and note
links to material in other lectures. Clarify terms and concepts that might
be confusing with short notes and definitions. Thinking through the ideas
of the lecture in this organized way will crystallize them for you, which
is the key step in learning. Also, simply rewriting your notes to make
them legible, neat, and tidy can be a tremendous improvement that will
further enhance your ease of learning (figure 0.1).

BIOLOGY & YOU Remembering and Forgetting


Improving Memory. There is an active mar-
ket on the Internet for commercial products that claim to
LEARNING OBJECTIVE 0.1.2 Name two things you can do to
improve your memory. Many of these products involve repetitive slow down the forgetting process.
games or other gimmicks; few have any lasting impact on
memory. Psychologists have carried out considerable research
Learning is the process of placing information in your memory. Just as
on this subject and have found that the best way to improve
in your computer, there are two sorts of memory. The first, short-term
memory seems to be to increase the supply of oxygen to the
brain. How do you do this? These researchers recommend aero-
memory, is analogous to the RAM (random access memory) of a computer,
bic exercise. Walking for three hours each week significantly holding information for only a short period of time. Like in your computer,
increases brain oxygen levels, as does swimming or cycling. this memory is constantly being “written over” as new information comes
One study found that chewing gum while studying will supply the in. The second kind of memory, long-term memory, consists of information
brain with enough oxygen to improve memorizing items simply that you have stored in your memory banks for future retrieval, like storing
because of the muscle movement. files on your computer’s hard drive. In its simplest context, learning is the
process of transferring information to your hard drive.
Chapter 0 Studying Biology5

Forgetting is the loss of information stored in memory. Most of what


we forget when taking exams is the natural consequence of short-term
memories not being effectively transferred to long-term memory. Forget-
ting occurs very rapidly, dropping to below 50% retention within one hour
after learning and leveling off at about 20% retention after 24 hours.
There are many things you can do to slow down the forgetting process.
Here are two important ones:
1. Recopy your notes as soon as possible after lecture. Remember,
there is about a 50% memory loss in the first hour. You should use your
textbook as well when recopying your notes.
2. Establish a purpose for reading. When you sit down to study your
textbook, have a definite goal to learn a particular concept. Each chap-
ter begins with a preview of its key concepts—let them be your guides.
Do not try and learn the entire contents of a chapter in one session;
break it up into small pieces that are “easily digested.”

Learning
LEARNING OBJECTIVE 0.1.3 List three general means of
rehearsal.

Learning may be viewed as the efficient transfer of information from your


short-term memory to your long-term memory. Learning strategists refer
to this transfer as rehearsal. As its name implies, rehearsal always involves
some form of repetition. There are four general means of rehearsal in the
jargon of education called “critical thinking skills” (figure 0.2).
Repeating. The most obvious form of rehearsal is repetition. To learn
facts, the sequence of events in a process, or the names of a group of things,
you write them down, say them aloud, and mentally repeat them over and
over until you have “memorized” them. This often is a first step on the
road to learning. Many students mistake this as the only step. It is not, as
it involves only rote memory instead of understanding. If all you do in this Figure 0.2 Learning requires work.
course is memorize facts, you will not succeed. Learning is something you do, not something that happens to you.

Organizing. It is important to organize the information you are attempt- IMPLICATION FOR YOU If you are honest with yourself, how
ing to learn, because the process of sorting and ordering increases reten- many of the four rehearsal techniques (critical thinking skills) do you
tion. For example, if you place a sequence of events in order, such as the use when you take a science course like this one? Do you think they
stages of mitosis, you will be able to recall the entire sequence if you can are as important in nonscience classes like English or history? Why?
remember what gets the sequence started.
Linking. Biology has a natural hierarchy of information, with terms and
concepts nested within other terms and concepts. You will learn facts and
concepts more easily if you attempt to connect them with something you
already know, linking them to some information that is already stored in
your memory. Throughout this textbook, you will see arrows, like the one
in figure 0.3, indicating such links. Use them to check back over concepts
and processes you have already learned. You will be surprised how much
doing this will help you learn the new material. Throughout the text, these arrows
will direct you back to related infor-
Connecting. You will learn biology much more effectively if you relate mation presented in an earlier
what you are learning to the world around you. The many challenges of chapter.

living in today’s world are often related to the information presented in


this course, and understanding these relationships will help you learn. In Figure 0.3 Linking concepts.
each chapter of this textbook you will encounter several Apps (Application These linking arrows, found throughout the text, will help you to form
dialogs) in the outer margins (there is a “BIOLOGY & YOU” App on the connections between seemingly discrete topics covered earlier in
facing page) that allow you to briefly explore a “real-world” topic related the text.
to what you are learning. Read them. You may not be tested on these Apps,
but reading them will provide you with another “hook” to help you learn
the material on which you will be tested.
6 Chapter 0 Studying Biology

Studying to Learn
LEARNING OBJECTIVE 0.1.4 Describe three strategies to
improve studying efficiency.

If I have heard it once, I have heard it a thousand times, “Gee, Professor John-
son, I studied for 20 hours straight and I still got a D.” By now, you should be
getting the idea that just throwing time at the material does not ensure a favor-
able outcome. Many students treat studying for biology like penance: If you
do it, you will be rewarded for having done so. Not always.
The length of time spent studying and the spacing between study or
reading sessions directly affect how much you learn. If you had 10 hours
to spend studying, you would be better off if you broke it up into 10 one-
hour sessions than to spend it all in one or two sessions. There are two
good reasons for this:
First, we know from formal cognition research (as well as from our
everyday life experiences) that we remember “beginnings” and “endings”
but tend to forget “middles.” Thus, the learning process can benefit from
many “beginnings” and “endings.”
Second, unless you are unusual, after 30 minutes or an hour your abil-
ity to concentrate is diminished. Concentration is a critical component of
studying to learn. Many short, topic-focused study sessions maximize your
ability to concentrate effectively. For most of us, effective concentration
also means a comfortable, quiet environment with no outside distractions
like loud music or conversations.
It is important to realize that learning biology is not something you can do
passively. Many students think that simply possessing a lecture video or a set
of class notes will get them through. In and of themselves, videos and notes
Figure 0.4 Critical learning occurs in the classroom.
are no more important than the Nautilus machine an athlete works out on. It
Learning occurs in at least four distinct stages: doing assigned is not the machine per se, but what happens when you use it effectively, that
textbook readings before lecture; attending class; listening and
taking notes during lecture; and recopying notes shortly after lecture.
is of importance.
If you are diligent in these steps, then studying lecture notes and Common sense will have a great deal to do with your success in learn-
text assignments before exams is much more effective. Skipping any ing biology, as it does in most of life’s endeavors. Your success in this
of these stages makes it far less likely that you will learn successfully. biology course will depend on simple, obvious things (figure 0.4):
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will hear things in lecture that will be familiar to you, a recognition
that is a vital form of learning reinforcement. Later you can go back
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double-breasted overcoat he was wearing. His voice was deep and
sympathetic in spite of his rather sombre appearance.
“So kind of you to accept my casual invitation,” he murmured.
“Come along, I’ve a decrepit vehicle waiting for us outside the front
of the theatre.”
The dining-room of the Royal Severn Hotel did not succeed any
better than most provincial hotels in suggesting an atmosphere of
nocturnal gaiety. The two waiters looked as if they had been dragged
out of bed by the hair of their heads in order to attend to the wants of
the unreasonable beings who required to be fed at this unnatural
hour. Most of the tables suggested that they would welcome more
cheerfully the eggs and bacon of the morning breakfast than the
lobster mayonnaise of supper. The very flowers in attendance
appeared heavy with sleep and resentful at not being allowed a
night’s repose with the other table decorations that were piled upon
one of the sideboards like wreaths upon a coffin. Half the room was
in twilight, so that the portion of it that was lighted was so
uncomfortably bright as to seem garish. At one end two members of
the chorus were trying to make a pair of youthful hosts feel at their
ease by laughter that sounded as thin as broken glass.
“I’m sorry to inflict this atmosphere of gloom upon you,” said Mr.
Kenrick. “Let’s try to dissipate it in a bottle of champagne. I did my
best to order a special supper, but my efforts were regarded with
suspicion by the management. Your fellow performers over there
seem to be enjoying themselves. Touring with them must be rather
like travelling with an aviary of large and noisy birds.”
“Oh, but they’re such dears,” Nancy exclaimed, in arms against
any criticism of her fellow players.
Mr. Kenrick put up a monocle and looked across at the group for a
moment. Then he let it fall without comment.
“You sang better than ever to-night,” he said gravely.
Nancy felt that she simpered.
“I’m in earnest, you know. What are you going to do about it?”
“My voice?”
He nodded.
“What can I do?”
“You could have it trained.”
“But, my dear man, do you realize that I’m twenty-eight? Rather
late in the day to be cultivating operatic ambitions.”
“Not at all when the voice is as good as yours, and if you go to the
right man.”
“And where is he to be found?”
“Naples.”
Nancy laughed.
“It’s like a fairy-story where the poor heroine is set an impossible
task by the wicked stepmother. How do you think I could afford to go
to Naples?”
“That’s just what I wanted to discuss with you,” said Kenrick.
“But wait a moment,” Nancy interrupted. “I have a little girl.”
“What has that got to do with training your voice?”
“Why, this. Every penny that I can save I am saving for her. She is
in a convent now, and when she leaves school in another twelve
years I want her to have a voice and be able to afford to pay for its
training. I want her to have everything that I lacked. I would be wrong
to spend the money I have saved in building castles in Spain for
myself.”
“But, my dear woman, if in another twelve years you are an
operatic star of some magnitude you’ll be able to do much more for
your daughter than you could with what you’ll save as a provincial
actress between now and then. But forgive me; you speak of a little
girl. You have a husband then?”
“My husband is dead. He died nearly four years ago.”
Kenrick nodded slowly.
“And—forgive my bluntness—you have no other entanglements?”
She flushed.
“My marriage was never an entanglement ... and if you mean ‘am I
in love with anybody now?’ why, no, I could never love anybody
again.”
“That’s a sad remark for twenty-eight. A woman’s grande passion
usually happens when she is thirty-three.”
“Mine won’t,” said Nancy obstinately.
“I shouldn’t dare the God of Love,” Kenrick warned her.
“Remember, he’s a mischievous boy and nothing gives him greater
delight than to behave as such. Never dare a boy to climb an apple-
tree or Cupid to shoot his arrows in vain. You offered him a fine
target by that remark of yours. But don’t let’s begin an argument
about love. It’s your voice I want to talk about. Surely you must
realise that you possess a contralto of the finest quality?”
“I thought it was a fairly good natural voice,” Nancy admitted. “But I
certainly never supposed it was of the finest quality.”
“Not only have you a marvellous voice, but you can act. Very few
contraltos can act. On the operatic stage they usually sound like
governesses who have drunk a little too much at a fancy-dress ball.”
“Rather voluptuous governesses usually,” Nancy laughed.
“Yes, but with the healthy voluptuousness of women who have
been eating plenty of the best butter and drinking quarts of the
richest cream. You would be different.”
“I hate to be rude,” Nancy said. “But do you know, it always seems
to me such a waste of time to talk about impossibilities. Perhaps I’ve
no imagination. I’ll talk as long and as earnestly as you like about the
best way of travelling from one town to another, or of any of life’s
small problems, but to discuss which seaside resort in the moon
would be the jolliest place to spend one’s holidays surely isn’t worth
while.”
“But why is your appearance in opera so remote from any
prospect of being realised?”
“I’ve told you, my dear man,” said Nancy impatiently. “I have
planned my life so that my small daughter may have what I could not
have. To indulge my own ambitions at her expense would be wrong.
I can’t pretend that I’m denying myself much, because, to be honest,
until I had your letter I had never contemplated myself as an operatic
star. I knew I had an unusually good contralto voice. I knew that I
could act as well as most women and a good deal better than some.
Your letter was a pleasure, because it is always a pleasure to feel
that one has interested somebody. I am grateful to you for inviting
me out to supper and saying nice things about my possibilities. But
now let’s talk of something else, for you’ll never infect me with any
ambition to do anything that could risk my ability to do what I can for
my daughter, just by acting quietly in the provinces as I am acting at
present.”
“Listen to me, Miss O’Finn,” said Kenrick earnestly. “I am a
business man. That is my inheritance from a hard-working father. But
I have one passion, and that is not business. My passion is the
opera; my dream is to make enough money to be able to help the
opera in England. But I am rich enough to do something for the
individual artist, and I beg you to let me help you. Let me guarantee
you what you would usually earn on the provincial stage. Let me pay
for your lessons. The maestro I want to teach you is an old friend of
mine. If at the end of six months he tells me that you are not the
finest contralto of the time, why, then you can go back to your life on
tour. At the worst you will have spent six months in Italy to gratify the
whim of an eccentric business man whose dreams are all of art. At
the best you will be able to do what you like for your daughter in
another ten years, and long, long before that. We’ll not talk about it
any more to-night. Go home and sleep over my proposal. Think over
it for a week. I must be back in town to-morrow. If at the end of a
week you feel that you can risk six months in Italy to have the world
at your feet, send me a line, and I will pay into your account the
necessary funds. You can leave this absurd company when you
like.”
“Och, I would have to give a fortnight’s notice,” said Nancy quickly.
Kenrick smiled.
“Very well, give your fortnight’s notice. To-day is the eleventh. If
you settle by next Saturday that will be the fifteenth. On the first of
November you can quit the fogs and be on your way to Naples. It will
probably be fine weather. It usually is about then in the south of
Italy.”
“You seem to have made up your mind that I’m going to accept
your generosity,” Nancy said.
“There is no generosity in gratifying one’s own desires,” Kenrick
observed. “But if you have any feelings of pride on the subject, why,
you can pay me back when your position is secure.”
“But why, really, are you doing this?” Nancy asked, looking deep
into the eyes of her host.
“Really and truly because I believe you have a great voice and
may become a great singer, and because if you did I should get as
much satisfaction from your success as if I had a voice and were a
great singer myself,” he replied.
The thin laughter of the chorus-girls at the other end of the room
commented upon this grave assertion. The waiter put up a grubby
hand to hide a yawn.
When Nancy woke next morning she felt like the heroine of an
Arabian Nights tale who has been carried half across Asia by a
friendly djinn. But when she called at the theatre for her letters, the
following note was a proof that she had not been dreaming:

Royal Severn Hotel,


Bristol.
October 12.
Dear Miss O’Finn,
Do think very hard over our talk last night. You can’t lose
anything by my offer; you may gain a very great deal. In
fact, I am positive that you will. Let me know your decision
at my London address, 42 Adelphi Terrace, and I will get
into communication with Maestro Gambone, and fix up
your lessons. I suggest you live at an Italian pensione in
Naples. The more Italian you can learn to speak, the
better you will sing it. I’ll find out a good place.
Good luck to you.
Yours sincerely,
John Kenrick.

It was a fine October day of rich white clouds and rain-washed


blue deeps between. A faint haze bronzed the lower air and lent the
roofs and chimneys of the city a mirrored peace, a mirrored
loveliness. Nancy wandered down by the docks and in contemplation
of the glinting masts tried to find an answer to the riddle of her future.
Suppose her voice turned out to be less good than he had
supposed? Well, that would be his bad judgment. But had she the
right to accept money from a stranger in the event of failure? It would
be his own fault if she proved a failure. It was a serious matter to
leave a company in which she had expected to be playing until next
summer. What would Sister Catherine say? Nancy remembered
what Sister Catherine had said about Italy that night they met in the
train. Sister Catherine would never be the one to blame her. She
took Letizia’s letter out of her bag and read it through again.

St. Joseph’s School,


5 Arden Grove,
N. W.
Sunday.
My dear Mother,
I hope you are very well. I am learning Italian with Sister
Catherine. It is very nice. I know twenty-two words now
and the present indicitive of “I am.” I like it very much. We
have a new girl called Dorothy Andrews. She is very nice.
She is eight and a half years old, but she is not so big as
me. I must stop now because the bell is ringing for
Vespurs and Benedicsion.
Your loving
Letizia.

She was safe for so many years, Nancy thought. Would it be so


very wrong to embark upon this adventure?
That night, when she was singing the first of her two songs, she
tried to imagine that the piece was Aïda and that she was Amneris.
“If I get a genuine encore,” she promised herself, “I’ll write to him
and accept.”
And she did get a most unmistakable encore.
“Your songs went very well to-night, dear,” said Miss Fitzroy
grudgingly. “Had you got any friends in front?”
The next day Nancy wrote to John Kenrick and told him that she
was going to accept his kind offer, and that on Sunday, October
23rd, she should be in London.
He telegraphed back: Bravo will meet train if you let me know time.
But she did not let him know the time of her arrival at Paddington,
for she thought that there was really no reason why he should want
to meet her train. Somehow it made his interest in her seem too
personal, and Nancy was determined that the whole affair should be
carried through on the lines of the strictest business. Besides, she
would be staying at the convent, and it would be so exciting to learn
her first words of Italian from Letizia.
CHAPTER XX
SOUTHWARD
St. Joseph’s School was a pleasant early Victorian house with
white jalousies encircled by a deep verandah of florid ironwork. The
garden, even for the spacious northwest of London, was
exceptionally large, and like all London gardens seemed larger than
it really was by the contrast between its arbours and the houses
entirely surrounding them. There was a mystery about its seclusion
that no country garden can possess, and one could imagine no fitter
tenants of its leafy recesses than these placid nuns and the young
girls entrusted to their tutelage. It seemed that in all those fortunate
windows of the houses which overlooked through the branches of
the great lime-trees this serene enclosure there must be sitting poets
in contemplation of the pastoral of youth being played below. The
flash of a white dress, the echo of a laugh, the flight of a tennis-ball,
the glint of tumbling curls, all these must have held the onlookers
entranced as by the murmur and motion and form and iridescence of
a fountain; and this happy valley among the arid cliffs of London
bricks must have appeared to them less credible than the green
mirages in desert lands that tease the dusty eyelids of travellers.
“I’m glad you have a friend of your own age,” Nancy said to
Letizia, when the morning after her arrival they were walking
together along the convent avenue strewn with October’s fallen
leaves.
“Well, she’s not a very great friend,” Letizia demurred.
“But I thought you wrote and told me that she was so very nice?”
“Well, she is very nice. Only I don’t like her very much.”
“But if she’s so very nice, why don’t you like her?”
“Well, I don’t like her, because she is so nice. Whenever I say,
‘Let’s do something,’ she says, ‘Oh, yes, do let’s,’ and then I don’t
want to do it so much.”
“Darling, isn’t that being rather perverse?”
“What’s ‘perverse,’ mother? Do tell me, because I’m collecting
difficult words. I’ve got thirty-eight words now, and when I’ve got fifty
I’m going to ask Hilda Moore what they all mean, and she’s twelve
and it’ll be a disgusting humiliation for her when she doesn’t know.
And that’ll be simply glorious, because she thinks she’s going to be a
yellow-ribbon presently.”
“But don’t you want to be a yellow-ribbon?”
“Oh, I don’t think it’s really worth while. Evelyn Joy who’s much the
nicest girl in the school has never been a ribbon. She said she
couldn’t be bothered. She’s frightfully nice, and I love her one of the
best six people in the world. She can’t be bothered about anything,
and most of the girls are always in a fuss about something. Dorothy
Andrews only wants to do what I want, because she thinks she ought
to. Fancy, she told me she simply longed to be a saint. And she said
if she died young she’d pray for me more than anybody, and I said,
‘Pooh, St. Maurice is always praying for me and he wears armour
and is very good-looking, so there’s no need for you to die young.’
And then she cried and said when she was dead I’d be sorry I’d
been so cruel.”
Nancy thought that Letizia was not less precocious than she had
always been, and she wondered if she ought to say anything to
Sister Catherine about it. She decided that Sister Catherine was
probably well aware of it and, not being anxious to give her the idea
that she was criticising the wonderful education that the nuns were
giving her little daughter, she resolved to say nothing.
She did, however, discuss with Sister Catherine her own project to
go to Italy and have her voice trained; and she was much relieved
when it was approved.
“It would be wrong not to avail yourself of such an opportunity,” the
nun exclaimed. “Even if it involved breaking into your own savings, I
should still urge you to go; but there seems no likelihood of that, and
there is no reason why you shouldn’t accept this Mr. Kenrick’s offer.
I’d no idea that you had a wonderful voice, and how delightful to be
going to Italy. Do sing for us one evening at Vespers before you go.
Sister Monica would be so pleased, and we shall all enjoy it so
much. We shall feel so grand.”
“But I’m just as much astonished to hear that I’ve got this
wonderful voice as you are,” Nancy said. “Nobody ever told me I
had, until this fairy prince arrived in Bristol.”
“Ah, but I think people are always so afraid to think anybody has a
good voice until somebody else has established the fact for them,”
Sister Catherine laughed. “It was just a piece of good luck that you
should be heard by somebody who understood what good singing
is.... I’m glad you think dear little Letizia is looking so well. She is a
great treasure, and we are all very proud of her. She has so much
personality, and I’m doing my best to let her keep it without spoiling
her.”
“I’m sure you are,” Nancy said. “And och, I wish I could ever tell
you how grateful I am to you.”
“There is no need of words, dear child,” said the nun, smiling. “You
prove it to us all the time. I heard from the Reverend Mother
yesterday, and she inquired most affectionately after you.”
That afternoon Nancy went to Mr. Kenrick’s flat in Adelphi Terrace.
He was so kind that she reproached herself for having refused so
brusquely to let him meet her at Paddington.
“Well, it’s all arranged with Maestro Gambone. He’s really the
kindest old man, though he may seem a little fierce before you know
him. Should he, on hearing your voice, decide it’s not worth training,
you’ll have to forgive me for rousing your ambitions and let me see
you through any difficulties you may have about getting another
engagement in England. I have taken a room for you with some
people called Arcucci who have a pensione in the Via Virgilio which
is close to Santa Lucia. Arcucci himself was a singer; but he lost his
voice through illness, poor chap. He never earned more than a local
reputation at the San Carlo Opera House; but he is full of stories
about famous singers, and you’ll get the right atmosphere from him.
His wife is a capable and homely woman who will make you as
comfortable as Neapolitans know how, which, to tell the truth, is not
saying much.”
While her patron was speaking, Nancy was gazing out of his study
window at the Thames and letting her imagination drift down on the
fast-flowing ebb with the barges that all seemed like herself bound
for some adventure far from this great city of London. Away on the
horizon beyond Lambeth the domes of the Crystal Palace sparkled
in the clearer sunshine. Even so, on an horizon much farther south
than Sydenham flashed the elusive diamonds of success and fame.
“Tuesday is no day to set out on a journey,” said Kenrick. “So, I’ve
taken your ticket for Wednesday. You’ll leave Paris that night from
the Gare de Lyon in the Rome express, and you’ll be at Naples on
Friday afternoon.”
He went to a drawer in his desk and took out the tickets.
“Good luck,” he said, holding Nancy’s hand.
She was again the prey of an embarrassment against which she
tried hard to struggle, because it seemed to smirch the spirit in which
she wanted to set out. This constraint prevented her from thanking
him except in clumsy conventional phrases.
“Now, will you dine with me to-night?”
She wanted to refuse even this, but she lacked the courage; in the
end she passed a pleasant enough evening, listening to her host
expatiate upon the career for which he assured her again and again
she was certainly destined. He wanted her to lunch and dine with
him on the next day too; but she pleaded the urgency of shopping
and packing and her desire to see something of her daughter.
“Very well then,” he said, as he put her into a hansom outside
Verrey’s where they had dined. “I’ll be at Victoria on Wednesday
morning.”
Nancy was glad to be jingling back to St. Joseph’s, alone with her
dreams in the sharp apple-sweet air of the October night.
The next day Mrs. Pottage arrived to say good-bye and help
Nancy with her shopping. By now she had long been an institution at
St. Joseph’s, where her conversation afforded the most intense
delight to the nuns.
“Well, when you wrote you was off to Italy I was in two minds if I
wouldn’t suggest coming with you. I don’t know what it is, whether
I’m getting old or ugly or both, but I’ve not had a single proposal for
eighteen months. I suppose it means I’ve got to be thinking of
settling down and giving some of the younger ones a chance. Well,
take care of yourself in Italy, and don’t eat too much ice-cream.
Funny thing, I-talians should eat so much ice-cream and yet be so
hot. There was an opera company came to Greenwich once, and the
tenor who was an I-talian stayed with me. ‘Well,’ I said to myself,
‘what he’ll want is plenty of macaroni and ice-cream.’ He looked a bit
surprised, I’m bound to say, when I give it him for breakfast on the
Sunday morning, but I thought he was only surprised at any one
knowing his tastes so well. But, will you believe me, when I give it
him for dinner again, he used language that was far from I-talian,
very far. In fact, I never heard any one swear so fluent in English
before or since. It quite dazed me for the moment. But we got on all
right as soon as I found he liked good old roast beef. He gave me
two passes for the Friday night, and Mrs. Bugbird and me thoroughly
enjoyed ourselves. The opera was called Carmen and Mrs. B.
thought it was going to be all about them, and when she found it was
actually the name of a woman she laughed herself silly. Every time
this Carmen came on she’d whisper to me, ‘a good pull up,’ and then
she’d start off shaking like a jelly. But there, she’s very quick to see
the radiculous side of anything, Mrs. Bugbird is. Well, good-bye,
dear, and take good care of yourself. You know your old Mrs.
Pottage wishes you all the best you can wish for yourself.”
Sister Catherine had repeated her request that Nancy should sing
to them, especially as it was the feast of All Saints. So after
practising with Sister Monica, who had charge of the music, she
sang Mozart’s motet Ave Verum Corpus at Benediction amid the
glowing candles and white chrysanthemums of the little chapel.
“Mother, you don’t often sing in church, do you?” Letizia asked.
“Didn’t I sing well?” said her mother with a smile.
“Yes, I expect you sang very well, but I thought it was a little loud,
didn’t you? Sometimes it sounded like a man singing. I think you
ought to be careful and not sing quite so loud, mother.”
Luckily the nuns themselves enjoyed Nancy’s rich contralto a great
deal more than did their pupils. The warmth of femininity spoke to
their hearts of something that they had lost, or rather of something
that most of them had never won. It was easy to understand and
sympathise with the readiness of the nuns to turn away for a few
minutes from the austere ecstasies of Gothic art to worship some
dolorous “Mother” of Guido Reni. A flush had tinged their cheeks so
virginally tralucent, as if a goblet of water had been faintly suffused
by a few drops of red wine.
Kenrick was at Victoria to see Nancy off next morning. Just as the
train started, she leaned out of the window of her compartment and
exclaimed breathlessly:
“Please don’t think me ungrateful. I do appreciate tremendously
what you are doing for me. Really, I do.”
His long, sombre face lit up with a smile, and he waved his hand
as Nancy withdrew from London into the train again.
France dreamed in a serenity of ethereal blue. In the little
wedding-cake cemeteries black figures were laying wreaths of
immortelles upon the graves. Nancy remembered with a pang that it
was All Souls’ Day and reproached her cowardice for not having laid
flowers on Bram’s grave at Greenwich before she left England. The
bunch of carnations with which Kenrick had presented her became
hateful to hold, and she longed to throw it out of the window. She
would have done so, if two English old maids had not been regarding
her curiously from the other side of the compartment, the one above
her Baedeker, the other above the Church Times. Why should
elderly English women travelling abroad look like butterfly-collectors?
“Parlez vous anglaise?” said one of them to the ticket-collector,
nodding her head and beaming as if she were trying to propitiate an
orang-utan.
“Yes, I spik English, madame,” he said coldly after punching the
tickets.
The other elderly lady congratulated her companion upon the
triumphant conversation.
“He undoubtedly understood perfectly what you were saying,
Ethel.”
“Oh, yes, I think we shall get along capitally after a time. I was
always considered very good at French in my schooldays, and it’s
just beginning to come back to me.”
Her ambition had been kindled by her success with the first ticket-
inspector. With the next one who invaded the compartment she took
a line of bold and direct inquiry.
“Paris, quand?”
The inspector stared back, indignation displayed upon his
countenance.
“Comment?”
“Non, quand,” said the elderly lady.
The inspector shrugged his shoulders and slammed the carriage-
door as he retired.
“That man seemed rather stupid, I thought, Ethel.”
“Most stupid,” the ambitious Ethel emphatically agreed.
Nancy felt thankful that Letizia would be taught French properly.
Sister Catherine had already suggested to her that when she was
twelve she should be sent for three years to a convent in Belgium
with which the Sisters of the Holy Infancy had an arrangement of
exchanging pupils. Nancy had been a little alarmed at first by the
prospect of sending Letizia abroad all that time; but after these two
absurd Englishwomen she felt no trouble was too great and no place
too far and no separation too long that would insure Letizia against
talking French like them in public.
But presently Nancy was too much occupied with her own
problems—transferring herself and her luggage from one station in
Paris to another, finding out how the wagon-lit toilet arrangements
worked, how to reply to the Italian examination of baggage in the Mt.
Cenis tunnel, and how to achieve the change at Rome into the
Naples train—either to criticise anybody else or even to dream and
speculate about her own operatic future.
Then Vesuvius loomed above the russet orchards and dishevelled
vines on the left of the railway. Nancy suddenly remembered that
when she and Bram were first married he had one day said how
much he should like to visit Naples with her. He had told her that he
had seen a picture of it when he was a boy and of what a thrill it had
given him. Now here it actually was, and he was not by her side to
behold it. Here Naples had been all these years, and he had never
seen it.
Time heals many wounds; but in some he makes a deeper gash
every year with his inexorable scythe.
CHAPTER XXI
CLASSIC GRIEF
Nancy was lost at first in the pensione to which Kenrick had
entrusted her. The bareness of it seemed to reflect the bareness of
her own mind amid the unmeaning sounds of a strange tongue.
During the first week she felt that she should never, stayed she in
Naples for years, acquire a single word of Italian, and the week after
she was convinced that she should never be able to say anything
more than the Italian for “yes,” “no,” “please,” “thanks,” “good night,”
“good morning,” and “bread.” For a fortnight she was so completely
stunned by the swarming rackety city that she spent all her spare
time in the aquarium, contemplating the sea-anemones. The stories
of great singers with which Signor Arcucci was to have entertained
her leisure seemed indefinitely postponed at her present rate of
progress with Italian. She should have to become proficient indeed
to follow the rapid hoarseness of that faded voice. Meanwhile, she
must wrestle with an unreasonable upside down language in which
aqua calda meant hot water and not, as one might suppose, cold.
Nancy cursed her lack of education a hundred times a day, and an
equal number of times she thanked Heaven that Letizia already
knew twenty-two Italian words and could say the present indicative
of the verb “to be.” Signora Arcucci was a plump waxen-faced
Neapolitan housewife who followed the English tradition of
supposing that a foreigner would understand her more easily if she
shouted everything she had to say about four times as loud as she
spoke ordinarily. She used to heap up Nancy’s plate with spaghetti;
and, as Nancy could not politely excuse herself from eating any
more, she simply had to work her way through the slithery pyramid
until she felt as if she must burst.
Nor did Maestro Gambone do anything to make up for the state of
discouragement into which her unfamiliar surroundings and her
inability to talk had plunged her. Nancy found his little apartment at
the top of a tall tumbledown yellow house that was clinging to the
side of the almost sheer Vomero. He was a tiny man with snow-white
hair and imperial and jet-black eyebrows and moustache. With his
glittering eyes he reminded her of a much polished five of dominos,
and when he wanted anything in a hurry (and he always did want
things in a hurry) he seemed to slide about the room with the rattle of
a shuffled domino. Although his apartment stood so high, it was in a
perpetual green twilight on account of the creepers growing in rusty
petrol tins that covered all the windows.
“You speaka italiano, madama?” he asked abruptly when Nancy
presented herself.
“No, I’m afraid I don’t.”
“Allora come canta? How you singa, madama?”
“I only sing in English at present.”
“What musica you havva?”
Nancy produced the stock-in-trade of ballads, which the maestro
fingered like noxious reptiles.
“E questo? Anna Lowrie o qualche nome indiavolato. Probiamolo.
Avanti!”
The little man sat down at the piano and was off with the
accompaniment on an instrument of the most outrageously tinny
timbre before Nancy had finished deciding that he was not so much
like a domino as a five-finger exercise.
“Eh, avanti!” he turned round and shouted angrily. “What for you
waita, madama? Di nuovo!”
In the green twilight of this little room hanging over the precipitous
cliff above the distant jangling of Naples Nancy could not feel that
Maxwellton Braes had ever existed. She made a desperate effort to
achieve an effect with the last lines.
“And for bonnie Annie Laurie
I would lay me down and dee.”
There was a silence.
Then the maestro grunted, twirled his moustache, rose from the
piano, and sat down at his desk.
“Here I writa when you come,” he said. “A rivederla e buon giorno.”
He thrust the paper into Nancy’s hand and with the same gesture
almost pushed her out of his apartment. The next thing of which she
was conscious was walking slowly down the Vomero in the honey-
coloured November sunshine and staring at the hours and days
written down upon the half-sheet of notepaper she held in her hand.
So the lessons began, and for a month she wondered why she or
anybody else should ever have suffered from a momentary delusion
that she could sing. She knew enough Italian by that time to
understand well enough that Maestro Gambone had nothing but
faults to find with her voice.
“Have I made any progress?” she found the courage to stammer
out one morning.
“Progresso? Ma che progresso? Non sa encora camminare.”
Certainly if she did not yet know how to walk she could not
progress. But when should she know how to walk? In her halting
Italian Nancy tried to extract from the maestro an answer to this.
“Quanda camminerà? Chi sa? Forse domani, forse giovedì, ma
forse mai.”
Perhaps to-morrow, perhaps on Thursday, but perhaps never!
Nancy sighed.
When she got back to the pensione she sat down and wrote to her
patron.

Pensione Arcucci,
Via Virgilio 49.
Napoli.
Dec. 8.
Dear Mr. Kenrick,
I really don’t think it’s worth your while to go on paying
for these singing lessons. Maestro Gambone told me to-
day that I might never know how to sing. I’m sure he’s
disgusted at my slowness. I’ve been having lessons for a
month now, and he has had ample time to judge whether
I’m worth his trouble. He evidently thinks I’m not. It’s a
great disappointment, and I feel a terrible fraud. But I’m
not going to reproach myself too bitterly, because, after all,
I would never have thought of becoming a singer if you
hadn’t put it into my head. So, next week I shall return to
England. I’m afraid your kindness has been....

Nancy put down her pen. Her struggles with Italian seemed to
have deprived her of the use of her own tongue. She could not
express her appreciation of what he had done for her except in a
bread-and-butter way that would be worse than writing nothing. For
all the sunlight flickering on the pink and yellow houses opposite she
felt overwhelmed by a wintry loneliness and frost. And then she
heard coming up from the street below the sound of bagpipes. She
went to the window and looked out. Two men in heavy blue cloaks
and steeple-crowned felt hats, two shaggy men cross-gartered, were
playing before the little shrine of the Blessed Virgin at the corner of
the Via Virgilio an ancient tune, a tune as ancient as the hills whence
every year they came down for the feast of the Immaculate
Conception to play their seasonable carols and grave melodies until
Christmas-tide. Nancy had been told about them, and here they
were, these—she could not remember their name, but it began with
“z”—these zamp something or other. And while she stood listening
by the window she heard far and wide the pipes of other pious
mountaineers piping their holy ancient tunes. Their bourdon sounded
above the noise of the traffic, above the harsh cries of the street-
vendors, above the chattering of people and the clattering of carts
and the cracking of whips, above the tinkling of mandolins in the
barber-shops, sounded remote and near and far and wide as the
bourdon of bees in summer.
The playing of these pipers calmed the fever of Nancy’s
dissatisfaction and seemed to give her an assurance that her failure
was not yet the sad fact she was imagining. She decided to
postpone for a little while her ultimatum to Kenrick and, tearing up
the unfinished letter, threw the pieces on the open brazier, over
which for so many hours of the wintry days Signor Arcucci used to
huddle, slowly stirring the charcoal embers with an iron fork and
musing upon the days when he sang this or that famous part. He
was out of the room for a moment, but presently he and his Signora,
as he called her, came in much excited to say that the zampognieri
were going to play for them. The pipers in the gimcrack room looked
like two great boulders from their own mountains, and the droning
throbbed almost unbearably in the constricted space. When
everybody in turn had given them a lira or two, they acknowledged
the offerings by presenting Nancy as the guest and stranger with a
large wooden spoon. She was taken aback for the moment by what
would have been in England the implication of such a gift. Even
when she had realised that it was intended as a compliment the
omen remained. She could not help wondering if this wooden spoon
might not prove to be the only gift she should ever take home from
Italy. Nevertheless, the zampognieri with their grave carols healed
her fear of discouragement, and during the next fortnight Maestro
Gambone on more than one occasion actually praised her singing
and found that at last she was beginning to place her voice
somewhat more approximately where it ought to be placed. It was as
if the fierce little black and white man had been softened by the spirit
of Christmas, of which those blue-cloaked pipers were at once the
heralds and the ambassadors with their bourdon rising and falling
upon the mandarin-scented air. Absence from home at this season
did not fill Nancy with sentimental regrets. Since Bram died
Christmas had not been a happy time for her, so intimately was its
festivity associated with that dreadful night at Greenwich four years
ago. She welcomed and enjoyed the different atmosphere of Natale,
and after so many grimy northern winters these days of turquoise,
these dusks of pearl and rose, these swift and scintillating nights.
On the anniversary of Bram’s death she drove out to Posilipo and
sat on a rock by the shore, gazing out across the milky cerulean
waters of the bay. For all the beauty of this classic view she was only
aware of it as one is aware of a landscape by Poussin or Claude,

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