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RUSSELL FOURTH CANADIAN EDITION
HERTZ
MCMILLAN
BIOLOGY
MAXWELL
HAFFIE
MILSOM
NICKLE
ELLIS
BIOLOGY
EXPLORING THE DIVERSITY OF LIFE
FOURTH CANADIAN EDITION
BIOLOGY
The textbook is very engaging and interesting, which makes it a lot easier for a student to go over
the chapters and learn the material. In addition, by removing unnecessary boxes, you have made
the chapters less dense and more “to the point.” I also like the additions made, including the Why
It Matters and Concept Maps, as they help ease the students into and out of the chapter.
— Student Advisory Board Member, E. Oliya, University of British Columbia
FOURTH
CANADIAN
EDITION
nelson.com
RUSSELL, HERTZ, MCMILLAN
FENTON, MAXWELL, HAFFIE, MILSOM, NICKLE, ELLIS
Paul E. Hertz was born and raised in New York City. Hughes Medical Institute since 1992. The Pipeline Project
He received a B.S. in Biology from Stanford University includes the Intercollegiate Partnership, a program for
in 1972, an A.M. in Biology from Harvard University in local community college students that facilitates their
1973, and a Ph.D. in Biology from Harvard University transfer to four-year colleges and universities. He teaches
in 1977. While completing field research for the doctor- one semester of the introductory sequence for Biology
ate, he served on the Biology Faculty of the University of majors and pre-professional students, lecture and labo-
Puerto Rico at Rio Piedras. After spending two years as ratory courses in vertebrate zoology and ecology, and
an Isaac Walton Killam Postdoctoral Fellow at Dalhousie a year-long seminar that introduces first-year students
Aaron Kinard
University, Paul accepted a teaching position at Barnard to scientific research. Paul is an animal physiological
College, where he has taught since 1979. He was named ecologist with a specific research interest in the thermal
Ann Whitney Olin Professor of Biology in 2000, and he biology of lizards. He has conducted fieldwork in the
received the Barnard Award for Teaching Excellence in West Indies since the mid-1970s, most recently focusing
2007. In addition to serving on numerous college com- on the lizards of Cuba. His work has been funded by the
mittees, Paul chaired Barnard’s Biology Department for NSF, and he has published his research in The American
eight years and served as Acting Provost and Dean of the Naturalist, Ecology, Nature, Oecologia, and Proceedings of
Faculty from 2011 to 2012. He is the founding Program the Royal Society. In 2010, he and his colleagues at three
Director of the Hughes Science Pipeline Project at other universities received funding from NSF for a proj-
Barnard, an undergraduate curriculum and research pro- ect designed to detect the effects of global climate warm-
gram that has been funded continuously by the Howard ing on the biology of Anolis lizards in Puerto Rico.
Beverly McMillan has been a science writer for numerous trade books on scientific subjects and has
more than 25 years. She holds undergraduate and gradu- worked extensively in educational and commercial pub-
ate degrees from the University of California, Berkeley, lishing, including eight years in editorial management
and is coauthor of a college text in human biology, now positions in the college divisions of Random House and
in its 11th edition. She has also written or coauthored McGraw-Hill.
Courtesy of Beverly McMillan
vi NEL
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Brief Contents
VOLUME 1: BIOLOGY OF THE CELL 1 25 Fungi 587
1 Light and Life 5 26 Plants 611
27 Animals 643
UNIT ONE SYSTEMS AND PROCESSES: THE 28 Conservation of Biodiversity 719
CELL 25
2 The Cell: An Overview 25 UNIT SIX ECOLOGY AND BEHAVIOUR 747
3 Energy and Enzymes 53 29 Population Ecology 747
4 Cell Membranes and Signalling 77 30 Species Interactions and Community Ecology 777
5 Cellular Respiration 101 31 Ecosystems 813
6 Photosynthesis 125 32 Animal Behaviour 845
NEL vii
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Contents
About the Canadian Authors iv 3.5 The Role of Enzymes in Biological Reactions 64
About the U.S. Authors vi 3.6 Factors That Affect Enzyme Activity 67
SUMMARY ILLUSTRATION FOR CHAPTER 3 72
Preface xv
New to This Edition xix 4 Cell Membranes and Signalling 77
Welcome to Biology: Exploring the Diversity 4.1 An Overview of the Structure of Membranes 78
of Life, 4Ce xxiv Figure 4.2 Experimental Research The Frye–Edidin Experiment
Active Learning xxvi Demonstrating That the Phospholipid Bilayer Is Fluid 79
Student and Instructor Resources xxx Figure 4.3 Research Method Freeze Fracture 80
Acknowledgements xxxii 4.2 The Lipid Fabric of a Membrane 80
4.3 Membrane Proteins 82
VOLUME 1: BIOLOGY OF THE CELL 1 4.4 Passive Membrane Transport 84
4.5 Active Membrane Transport 88
1 Light and Life 5
4.6 Exocytosis and Endocytosis 90
1.1 The Physical Nature of Light 6
4.7 Role of Membranes In Cell Signalling 92
1.2 Light as a Source of Energy 8
SUMMARY ILLUSTRATION FOR CHAPTER 4 96
1.3 Light as a Source of Information 10
1.4 The Uniqueness of Light 14 5 Cellular Respiration 101
1.5 Light Can Damage Biological Molecules 14 5.1 The Chemical Basis of Cellular Respiration 102
Figure 1.19 Research Method Using Spectrophotometry to 5.2 Cellular Respiration: An Overview 104
Determine an Absorption Spectrum 16 5.3 Glycolysis: The Splitting of Glucose 105
1.6 Using Light to Tell Time 17 5.4 Pyruvate Oxidation and the Citric Acid Cycle 106
1.7 The Role of Light in Behaviour and Ecology 20 5.5 Oxidative Phosphorylation: Electron Transport and
1.8 Organisms Making Their Own Light: Bioluminescence 21 Chemiosmosis 109
5.6 The Efficiency and Regulation of Cellular Respiration 114
UNIT ONE SYSTEMS AND PROCESSES: 5.7 Oxygen and Cellular Respiration 116
THE CELL 25 SUMMARY ILLUSTRATION FOR CHAPTER 5 120
2.1 Basic Features of Cell Structure and Function 26 6.1 Photosynthesis: An Overview 126
2.2 Prokaryotic Cells 30 6.2 The Photosynthetic Apparatus 128
2.3 Eukaryotic Cells 31 6.3 The Light Reactions 132
Figure 2.8 Research Method Cell Fractionation 32 6.4 The Calvin Cycle 134
6.5 Photorespiration and CO2-Concentrating Mechanisms 137
2.4 Specialized Structures of Plant Cells 43
6.6 Photosynthesis and Cellular Respiration Compared 142
2.5 The Animal Cell Surface 45
SUMMARY ILLUSTRATION FOR CHAPTER 6 144
SUMMARY ILLUSTRATION FOR CHAPTER 2 48
UNIT TWO GENES 149
3 Energy and Enzymes 53
3.1 Energy and the Laws of Thermodynamics 54 7 Cell Cycles 149
3.2 Free Energy and Spontaneous Processes 57 7.1 The Cycle of Cell Growth and Division: An Overview 150
3.3 Thermodynamics and Life 59 7.2 The Cell Cycle in Prokaryotic Organisms 151
3.4 Overview of Metabolism 61 7.3 Mitosis and the Eukaryotic Cell Cycle 152
viii NEL
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7.4 Formation and Action of the Mitotic Spindle 158 UNIT THREE DNA AND GENE EXPRESSION 251
7.5 Cell Cycle Regulation 162
11 DNA Structure, Replication, and Repair 251
Figure 7.19 Experimental Research Movement of Chromosomes
during Anaphase of Mitosis 164 11.1 Establishing DNA as the Hereditary Molecule 252
Figure 11.1 Experimental Research Griffith’s Experiment with
SUMMARY ILLUSTRATION FOR CHAPTER 7 168
Virulent and Nonvirulent Strains of Streptococcus pneumoniae 253
8 Genetic Recombination 173
Figure 11.2 Experimental Research The Hershey and Chase
8.1 Mechanism of Genetic Recombination 174 Experiment Demonstrating That DNA Is the Hereditary Molecule 255
8.2 Genetic Recombination in Bacteria 175
11.2 DNA Structure 255
Figure 8.2 Research Method Replica Plating 176
11.3 DNA Replication 258
Figure 8.3 Experimental Research Genetic Recombination in
Figure 11.9 Experimental Research The Meselson and Stahl
Bacteria 177
Experiment Demonstrating the Semiconservative Model for DNA
8.3 Genetic Recombination Occurs in Eukaryotes during Replication to Be Correct 261
Meiosis 183
11.4 Repair of Damage in DNA 270
SUMMARY ILLUSTRATION FOR CHAPTER 8 194
SUMMARY ILLUSTRATION FOR CHAPTER 11 272
9 The Chromosomal Basis of Mendelian
Inheritance 199 12 Gene Structure, Expression, and Mutation 277
9.1 The Beginnings of Genetics: Mendel’s Garden Peas 200 12.1 The Connection between DNA, RNA, and Protein 278
Figure 9.2 Research Method Making a Genetic Cross between Two Figure 12.2 Experimental Research The Gene–Enzyme
Pea Plants 201 Relationship 280
Figure 9.4 Experimental Research The Principle of Segregation: 12.2 Transcription: DNA-Directed RNA Synthesis 283
Inheritance of Flower Colour in Garden Peas 204 12.3 Processing of mRNAs in Eukaryotes 285
12.4 Translation: mRNA-Directed Polypeptide Synthesis 289
Figure 9.7 Experimental Research Testing the Predicted Outcomes
of Genetic Crosses 208 12.5 Mutations Can Affect Protein Structure and Function 299
SUMMARY ILLUSTRATION FOR CHAPTER 12 302
Figure 9.8 Experimental Research The Principle of Independent
Assortment 209 13 Regulation of Gene Expression 307
9.2 Later Modifications and Additions to Mendel’s 13.1 Regulation of Gene Expression in Prokaryotic Cells 308
Hypotheses 212 13.2 Regulation of Transcription in Eukaryotes 314
Figure 9.12 Experimental Research Experiment Showing 13.3 Posttranscriptional, Translational, and Posttranslational
Incomplete Dominance of a Trait 213 Regulation 321
13.4 The Loss of Regulatory Controls in Cancer 325
SUMMARY ILLUSTRATION FOR CHAPTER 9 220
SUMMARY ILLUSTRATION FOR CHAPTER 13 328
10 Genetic Linkage, Sex Linkage, and Other Non-
Mendelian Inheritance Mechanisms 225 14 DNA Technologies 333
10.1 Genetic Linkage and Recombination 226 14.1 DNA Cloning 334
Figure 10.2 Experimental Research Evidence for Gene Figure 14.3 Research Method Identifying a Recombinant Plasmid
Linkage 228 Containing a Gene of Interest 337
10.2 Sex-Linked Genes 231 Figure 14.4 Research Method Synthesis of DNA from mRNA Using
Figure 10.8 Experimental Research Evidence for Sex-Linked Reverse Transcriptase 338
Genes 234
Figure 14.5 Research Method The Polymerase Chain Reaction
10.3 Chromosomal Mutations That Affect Inheritance 236 (PCR) 339
10.4 Human Genetic Traits, Pedigree Analysis, and Genetic
Counselling 240 Figure 14.6 Research Method Separation of DNA Fragments by
Agarose Gel Electrophoresis 340
10.5 Additional Non-Mendelian Patterns of Inheritance 244
SUMMARY ILLUSTRATION FOR CHAPTER 10 246 14.2 Applications of DNA Technologies 341
NEL C O N T E N T S
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Figure 14.8 Research Method Southern Blot Analysis 343 17.4 Non-random Mating 423
17.5 Maintaining Genetic and Phenotypic Variation 425
Figure 14.11 Research Method Making a Knockout Mouse 347
Figure 17.16 Experimental Research Sexual Selection in Action 426
Research in Biology CRISPR: A Programmable RNA-Guided Genome
Editing System 348 SUMMARY ILLUSTRATION FOR CHAPTER 17 428
Figure 14.13 Experimental Research The First Cloning of a 18 Speciation and Macroevolution 433
Mammal 351 18.1 What Is a Species? 434
18.2 Maintaining Reproductive Isolation 437
Figure 14.15 Research Method Using the Ti Plasmid of Rhizobium
radiobacter to Produce Transgenic Plants 352 18.3 The Geography of Speciation 440
18.4 Genetic Mechanisms of Speciation 443
SUMMARY ILLUSTRATION FOR CHAPTER 14 356
Figure 18.16 Observational Research Chromosomal Similarities
15 Genomes 361 and Differences among Humans and the Great Apes 447
Figure 16.16 Experimental Research Adaptation of E. coli to a 20.3 Human Features That Do Not Fossilize 484
Change in Temperature 404 20.4 Dispersal of Early Humans 485
20.5 Hominins and the Species Concepts 485
SUMMARY ILLUSTRATION FOR CHAPTER 16 406
SUMMARY ILLUSTRATION FOR CHAPTER 20 488
17 Microevolution: Changes within Populations 411
17.1 Variation in Natural Populations 412
17.2 Population Genetics 414
UNIT FIVE THE DIVERSITY OF LIFE 493
17.3 The Agents of Microevolution 416 21 Defining Life and Its Origins 493
Figure 17.7 Research Method Using the Hardy–Weinberg 21.1 What Is Life? 494
Principle 417 21.2 The Chemical Origins of Life 494
Figure 17.12 Experimental Research Do Humans Experience 21.3 From Macromolecules to Life 499
Stabilizing Selection? 422 21.4 Evidence of the Earliest Life 502
x C O N T E N T S NEL
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21.5 Eukaryotes and the Rise of Multicellularity 504 Figure 25.20 Experimental Research Hidden Third Partner in Lichen
21.6 The Fossil Record 507 Symbiosis 602
21. 7 The Tree of Life 509 SUMMARY ILLUSTRATION FOR CHAPTER 25 606
Figure 21.23 Research Method Radiometric Dating 511
26 Plants 611
SUMMARY ILLUSTRATION FOR CHAPTER 21 516 26.1 Defining Characteristics of Land Plants 612
26.2 The Transition to Life on Land 613
22 Viruses, Viroids, and Prions: Infectious Biological
Particles 521 26.3 Bryophytes: Nonvascular Land Plants 619
22.1 What Is a Virus? Characteristics of Viruses 522 26.4 Seedless Vascular Plants 622
22.2 Viruses Infect Bacterial, Animal, and Plant Cells by Similar 26.5 Gymnosperms: The First Seed Plants 628
Pathways 524 26.6 Angiosperms: Flowering Plants 632
22.3 Treating and Preventing Viral Infections 528 Figure 26.30 Experimental Research Exploring a Possible Early
Figure 22.7 Experimental Research A New Discovery for Hepatitis C Angiosperm Adaptation for Efficient Photosynthesis in Dim
Therapy 529 Environments 635
SUMMARY ILLUSTRATION FOR CHAPTER 26 638
22.4 Viruses May Have Evolved from Fragments of Cellular DNA or
RNA 530
27 Animals 643
22.5 Viroids and Prions Are Infective Agents Even Simpler in
Structure than Viruses 530 27.1 What Is an Animal? 644
SUMMARY ILLUSTRATION FOR CHAPTER 22 532 27.2 Key Innovations in Animal Evolution 645
27.3 Molecular Phylogenetics and Classification 649
23 Bacteria and Archaea 537 27.4 The Basal Phyla 650
23.1 The Full Extent of the Diversity of Bacteria and Archaea Is 27.5 The Protostomes 658
Unknown 538 27.6 Lophotrochozoa Protostomes 659
23.2 Prokaryotic Structure and Function 538 27.7 Ecdysozoa Protostomes 670
Figure 23.5 Experimental Research Genetic Recombination in 27.8 The Deuterostomes 680
Bacteria 541
Research in Biology The Tully Monster 686
23.3 The Domain Bacteria 548
27.9 The Origin and Diversification of Vertebrates 687
23.4 The Domain Archaea 550
27.10 Agnathans: The Jawless Fishes 689
SUMMARY ILLUSTRATION FOR CHAPTER 23 554
27.11 Jawed Fishes: Jaws Meant New Feeding Opportunities 690
24.1 The Vast Majority of Eukaryotes Are Protists 560 27.13 The Origin and Mesozoic Radiations of Amniotes 698
24.2 Characteristics of Protists 561 27.14 Turtles and Tortoises (Subclass Testudinata) 702
24.3 Protists’ Diversity Is Reflected in Their Metabolism, 27.15 Living Diapsids: Sphenodontids, Squamates, and
Reproduction, Structure, and Habitat 562 Crocodylians 702
24.4 The Eukaryotic Supergroups and Their Key Protist Lineages 563 27.16 Birds 704
27.17 Mammalia: Monotremes, Marsupials, and Placentals 708
Figure 24.8 Observational Research Isolation and Identification
of Marine Diplonemids, Potentially the Most Abundant Marine SUMMARY ILLUSTRATION FOR CHAPTER 27 714
Organism 565
28 Conservation of Biodiversity 719
24.5 Some Protist Lineages Arose from Primary Endosymbiosis and
28.1 The Anthropocene 720
Others from Secondary Endosymbiosis 580
28.2 Vulnerability to Extinction 721
SUMMARY ILLUSTRATION FOR CHAPTER 24 582
28.3 Climate Change Can Cause Extinction 725
25 Fungi 587 28.4 Protecting Species 726
25.1 General Characteristics of Fungi 588 28.5 Protecting What? 727
25.2 Evolution and Diversity of Fungi 590 28.6 Conservation and Agriculture 729
25.3 Fungal Lifestyles 600 28.7 Contaminating Natural Systems 730
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28.8 Motivation 736 31 Ecosystems 813
28.9 Effecting Conservation 737 31.1 Connections Within and Among Ecosystems 815
Figure 28.33 Observational Research Near-Complete Extinction of 31.2 Ecosystems and Energy 816
Small Mammals in Tropical Forest Fragments 738 31.3 Nutrient Cycling in Ecosystems 823
28.10 Human Population: A Root Problem for Conservation 739 31.4 Carbon: A Disrupted Cycle 830
28.11 Signs of Stress: Systems and Species 739 31.5 Ecosystem Modelling 832
SUMMARY ILLUSTRATION FOR CHAPTER 28 742 31.7 Three Sample Ecosystems 834
SUMMARY ILLUSTRATION FOR CHAPTER 31 840
xii C O N T E N T S NEL
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Carbohydrates F-24 Figure 37.4 Experimental Research Two Experiments by Frits
Proteins F-28 Went Demonstrating the Effect of Indoleacetic Acid (IAA) on an Oat
Nucleic Acids F-36 Coleoptile 983
34.3 Long-Distance Transport of Water and Minerals in the Xylem 919 39.1 Nutrients Are Essential Components of Any Diet 1030
34.4 Transport of Organic Substances in the Phloem 914 39.2 Feeding: Obtaining Nutrients 1034
Figure 34.13 Experimental Research Translocation Pressure 925 39.3 Digestive Processes 1036
39.4 Structure and Function of the Mammalian Digestive Tract 1039
SUMMARY ILLUSTRATION FOR CHAPTER 34 928
39.5 Regulation of Digestive Processes 1048
35 Reproduction and Development in Flowering Plants 933 Figure 39.20 Experimental Research Association of Bacterial
Populations in the Gut Microbiome with Obesity in Humans 1050
35.1 Overview of Flowering Plant Reproduction 934
35.2 Flower Structure and Formation of Gametes 936 SUMMARY ILLUSTRATION FOR CHAPTER 39 1052
35.3 Pollination, Fertilization, and Germination 940
35.4 Asexual Reproduction in Flowering Plants 947 40 Gas Exchange: The Respiratory System 1057
40.1 General Principles 1058
Figure 35.16 Research Method Plant Tissue Culture Protocol 948
40.2 Gas Exchange Organs 1061
35.5 Early Development of Plant Form and Function 949
40.3 The Mammalian Respiratory System 1065
SUMMARY ILLUSTRATION FOR CHAPTER 35 950 40.4 Exchange of Gas with Blood 1068
36.1 Plant Nutritional Requirements 956 Figure 40.21 Experimental Research Demonstration of a Molecular
Basis for High-Altitude Adaptation in Deer Mice 1073
Figure 36.2 Research Method Hydroponic Culture 957
Figure 37.3 Experimental Research The Darwins’ Experiments on 41.4 Blood Vessels of the Circulatory System 1090
Phototropism 982 41.5 Maintaining Blood Flow and Pressure 1093
NEL C O N T E N T S
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Figure 41.17 Experimental Research Demonstration of a 44.3 Mechanisms of Sexual Reproduction 1164
Vasodilatory Signalling Molecule 1094 44.4 Sexual Reproduction in Mammals 1171
41.6 The Lymphatic System 1095 Figure 44.11 Experimental Research Vocal Cues to Ovulation in
Human Females 1172
SUMMARY ILLUSTRATION FOR CHAPTER 41 1098
44.5 Development 1180
42 Regulation of the Internal Environment: Water, Solutes,
and Temperature 1103 SUMMARY ILLUSTRATION FOR CHAPTER 44 1194
42.1 Introduction to Osmoregulation and Excretion 1104 45 Control of Animal Processes: Neural Control 1199
42.2 Osmoregulation and Excretion in Invertebrates 1108 45.1 The Basis of Information Flow in Nervous Systems:
42.3 Osmoregulation and Excretion in Non-mammalian An Overview 1200
Vertebrates 1110
Figure 45.13 Experimental Research Demonstration of Chemical
42.4 Osmoregulation and Excretion in Mammals 1112 Transmission of Nerve Impulses at Synapses 1212
Figure 42.15 Experimental Research ADH-Stimulated Water
45.2 Sensory Inputs: Reception 1214
Reabsorption in the Kidney Collecting Duct 1118
Figure 45.25 Experimental Research How Do Sea Urchins Detect
42.5 Introduction to Thermoregulation 1120 Light? 1223
42.6 Ectothermy 1122
Figure 45.42 Experimental Research Magnetic Sense in Sea
42.7 Endothermy 1125
Turtles 1234
SUMMARY ILLUSTRATION FOR CHAPTER 42 1130
45.3 The Central Nervous System: Integration 1236
43 Control of Animal Processes: Endocrine Control 1135 45.4 The Peripheral Nervous System: Transmission and
43.1 Hormones and Their Secretion 1136 Response 1247
43.2 Mechanisms of Hormone Action 1139 SUMMARY ILLUSTRATION FOR CHAPTER 45 1250
Figure 43.6 Experimental Research Demonstration That
Epinephrine Acts by Binding to a Plasma Membrane Receptor 1142
46 Muscles, Skeletons, and Body Movements 1255
46.1 Vertebrate Skeletal Muscle: Structure and Function 1256
43.3 The Hypothalamus and Pituitary 1145
Figure 46.5 Experimental Research The Sliding Filament Model of
43.4 Other Major Endocrine Glands of Vertebrates 1148 Muscle Contraction 1259
43.5 Endocrine Systems in Invertebrates 1153
46.2 Skeletal Systems 1264
Figure 43.16 Experimental Research Demonstration That Growth
46.3 Vertebrate Movement: The Interactions between Muscles and
and Moulting in Insects Is Hormonally Controlled 1155
Bones 1267
SUMMARY ILLUSTRATION FOR CHAPTER 43 1156 SUMMARY ILLUSTRATION FOR CHAPTER 46 1272
xiv C O N T E N T S NEL
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Preface
Welcome to an exploration of the diversity of life. The main goal Emphasizing the big picture …
of this textbook is to guide you on a journey of discovery about
Many biology textbooks use the first few chapters to review fun-
life’s diversity across levels ranging from molecules to genes,
damentals of chemistry and biochemistry as well as information
cells to organs, and species to ecosystems. Along the way, we
on the scientific method. Instead of focusing on this background
will explore many questions about the mechanisms underlying
information, we have used the first chapter, in particular, to
diversity as well as the consequences of diversity, for our own
immediately engage students by conveying the excitement that
species and for others.
is modern biology. We have put important background infor-
mation in the centre of the book as a distinct reference section
An emphasis on the diversity of life … entitled The Chemical and Physical Foundations of Biology. With
their purple borders, these pages are distinct and easy to find,
At first glance, the riot of life that animates the biosphere over-
and have become affectionately known as The Purple Pages.
whelms our minds. One way to begin to make sense of this
These pages enable information to be readily identifiable and
diversity is to divide it into manageable sections on the basis of
accessible to students as they move through the textbook rather
differences. We also consider features found in all life forms to
than being tied to a particular chapter. In this edition, the con-
stress similarities as well as differences. We examine how dif-
cepts of atoms, molecules, and macromolecules are connected
ferent organisms solve the common problems of finding nutri-
through the theme of “emergent properties.” By considering
ents, energy, and mates on the third rock from our Sun. What
how the “stuff of life” interrelates as a function of increasing
basic evolutionary principles inform the relationships among
complexity rather than just memorizing the attributes of indi-
life forms regardless of their different body plans, habitats, or
vidual items, students can better grasp why biology works the
life histories? Unlike many other first-year biology texts, this
way it does, rather than be awed by how much information we
book has chapters integrating basic concepts such as the effects
know about it.
of genetic recombination, light, and domestication across the
We hope that Canadian students will find the subject of
breadth of life from microbes to mistletoe to moose. As you read
biology as it is presented here accessible and engaging because it
this book, you will be referred frequently to other chapters for
is presented in familiar contexts. We have highlighted the work
linked information that expands the ideas further.
of Canadian scientists, used examples of Canadian species, and
Evolution provides a powerful conceptual lens for viewing
referred to Canadian regulations and institutions.
and understanding the roots and history of the diversity of
living things. We will demonstrate how knowledge of evolution
helps us appreciate the changes we observe in organisms. Focusing on research to help students engage
Whether the focus is the conversion of free-living prokaryotic
the living world as scientists …
organisms into mitochondria and chloroplasts or the steps
involved in the domestication of rice, selection for particular A primary goal of this book is to evoke and sustain students’
traits over time can explain the current condition. curiosity about biology, rather than dulling it with a moun-
Examining how biological systems work is another theme tain of disconnected facts. We can help students develop the
pervading this text and underlying the idea of diversity. We have mental habits of scientists and a fascination with the living
intentionally tried to include examples that will tax your imagi- world by conveying our passion for biological research. We
nation, from sea slugs that steal chloroplasts for use as solar want to excite students not only with what biologists know
panels, to the molecular basis of high altitude adaptations in about the living world but also with how they know it and
deer mice, to adaptive radiation of viruses. In each situation, we what they still need to learn. In doing so, we can encourage
examine how biologists have explored and assessed the inner some students to accept the challenge and become biologists
workings of organisms, from gene regulation to the challenges themselves, posing and answering important new questions
of digesting cellulose. through their own innovative research. For students who
Solving problems is another theme that runs throughout pursue other careers, we hope that they will leave their
the book. Whether the topic is gene therapy to treat a disease in introductory—and perhaps only—biology course armed with
people, increasing crop production, or reducing the incidence of intellectual skills that will enable them to evaluate future
human obesity, both the problem and the solution lie in biology. knowledge with a critical eye.
We will explore large problems facing planet Earth and the In this book, we introduce students to a biologist’s “ways of
social implications that arise from them. learning.” Research biologists constantly integrate new observations,
NEL xv
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hypotheses, questions, experiments, and insights with existing examples and tells the story of how a researcher or researchers
knowledge and ideas. To help students engage the world as arrived at a key insight, or how biological research solved a
biologists do, we must not simply introduce them to the cur- major societal problem, explained a fundamental process, or
rent state of knowledge, we must also foster an appreciation of elucidated a phenomenon. The Why It Matters feature links
the historical context within which those ideas developed, and the insight from the vignette to the contents of the chapter to
identify the future directions that biological research is likely to spark student interest in the topic at hand.
take. ●● Three types of specially designed research figures provide
Because advances in science occur against a background of more detailed information about how biologists formulate
research, we also give students a feeling for how biologists of the specific hypotheses and test them by gathering and inter-
past formulated basic knowledge in the field. By fostering an preting data. Experimental Research figures describe spe-
appreciation of such discoveries, given the information and the- cific studies in which researchers used both experimental
ories available to scientists in their own time, we can help stu- and control treatments, either in the laboratory or in the
dents understand the successes and limitations of what we field, to test hypotheses or answer research questions by
consider cutting edge today. This historical perspective also manipulating the system they studied. Observational
encourages students to view biology as a dynamic intellectual Research figures describe specific studies in which biolo-
enterprise, not just a collection of facts and generalities to be gists have tested hypotheses by comparing systems under
memorized. varying natural circumstances. Research Method figures
We have endeavoured to make the science of biology come provide examples of important techniques, such as light
alive by describing how biologists formulate hypotheses and and electron microscopy, the polymerase chain reaction,
evaluate them using hard-won data; how data sometimes tell making a knockout mouse, DNA microarray analysis, plant
only part of a story; and how the results of studies often end up cell culture, producing monoclonal antibodies, radiometric
posing more questions than they answer. Our exploration of the dating, and cladistic analysis. Each Research Method figure
Tully Monster in Chapter 27 is a case in point. Since its fossil leads a student through the purpose of the technique and
discovery and description, this mainly soft-bodied animal has protocol, and describes how scientists interpret the data it
been tentatively classified with species in five different groups of generates.
animals. Through this example, and throughout Chapter 27, we
explore the current recognition that the historical and tradi-
tional grouping of animals into protostomes and deuterostomes Integrating effective, high-quality visuals into
is more artificial than real. the narrative …
Although students might prefer simply to learn the “right”
answer to a question, they must be encouraged to embrace Today’s students are accustomed to receiving ideas and informa-
“the unknown,” those gaps in knowledge that create opportu- tion visually, making the illustrations and photographs in a text-
nities for further research. An appreciation of what biologists book and the fully integrated online resources critically
do not yet know will draw more students into the field. And by important. From the first Canadian edition, our illustration pro-
defining why scientists do not understand interesting phe- gram has provided an exceptionally clear supplement to the nar-
nomena, we encourage students to think critically about pos- rative in a style that is consistent throughout the book. Graphs
sible solutions and to follow paths dictated by their own and anatomical drawings are annotated with interpretive expla-
curiosity. We hope that this approach will encourage students nations that lead students, step by step, through the major points
to make biology a part of their daily lives by having informal they convey.
discussions and debates about new scientific discoveries. Over subsequent editions, we have continued to enhance
the illustration program, focusing on features that reviewers and
users of the book identified as the most useful pedagogical tools.
Presenting the story line of the research In revising the text, we reevaluated each illustration and photo-
process … graph, and made appropriate changes to improve their utility as
teaching tools.
Science is by its nature a progressive enterprise in which answers
For this most recent edition, we have made some exciting
to questions open new questions for consideration. In preparing
new additions to our illustration program through the creation
this book, we developed several special features to help students
of Chapter Roadmaps and Summary Illustrations for every
broaden their understanding of the material presented and of
chapter the book. Chapter Roadmaps appear at the beginning of
the research process itself:
each chapter and provide a visual overview of the chapter con-
●● The chapter openers, titled Why It Matters, are engaging, tents. Connections between topics across chapters are empha-
short vignettes designed to capture students’ imaginations sized to give students a sense of how the content of each chapter
and whet their appetites for the topic that the chapter fits within the larger context of the book, and biology as a whole.
addresses. In many cases, this feature uses current Canadian At the end of each chapter, we have created vivid and engaging
xvi P R E FA C E NEL
Copyright 2019 Nelson Education Ltd. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content
may be suppressed from the eBook and/or eChapter(s). Nelson Education reserves the right to remove additional content at any time if subsequent rights restrictions require it.
Summary Illustrations that depict the core concepts—and Effectively introducing digital solutions into
teaching heart—of the chapter. These illustrations provide stu- your classroom—online or in class—is now
dents with a visual overview of the connections between key
concepts, and provide a unique touchstone to review and gauge
easier than ever …
understanding of the chapter contents. The fourth Canadian edition of Biology: Exploring the Diversity
of Life represents a fully integrated package of print and media,
providing comprehensive learning tools and flexible delivery
options. In preparing this edition, we conducted extensive
Organizing chapters around important research to determine how instructors prefer to present online
concepts … learning opportunities. The result of this research is a new
As authors and university teachers, we understand how easily MindTap course organized around the instructors’ preferred
students can get lost within a chapter. When students request workflow. Instructors can now select just the content they want
advice about how to read a chapter and learn the material in it, to assign, chosen from a comprehensive set of learning materials
we usually suggest that, after reading each section, they pause provided with the course for each chapter. Many types of
and quiz themselves on the material they have just encountered. learning activities are assignable and offer students immediate
After completing all the sections in a chapter, they should quiz feedback and automated instructor assessment.
themselves again, even more rigorously, on the individual sec- Research also indicates that online content is most effective
tions and, most important, on how the concepts that were devel- when it enhances conceptual understanding through the use of
oped in the different sections fit together. Accordingly, we have relevant applications. In this edition, we have developed new
adopted a structure for each chapter to help students review assessable online learning activities that provide students the
concepts as they learn them. opportunity to explore and practice biology the way scientists
practice biology:
●● The organization within chapters presents material in digest-
Interpret the Data exercises have been enhanced by an addi-
ible sections, building on students’ knowledge and under-
●●
NEL P R E FA C E
xvii
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may be suppressed from the eBook and/or eChapter(s). Nelson Education reserves the right to remove additional content at any time if subsequent rights restrictions require it.
alance between factual and conceptual material, taking great
b search for understanding, and, most important, observe and
care to provide clear explanations of how scientists draw conclu- enjoy the diversity of life around you.
sions from empirical data. Our approach helps students under- M. Brock Fenton
stand how we achieved our present knowledge. Clarity of Denis Maxwell
presentation, thoughtful organization, a logical and seamless Tom Haffie
flow of topics within chapters, and carefully designed illustra- Bill Milsom
tions are key to our approach. Todd Nickle
We hope that you are as captivated by the biological world Shona Ellis
as we are, and are drawn from one chapter to another. But don’t London, Calgary, and Vancouver
stop there; use the digital and other resources to broaden your January 2018
xviii P R E FA C E NEL
Copyright 2019 Nelson Education Ltd. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content
may be suppressed from the eBook and/or eChapter(s). Nelson Education reserves the right to remove additional content at any time if subsequent rights restrictions require it.
New to This Edition
The enhancements we have made in the fourth Canadian edition of behind Biology,” and “Life on the Edge” have also been moved
Biology: Exploring the Diversity of Life reflect our commitment to from the book to the Instructor’s Manual, allowing instructors to
providing a textbook that introduces students to new developments continue to draw upon these engaging stories and vignettes,
in biology while fostering active learning and critical thinking. without increasing the length of the textbook.
Our revisions to the new edition were guided by five impor-
tant principles: Engaging and relevant content …
●● Reduce the size of the book From personal genome reports to cues to recognizing human
●● Ensure content is relevant and engaging for students and female ovulation, the new edition is full of engaging examples
instructors that reflect everyday biology and its impact on society. In addi-
●● Emphasize connections tion to references to Canadian research and researchers
●● Support concepts with visuals wherever possible throughout the book, our MindTap features profiles of 13 former
●● Extensively revise and rewrite Unit Four: Evolution and biology students, and what they have done with their biology
Classification degrees in “Where Are They Now?”
NEL xix
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may be suppressed from the eBook and/or eChapter(s). Nelson Education reserves the right to remove additional content at any time if subsequent rights restrictions require it.
of evolutionary principles; whether it’s the importance of chance Chapter 6: Photosynthesis
mutation as a driver of evolutionary change or that evolution
can occur in the absence of natural selection. ●● Clarified and improved selected figures
With this in mind, we have extensively revised this unit to
focus more clearly on conveying the fundamental concepts of Chapter 7: Cell Cycles
evolution, to provide greater clarity on the processes that cause
NEW Why It Matters about algal blooms in Lake Erie
evolutionary change, as well as to make critical connections
●●
chromosomes
focused on experimental evolution in E. coli, as well as a con-
cluding figure that explains the major misconceptions students
have concerning evolution and natural selection. Chapter 17: Chapter 8: Genetic Recombination
Microevolution: Changes Within Populations has been rewritten
●● Added explicit reference to cytokinesis
to make stronger connections to genetics, which are often not
●● Specified creation of haploid cells
made in the context of evolution, by fully explaining terms such
as allele, gene, gene pool, and locus. This chapter also emphasizes
the role of random mutation in evolution and its importance in Chapter 9: The Chromosomal Basis of Mendelian
introducing genetic novelty. Chapter 18: Speciation and Macro- Inheritance
evolution has improved flow and clarity, including simpler and
more informative figures. Chapter 19: Systematics and Phyloge-
●● NEW Canadian Why It Matters about the spirit bears of
netics: Revealing the Tree of Life is now its own dedicated British Columbia
chapter. This allows for more clear discussion of the tools and
●● Enhanced discussion connecting genes/alleles to proteins
approaches used today to infer evolutionary histories. Great and protein products, and to the expression of alleles in the
care has been taken to clearly define and present concepts of phenotype as dominant/recessive
homology and convergent evolution.
Chapter 10: Genetic Linkage, Sex Linkage, and
Other Non-Mendelian Inheritance Mechanisms
Major revisions to selected chapters are listed
NEW Canadian Why It Matters about disease incidence in
below:
●●
Quebec
Chapter 1: Light and Life ●● NEW figures and examples dealing with translocations,
imprinting, and pedigree analysis
●● Streamlined to be more concise
thermodynamics
●● Improved and clarified figures related to exergonic and end- Mutation
ergonic reactions ●● NEW Canadian Why It Matters about poisonous mush-
rooms in British Columbia
Chapter 4: Cell Membranes and Signalling ●● Expanded material on mutations and how they can affect
protein function
●● NEW Research Figure: Frye–Edidin Experiment Demon- ●● NEW discussion about ENCODE versus the junk DNA
strating that the Phospholipid Bilayer Is Fluid debate
●● NEW Research Figure: Freeze Fracture ●● Expanded and clarified discussion of mutagenesis
Copyright 2019 Nelson Education Ltd. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content
may be suppressed from the eBook and/or eChapter(s). Nelson Education reserves the right to remove additional content at any time if subsequent rights restrictions require it.
●● Updated material on lncRNA ●● The concepts of homology and convergent evolution are
●● Updated material on cancer genetics more clearly presented.
Diplonemids
●● Emphasis on the role of random mutation, the different
types and when they occur, that may drive the introduction
of genetic novelty Chapter 25: Fungi
●● Incorporated information on a recent discovery in lichens
Chapter 18: Speciation and Macroevolution related to the third symbiont, a basidiomycete yeast that is
●● Improved flow and clarity of the writing part of the symbiosis that influences the morphology of
●● Simpler and more informative figures lichen
●● Added material on the Puccinia–grain interaction
NEW Research Figure: Hidden Third Partner in Lichen
Chapter 19: Systematics and Phylogenetics: ●●
Symbiosis
Revealing the Tree of Life
●● In the previous edition, systematics and phylogenetics were Chapter 26: Plants
grouped with the history of life (geological record) as a single
chapter. This made it somewhat disjointed. In this edition, ●● NEW Research Figure: Exploring a Possible Early Angio-
systematics and phylogenetics is its own dedicated chapter. sperm Adaptation for Efficient Photosynthesis in Dim
This allows for clearer discussion of the tools and approaches Environments
used today to infer evolutionary histories. ●● Extensive revision of key figures for clarity
NEL N E W TO T H I S E D I T I O N
xxi
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may be suppressed from the eBook and/or eChapter(s). Nelson Education reserves the right to remove additional content at any time if subsequent rights restrictions require it.
Chapter 27: Animals Chapter 35: Reproduction and Development in
●● NEW chapter that combines both protostomes and Flowering Plants
deuterostomes ●● NEW section explaining the genetics behind the ABC model
●● Includes latest research on phylogenetic tree of floral development
●● NEW Research in Biology box on the Tully Monster ●● NEW section showing how plant tissue culture can generate
virus-free plants from infected donors
Chapter 28: Conservation of Biodiversity
●● NEW Why It Matters featuring the extinction of passenger Chapter 36: Plant Nutrition
pigeons ●● Assimilation of nutrients connected with material in The
●● NEW section on the Anthropocene Purple Pages
●● Enhanced discussion of human impact on landscapes
NEW material on ecosystem services
Chapter 37: Plant Signals and Responses to the
●●
●● NEW Why It Matters about cave ecosystems Chapter 40: Gas Exchange: The Respiratory
Updated discussion of mass mortality
System
●●
Copyright 2019 Nelson Education Ltd. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content
may be suppressed from the eBook and/or eChapter(s). Nelson Education reserves the right to remove additional content at any time if subsequent rights restrictions require it.
●● Added new section on the regulation of mammalian kidney Chapter 45: Control of Animal Processes: Neural
function Control
●● Refined discussion to clarify the difference between osmo-
lality and osmolarity ●● NEW chapter created by combining neural control and
integration
Clarified the difference between the spike initiation zone
Chapter 43: Control of Animal Processes:
●●
development
●● Added sexual reproduction as a route for infection ●● NEW Concept Fix addressing the misconceptions about
●● Clarified discussion of where organelles were in the muscles getting smaller as they contract
sperm ●● More human examples integrated throughout
●● NEW Research Figure: Vocal Cues of Ovulation in Human ●● NEW Research Figure: The Sliding Filament Model of
Females Muscle Contraction
NEL N E W TO T H I S E D I T I O N
xxiii
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may be suppressed from the eBook and/or eChapter(s). Nelson Education reserves the right to remove additional content at any time if subsequent rights restrictions require it.
Another random document with
no related content on Scribd:
Fredensborg zu fahren. Sie sah, daß es ihm gut tat, wenn sie sich
munter und frisch gab.
Sie dachte kaum mehr nach. Ohne Anstrengung konnte sie ihre
Gedanken ausschalten. In Wirklichkeit war ihr Gehirn kraftlos. Wie
ein dauerndes mahnendes Erinnern war es nur, daß sich ihr die
Brust schmerzhaft spannte und daß das Korsett sie behinderte.
Jenny hatte sich bei einer Lehrerswitwe auf Westseeland
eingemietet. Gram begleitete sie dorthinaus und reiste am Abend
nach Kopenhagen zurück. So war sie endlich allein.
Sie hatte aufs Geratewohl gemietet. Während ihres
Studienaufenthaltes in Kopenhagen war sie einen Tag über mit
einigen Kameradinnen in dem Dorfe gewesen; sie hatten im Krug
gegessen und bei den Dünen gebadet. Sie entsann sich, daß es dort
schön war, und als auf ihre Anzeige eine Frau Rasmussen dort sich
erboten hatte, die junge Dame aufzunehmen, die ihre Niederkunft
erwartete, da griff sie zu.
Eigentlich fühlte sie sich wohl. Allerdings wohnte die
Lehrerswitwe in einem elend häßlichen, winzigkleinen gelben
Backsteinhaus etwas außerhalb des Dorfes, an der Landstraße, die
sich staubig und ohne Ende zwischen offenen, bestellten Feldern
hinzog. Aber Jenny mochte ihr Zimmer gern mit der Tapete in
Berlinerblau und den Lithographien nach Exner an den Wänden, mit
den weißen, gehäkelten Deckchen ringsumher, auf dem Bett, dem
amerikanischen Schaukelstuhl und der Kommode, auf die Frau
Rasmussen bei Jennys Ankunft einen großen Rosenstrauß gestellt
hatte.
Draußen vor den beiden Fensterchen lief die Landstraße vorbei.
Im Vorgärtchen blühten Rosen, Geranien und „Christi Blutstropfen“,
ungeachtet all des Staubes, mit dem sie gepudert waren. Jenseits
der Straße erhob sich ein nackter Hügelkamm im Acker. Steinwälle,
an deren Hängen struppige, leuchtende Herbstblumen zwischen
Brombeerhecken wucherten, teilten den Hügel in weiße
Stoppelfelder, blaugrüne Rübenäcker und braungrüne Wiesen,
umrändert von zackigen, zerzausten Weidenbüschen. War die
Abendsonne aus Jennys Kammer geschwunden, so flammte der
Himmel über dem Hügelkamm und den spärlichen Zweiglein der
Weiden auf.
Hinter ihrem Zimmer ging eine kleine puppenstubenartige Küche
mit rotem Backsteinfußboden, auf den Hof hinaus, wo die Hühner
der Witwe gackerten und die Tauben gurrten. Ein kleiner Flur lief
quer durch das ganze Haus. Auf der anderen Seite lag Frau
Rasmussens Stube mit Blumen vor den Fenstern und gehäkelten
Decken überall, Daguerreotypien und Photographien an den
Wänden, einem kleinen Bücherschrank mit religiösen Schriften in
schwarzem Pappeinband, einigen Jahrgängen von „Frem“ und
Gyldendals Serien in Prachtbänden. Dahinter befand sich ein
Zimmerchen, in dem die Luft immer merkwürdig dick war, und das
ein unbestimmbarer Geruch erfüllte, obgleich es vor Sauberkeit
blitzte. Hier schlief sie selbst und konnte nicht hören, wenn sich ihre
Pensionärin jenseits des Ganges Nacht für Nacht in den Schlaf
weinte.
Frau Rasmussen war übrigens nicht so schlimm. Groß und
schlottrig schlürfte sie in einer Art von Filzschuhen leise umher;
immer sah sie gleichmäßig sorgenvoll aus mit ihrem langen gelben
Pferdegesicht, unter dem graugesprenkelten Haar, das mit einem
drolligen kleinen Schwung über jedem Ohr weggestrichen war. Sie
sprach fast gar nicht, höchstens stellte sie einige besorgte Fragen,
ob das gnädige Fräulein mit dem Essen und dem Zimmer zufrieden
sei. Selbst wenn Jenny nach dem Mittagessen sich hin und wieder
mit ihrer Handarbeit in das Wohnzimmer zu Frau Rasmussen setzte,
schwiegen sie still. Jenny war ihr besonders dafür dankbar, daß Frau
Rasmussen niemals ihren Zustand erwähnte; nur ein einziges Mal
hatte sie ängstlich gefragt, als Jenny mit ihrem Malgerät hinausging,
ob das wohl dem gnädigen Fräulein nicht schaden könnte.
Sie arbeitete in der ersten Zeit eifrig, stand hinter einem Steinwall
mit ihrer Feldstaffelei, die der scharfe Wind fast umblies. Unter dem
Wall erstreckte sich das gelbe Stoppelfeld eines endlosen
Roggenackers bis zum Moor hinab, wo sich weißes Wollgras an
blauen Wasserlöchern hinzog, wo samtschwarze Torfmieten auf
saftiggrünem Wiesengrund lagen. Hinter dem Sumpf wellte sich das
Land mit grünen Rübenfeldern, Wiesen und gemähten
Roggenäckern, mit kalkweißen Bauernhöfen in üppigen,
dunkelgrünen Hainen — bis hinüber zum frischen, blauen Fjord. Der
Strand lief in Bogen und Zungen, mit weißgelbem Sand und kurzem,
vergilbtem Gras, in die See hinaus. Gegen Norden fiel der Hügel,
vom Heidekraut braun gefärbt, mit der Windmühle auf der Spitze, in
steilen, gelben Sanddünen zum blauen Fjord ab. Schatten und Licht
wechselten auf dem offenen Lande ab, je nachdem wie die Wolken
über den weißen, ewig blauen unruhigen Himmel wanderten.
Wenn Jenny müde wurde, legte sie sich am Walle nieder, starrte
in den Himmel und über den Fjord. Sie konnte nicht längere Zeit
hintereinander stehen, doch das reizte sie nur, weiterzuarbeiten.
Zwei kleinere Bilder vollendete sie oben auf dem Wall, und freute
sich selbst an ihnen. Eines malte sie vom Dorfe unten, wo die
weißgekalkten Häuser, umgeben von Kletterrosen und Georginen,
um einen sammetgrünen Dorfteich lagen. Ihre Strohdächer hingen
bis über die Fensterscheiben hinab, die rote Backsteinkirche erhob
ihren treppengiebligen Turm über die Laubmassen des
Pfarrhausgartens. Es machte sie aber nervös, daß die Leute zu ihr
kamen und ihr zusahen; die weißhaarigen Jungen umstanden sie in
Knäueln, während sie malte. Als das Bild dann fertig war, zog sie mit
ihrer Staffelei wieder hinauf zum Wall, der See entgegen.
Dann kam aber im Oktober der Regen; es goß ein oder zwei
Wochen lang. Ab und zu klärte es sich etwas auf und ein trüber,
gelber Lichtstreifen glitt durch die Wolken, über dem Hügel mit den
traurigen Weidenbüschen hin, und die Wasserpfützen lagen ein
Weilchen blank da. Dann regnete es wieder.
Jenny lieh sich Frau Rasmussens Bücher und ließ sich die
Muster der gestrickten Spitzen an ihren Gardinen zeigen. Aber es
wurde nicht viel, weder mit dem Lesen noch mit dem Stricken. Sie
saß den lieben langen Tag im Schaukelstuhl am Fenster und war
nicht einmal imstande, sich ordentlich anzuziehen, sondern schlüpfte
nur in ihren verwaschenen Kimono.
Sie litt furchtbar darunter, daß ihre Schwangerschaft nach und
nach sichtbar wurde.
Da meldete Gert Gram seinen Besuch. Schon zwei Tage später
kam er am frühen Morgen in strömendem Regen angefahren. Er
blieb eine Woche, wohnte im Bahnhofshotel, eine halbe Meile
entfernt, war aber den ganzen Tag bei ihr draußen. Als er abreiste,
versprach er, bald wiederzukommen, vielleicht schon in sechs
Wochen.
Jenny lag die Nächte hindurch bei brennender Lampe wach. Sie
wußte nur, sie konnte das nicht mehr ertragen. Es war zu furchtbar
gewesen.
Unerträglich war es — alles — von seinem ersten
teilnehmenden, besorgten Blick an, als er sie in dem neuen
dunkelblauen Hängerkleid erblickte, welches das Nähmädchen im
Dorf für sie angefertigt hatte. „Wie schön du bist,“ hatte er gesagt
und gemeint, sie gliche einer Madonna. Reizende Madonna! O ja. —
Sein vorsichtiger Arm um ihren Leib, seine langen behutsamen
Küsse auf ihre Stirn — ihr war, als sollte sie sterben vor Scham. Ja,
wie er sie gepeinigt hatte mit seiner liebevollen Besorgnis um ihre
Gesundheit, mit seinen Ermahnungen, für Bewegung zu sorgen. Als
einmal eine Pause zwischen den Regenschauern eintrat, hatte er sie
mit hinaus auf einen Spaziergang geschleppt, und sie mußte sich
um jeden Preis bei ihm einhängen und sich auf seinen Arm stützen.
Eines Abends besah er verstohlen ihre Handarbeit — er hatte sicher
erwartet, daß sie damit beschäftigt sei, Windeln zu säumen.
Es war ja keine böse Absicht von ihm. Aber darum war auch
keine Hoffnung vorhanden, daß es besser sein würde, wenn er
wiederkam, im Gegenteil. Aber sie konnte auch nicht mehr. —
Eines Tages bekam sie einen Brief von ihm, in dem er unter
anderem schrieb, daß sie auf jeden Fall einen Arzt zu Rate ziehen
müsse.
Am nämlichen Abend schrieb sie einen kurzen Brief an Gunnar
Heggen: sie erwarte im Februar ein Kind; ob er ihr die Adresse eines
stillen Ortes in Deutschland verschaffen könne, wo sie bleiben
könne, bis es überstanden sei.
Er antwortet umgehend:
Liebe Jenny!
Lieber Freund!
Ich habe einen Entschluß gefaßt, der Dir, wie ich fürchte,
wehe tun wird. Aber Du darfst mir nicht zürnen. Ich bin so
müde und nervös, weiß selbst, daß ich ungerecht und häßlich
gegen Dich war, als Du hier warst, und das möchte ich so
ungern. Daher will ich Dich nicht eher sehen, als bis alles
überstanden ist und ich wieder in normalem Zustande bin. Ich
reise morgen früh ins Ausland — meine Adresse gebe ich
vorläufig nicht an, Briefe kannst Du mir aber durch Frau
Franziska Ahlin, Varberg, Schweden, senden; ich schreibe
vorläufig über sie an Dich. Du darfst Dich meinetwegen nicht
ängstigen; ich bin frisch und es geht mir recht gut, aber,
Lieber, versuche nicht, bis auf weiteres anders mit mir in
Verbindung zu kommen, ich bitte Dich inständig. Und sei mir
nicht allzu böse, aber ich glaube, dieser Ausweg ist für uns
beide der beste. Versuche, um meinetwillen so wenig betrübt
und besorgt zu sein, wie es Dir möglich ist.
Deine
Jenny Winge.
Ich danke Dir für Deinen letzten Brief. Zu allererst muß ich
Dir sagen, da Du Dir scheinbar Vorwürfe machst in bezug auf
Dein Verhältnis zu mir; liebes kleines Mädchen, ich mache Dir
ja keine, und darum darfst Du es auch nicht. Du bist ja immer
nur gut und weich und liebevoll gegen Deinen Freund
gewesen. Nie werde ich Deine Zärtlichkeit und Deine Wärme
aus der kurzen Zeit, da Du mich liebtest, vergessen — Deine
süße Jungfräulichkeit und feine, sanfte Hingebung in den
Tagen unseres kurzen Glücks.
Unser Glück konnte nur kurz sein; das hätten wir beide
wissen müssen. Ich hätte es wissen m ü s s e n . Du hättest es
wohl wissen k ö n n e n , wenn Du nachgedacht hättest; aber
was denken zwei Menschen, die sich zu einander hingezogen
fühlen? Daß Du eines Tages aufhörtest, mich zu lieben —
glaubst Du, ich werfe Dir das vor? Wenn es mir auch das
bitterste Leid verursachte in meinem sonst nicht eben
glücklichen Leben — doppelt bitter für mich, da ich
gleichzeitig erfuhr, daß Du für unser Verhältnis nun durch dein
ganzes Leben büßen mußt.
Aber nun sehe ich aus Deinem Briefe, daß diese Folgen,
über die ich sicher viel verzweifelter war als Du, was Du auch
an Sorgen und körperlichen Leiden durchgemacht haben
magst, Dir dennoch eine tiefere Freude, ein größeres Glück
geschenkt haben, als es Dir sonst im Leben begegnet ist. Ich
sah, daß die Mutterfreude Dich ganz mit Frieden, Lebensmut
und Zufriedenheit erfüllte, so daß Du meinst, mit Deinem
Kinde im Arm genug Kraft zu besitzen, um alle
Schwierigkeiten, ökonomische wie soziale zu überwinden, die
die Zukunft einer jungen Frau in Deiner Lage bringen kann.
Daß Du dies schreibst, macht mich froher, als Du ahnen
kannst. Für mich ist dies wiederum ein Beweis für das Walten
jener ewigen Gerechtigkeit, an der ich ja nicht zweifle. Dir, die
Du einen Irrtum begingst, weil Dein Herz warm und zärtlich
war und nach Zärtlichkeit dürstete, wird gerade dieser Irrtum,
der Dir so verzweifelte Stunden gebracht hat, schließlich all
das bescheren, wonach Du so brennend verlangtest, besser,
schöner und reiner, als Du es je erträumt. Schon jetzt, da dein
Herz ganz von Liebe zu Deinem Kinde erfüllt ist und später in
noch höherem Maße, wenn der kleine Bursche heranwächst,
seine Mutter kennen lernt, sich an sie hängt und ihre Liebe
erwidern kann, stärker, tiefer und bewußter mit jedem Jahre,
das dahingeht.
Und mir, der ich Deine Liebe entgegennahm, obgleich ich
hätte wissen müssen, daß ein Liebesverhältnis zwischen uns
unmöglich und unnatürlich war — mir haben diese Monate
unerträgliches Leiden und Trauern gebracht — und einen
Verlust, Jenny, einen Verlust, wie Du ihn Dir nicht vorstellen
kannst, den Verlust Deiner Person, Deiner Jugend, Deiner
Schönheit, Deiner gesegneten Liebe. Jede kleinste
Erinnerung an diese Dinge war durch die Reue verbittert —
diese ständig nagende Frage, wie konnte ich sie es tun
lassen, wie konnte ich es annehmen, wie konnte ich an ein
Glück für mich mit ihr glauben? Ja, Jenny, ich habe daran
geglaubt, so wahnsinnig es auch klingt, weil ich mich bei Dir
so jung fühlte. Vergiß nicht, daß ich meiner eigenen Jugend
verlustig ging, und dies, als ich weit jünger war als Du jetzt
bist; der Jugend arbeitsfrohes Leben und frohes Liebesglück
durfte ich — durch eigene Schuld — nicht kennen lernen. Und
dies war die Strafe. Gespenstisch kehrte meine tote Jugend
zurück, als ich Dich gesehen — mein Herz fühlte sich nicht
älter als das Deine. Oh, Jenny, nichts auf der Welt ist
fürchterlicher, als wenn ein Mann alt und sein Herz jung
geblieben ist.
Du schreibst, Du sähest es gern, wenn ich später, sobald
der Knabe größer geworden ist, Dich besuchte, um mir unser
Kind anzusehen. Unser Kind — es ist ein so widersinniger
Gedanke. Weißt Du, woran ich dauernd denken muß? Kannst
Du Dich des alten Joseph entsinnen auf den italienischen
Altarbildern, der immer abseits oder im Hintergrunde zur Seite
steht, zärtlich und wehmütig das göttliche Kind und dessen
junge, herrliche Mutter betrachtend, diese beiden, die ganz
von einander in Anspruch genommen sind, und seine
Anwesenheit gar nicht beachten. Liebe Jenny, mißverstehe
mich nicht, ich weiß ja, daß das Kindchen, das jetzt in Deinem
Schoß liegt, auch mein Fleisch und Blut ist und doch — wenn
ich jetzt an Dich denke, die Mutter geworden ist, dann komme
ich mir wie der arme alte Joseph vor, der draußen steht.
Aber deshalb solltest Du ebensowenig Bedenken tragen,
den Namen als meine Gattin anzunehmen und den Schutz,
der für Dich und Dein Kind darin liegt, wie Maria, als sie sich
dem Joseph anvertraute. Eigentlich finde ich, es ist nicht ganz
richtig von Dir, dem Kinde den Vatersnamen zu rauben, auf
den es doch ein Anrecht hat — Du magst soviel
Selbstvertrauen haben wie Du willst. Selbstverständlich ist es,
daß Du im Falle einer solchen Ehe ebenso frei und
ungebunden bleibst wie sonst, und daß diese Ehe auch,
sobald du es wünschest, gesetzlich aufgehoben wird. Ich bitte
Dich inständig, Dir dies zu überlegen. Wir können uns im
Auslande trauen lassen, wenn Du es wünschest, und schon
einige Monate danach können Schritte zur Trennung getan
werden. Du brauchst nie wieder nach Norwegen
zurückzukehren, geschweige denn unter einem Dach mit mir
zu wohnen.
Von mir selbst ist nicht viel zu berichten. Ich habe zwei
kleine Zimmer hier oben auf dem Haegdehaug ganz in der
Nähe jenes Landhauses, in dem ich geboren bin und bis zu
meinem zehnten Jahre gelebt habe, als mein Vater im
Numetal Vogt wurde. Von meinem Fenster aus sehe ich die
Spitzen der beiden großen Kastanien an der Eingangstür
meines Vaterhauses. Sie haben sich nicht sonderlich
verändert. Hier oben beginnen die Abende bereits lang und
licht und lenzhaft zu werden, die Bäume zeichnen ihre
nackten braunen Kronen in den fahlgrünen Himmel, an dem
einzelne goldene Sterne durch die scharfe klare Luft funkeln.
Abend für Abend sitze ich hier an meinem Fenster und starre
in die Ferne, während mein ganzes Leben in Träumen und
Erinnerungen an mir vorüberzieht. Ach, Jenny, wie hatte ich
jemals vergessen können, daß ein ganzes Leben zwischen
Dir und mir lag, ein Leben, fast doppelt so lang wie das
Deine, ein Leben, von dem mehr als die Hälfte in
ununterbrochener Demütigung, Niederlage und Schmerz
dahingeschleppt worden ist. —
Daß Du ohne Zorn und Bitterkeit an mich denkst, ist mehr,
als ich erhofft und erwartet habe. Das Glück, das durch jede
Zeile Deines Briefes atmet, hat mir so unsagbar wohlgetan.
Gott segne und behüte mein Kind und Dich; alles Glück der
Welt wünsche ich auf Dich und das Kind herab. Ich habe Dich
so unsäglich lieb, Du kleine Jenny, die einst mein war.
Dein treuer
Gert Gram.
VII.
Jenny blieb bei Frau Schlessinger wohnen. Dort war es billig —
und sie wußte nicht, wohin mit sich.
Es lag Lenzeswehen in der Luft, über die gewaltige, offene
Himmelskuppel hin segelten schwere, vom Sonnenlicht verbrämte
Wolken, die wie Gold und Blut brannten und sich an den Abenden im
unruhigen Meer spiegelten, wenn sie draußen auf der Mole war. Die
trübseligen, dunklen Flächen im Lande wurden lichtgrün, die
Pappeln schimmerten braunrot von neuem Sproß, und dufteten lind
und weich. Am Eisenbahndamm wimmelte es von Veilchen und
kleinen weißen und gelben Blumen. Schließlich war die Ebene üppig
grün, es sprühte von Farben an den Wegrainen, schwefelgelbe Iris
und große weiße Doldenpflanzen spiegelten sich in den
Wasserlöchern der Torfmieten. Eines schönen Tages strömte süßer
Heuduft über Land, der sich in dem salzigen Algengeruch vom
Strande her mischte.
Das Badehotel wurde eröffnet und Sommergäste zogen in die
kleinen Häuser an der Mole. Es wimmelte von Kindern auf dem
weißen Sandstreifen. Sie kugelten sich im Sand und platschten
barfuß ins Wasser hinaus, Mütter, Kindermädchen und Ammen in
Spreewäldertracht saßen nähend im Grase und beaufsichtigten sie.
Die Badehäuschen waren ins Wasser gerollt worden, und kleine
deutsche Backfische schrien und juchten dort draußen. Luxussegler
legten an der Mole an; Besuch kam aus der Stadt, abends war Tanz
im Badehotel; die kleine Tannenplantage war voller Spaziergänger.
Hier hatte Jenny zu Beginn des Frühlings in dem struppigen Gras
gelegen, dem Wellenschlag und dem Sausen des Windes in den
zerzausten Kronen lauschend.
Diese oder jene der Damen sandte ihr einen interessierten oder
teilnehmenden Blick nach, wenn sie den Weg am Badestrand
entlang spazierte, mit ihrem schwarzweißen Sommerkleide angetan.
Die Badegäste im Ort hatten natürlich erfahren, daß sie eine junge
Norwegerin war, die ein Kind bekommen hatte, über dessen Tod sie
so furchtbar trauerte. Einige waren auch darunter, die es mehr
rührend als skandalös fanden.
Im übrigen wanderte sie meist landeinwärts; dorthin kamen
niemals Sommergäste. Ganz selten ging sie bis hinauf zur Kirche
und zum Kirchhof, wo der Knabe lag. Sie saß dann und starrte auf
das Grab, das sie nicht hatte herrichten lassen. Sie legte dann einige
wilde Blumen nieder, die sie unterwegs gepflückt hatte, aber ihre
Phantasie weigerte sich, den kleinen, grauen Erdhügel, auf welchem
Unkraut und Gräser in die Höhe schossen, mit ihrem Bübchen in
Verbindung zu bringen.
An den Abenden saß sie in ihrem Zimmer mit einer Handarbeit,
die sie nicht anrührte, und starrte in die Lampe. Sie dachte immer an
das Gleiche, rief die Tage wieder zurück, als sie ihren Jungen
besessen hatte, die erste Zeit, das matte, friedliche Glück, während
sie lag und genas, später, wenn sie aufrecht im Bett saß und Frau
Schlessinger ihr das Baden und Wickeln, das An- und Auskleiden
des Kindes zeigte, dann, als sie zusammen nach Warnemünde
reisten, um feinen Stoff, Spitzen und Band zu kaufen, als sie
heimkehrte, zuschnitt, nähte, zeichnete und stickte — ihr Junge
sollte feine Sachen haben statt des schlechten fertiggekauften
Zeuges, das sie aus Berlin bestellt hatte. Eine drollige Gartenspritze
hatte sie gekauft mit Abziehbildern auf dem grünbemalten Blech: ein
Löwe und ein Tiger standen zwischen Palmen an einem
himmelblauen Meer und betrachteten entsetzt die deutschen
Panzerungetüme, die den afrikanischen Besitzungen des Reichs
zudampften. Sie fand das Ding so lustig — Bübchen sollte es zum
Spielen haben, wenn er einmal groß genug geworden war. Erst
mußte er ja Mutters Brust finden, an der er sich jetzt nur blind
festsog — und seine eigenen kleinen Finger, die er nicht
voneinander bekommen konnte, sobald er sie ineinander verfilzt
hatte — bald würde er die Mutter kennen, nach der Lampe blinzeln
und nach Mutters Uhr, die sie vor ihm schaukeln ließ — da war so
viel, was Bübchen lernen mußte.