(Download PDF) Biology Exploring The Diversity of Life 4Th Canadian Edition Edition Peter Russell Full Chapter PDF

Biology: Exploring the Diversity of Life
4th Canadian Edition Edition Peter

Russell
Visit to download the full and correct content document:
https://ebookmass.com/product/biology-exploring-the-diversity-of-life-4th-canadian-ed
ition-edition-peter-russell/
More products digital (pdf, epub, mobi) instant
download maybe you interests ...
Biology: Exploring the Diversity of Life, Fifth

Canadian Edition Shelby
https://ebookmass.com/product/biology-exploring-the-diversity-of-
life-fifth-canadian-edition-shelby/
Biology : the unity and diversity of life 15th Edition

Cecie Starr
https://ebookmass.com/product/biology-the-unity-and-diversity-of-
life-15th-edition-cecie-starr/
(eBook PDF) Campbell Biology, Third Canadian Edition
https://ebookmass.com/product/ebook-pdf-campbell-biology-third-
canadian-edition/
The Diversity of Fishes: Biology, Evolution and

Ecology, 3rd Edition Douglas E. Facey
https://ebookmass.com/product/the-diversity-of-fishes-biology-
evolution-and-ecology-3rd-edition-douglas-e-facey/
Campbell Biology, Third Canadian Edition (3rd Edition)
Urry & Wasserman
https://ebookmass.com/product/campbell-biology-third-canadian-
edition-3rd-edition-urry-wasserman/
Biology 12th Edition Peter H. Raven
https://ebookmass.com/product/biology-12th-edition-peter-h-raven/
Marine Biology 10th Edition Peter Castro
https://ebookmass.com/product/marine-biology-10th-edition-peter-
castro/
Marine Biology 12th Edition Peter Castro
https://ebookmass.com/product/marine-biology-12th-edition-peter-
castro/
Principles of Biology 4th Edition Robert J. Brooker
https://ebookmass.com/product/principles-of-biology-4th-edition-
robert-j-brooker/
RUSSELL FOURTH CANADIAN EDITION
HERTZ
MCMILLAN
EXPLORING THE DIVERSITY OF LIFE

FENTON
BIOLOGY
MAXWELL
HAFFIE
MILSOM
NICKLE
ELLIS
BIOLOGY
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
9780176718886_CVR_hr.indd 1 12/21/17 12:50 PM

bowdenimages/Getty Images
MindTap empowers students.
Personalized content in an easy-to-use interface
helps you achieve better grades.
The new MindTap Mobile App allows The MindTap Reader lets you highlight
for learning anytime, anywhere with and take notes online, right within the
flashcards, quizzes and notifications. pages, and easily reference them later.
nelson.com/mindtap
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.
FOURTH CANADIAN EDITION
BIOLOGY
Peter J. Russell M. Brock Fenton
Western University
Paul E. Hertz
Denis Maxwell
Beverly McMillan
Western University
Tom Haffie
Western University
Bill Milsom
University of British Columbia
Todd Nickle
Mount Royal University
Shona Ellis
With contributions by Ivona Mladenovic, Simon Fraser University
Biology, Fourth Canadian Edition
by Peter J. Russell, Paul E. Hertz,

Beverly McMillan, M. Brock Fenton,
Denis Maxwell, Tom Haffie, Bill
Milsom, Todd Nickle, Shona Ellis
VP, Product Solutions, K–20: Copy Editor: Cover Image:

Claudine O’Donnell Frances Robinson © Seth Casteel
Senior Publisher, Digital and Print Proofreader: Art Coordinator:
Content: MPS Limited Suzanne Peden
Paul Fam
Indexer: Managing Designer:
Marketing Manager: MPS Limited Courtney Hellam
Tia Nguyen
Design Director: Illustrator(s):
Content Manager: Ken Phipps Articulate Graphics, Steve Corrigan,
Toni Chahley Crowle Art Group, Patrick Gnan,
Higher Education Design Project Dave McKay, MPS Limited, Allan
Photo and Permissions Researcher: Manager: Moon, Ann Sanderson, Ralph Voltz
Kristiina Paul Pamela Johnston
Compositor:
Senior Production Project Manager: Interior Design Modifications: MPS Limited
Imoinda Romain Ken Cadinouche
Production Service: Cover Design:
MPS Limited Courtney Hellam
COPYRIGHT © 2019, 2016 by Nelson ALL RIGHTS RESERVED. No part of Library and Archives Canada
Education Ltd. this work covered by the copyright Cataloguing in Publication
herein may be reproduced,
Adapted from Biology, Fourth Russell, Peter J., author
transcribed, or used in any form
Edition, by Peter J. Russell, Paul Biology : exploring the diversity
or by any means—graphic,
E. Hertz, and Beverly McMillan, of life / Peter J. Russell, Paul E.
electronic, or mechanical, including
published by Cengage Learning. Hertz, Beverly McMillan, M. Brock
photocopying, recording, taping,
Copyright ©2017 by Cengage Fenton, University of Western
Web distribution, or information
Learning. Ontario, Denis Maxwell, University
storage and retrieval systems—
without the written permission of of Western Ontario, Tom Haffie,
Printed and bound in Canada
the publisher. University of Western Ontario,
1 2 3 4 21 20 19 18
Bill Milsom, University of British
For more information contact For permission to use material Columbia, Todd Nickle, Mount Royal
Nelson Education Ltd., from this text or product, University, Shona Ellis, University of
1120 Birchmount Road, Toronto, submit all requests online at British Columbia ; with contributions
Ontario, M1K 5G4. Or you can visit cengage.com/permissions. by Ivona Mladenovic, Simon Fraser
our Internet site at nelson.com Further questions about University. — Fourth Canadian edition.
permissions can be emailed to
Includes index.
Cognero and Full-Circle Assessment permissionrequest@cengage.com
Issued also in 3 volumes.
are registered trademarks of
Every effort has been made to Issued in print and electronic
Madeira Station LLC.
trace ownership of all copyrighted formats.
material and to secure permission ISBN 978-0-17-671888-6 (hardcover).—
from copyright holders. In the event ISBN 978-0-17-682709-0 (PDF)
of any question arising as to the use 1. Biology—Textbooks.
of any material, we will be pleased 2. Textbooks. I. Title.
to make the necessary corrections in
future printings. QH308.2.R88 2018 570 C2017-904622-5
C2017-904623-3
ISBN-13: 978-0-17-671888-6
ISBN-10: 0-17-671888-5
For, and because of,
our generations of students.
M.B. Fenton
About the Canadian Authors
M. B. (Brock) Fenton received his Ph.D. from the of Biological Societies; Honorary Life Membership, Science
University of Toronto in 1969. Since then, he has been a North, Sudbury, Ontario; Canadian Council of University
faculty member in biology at Carleton University, then at Biology Chairs Distinguished Canadian Biologist Award;
York University, and then at Western University. In addi- The McNeil Medal for the Public Awareness of Science
tion to teaching parts of first-year biology, he has also of the Royal Society of Canada; and the Sir Sandford
taught vertebrate biology, animal biology, and conserva- Fleming Medal for public awareness of Science, the Royal
Canadian Institute). He also received the C. Hart Merriam
M. B. Fenton
tion biology, as well as field courses in the biology and

behaviour of bats. He has received awards for his teaching Award from the American Society of Mammalogists for
(Carleton University Faculty of Science Teaching Award; excellence in scientific research. Bats and their biology,
Ontario Confederation of University Faculty Associations behaviour, evolution, and echolocation are the topics of his
Teaching Award; and a 3M Teaching Fellowship, Society research, which has been funded by the Natural Sciences
for Teaching and Learning in Higher Education) in addi- and Engineering Research Council of Canada (NSERC).
tion to recognition of his work on public awareness of sci- In November 2014, Brock was inducted as a Fellow of the
ence (Gordin Kaplan Award from the Canadian Federation Royal Society of Canada.
Denis Maxwell received his Ph.D. from the University University of New Brunswick in 2000, he moved in 2003
of Western Ontario in 1995. His thesis, under the supervi- to the Department of Biology at Western University. Denis
sion of Norm Hüner, focused on photosynthetic acclima- served as Associate Chair for Undergraduate Education
tion in green algae. Following his doctorate, he undertook for the Department of Biology from 2009 to 2016.
postdoctoral training at the Department of Energy Plant Currently, he is Assistant Dean for the Faculty of Science,
Research Laboratory at Michigan State University, where with a portfolio that includes Recruitment and First-Year
he studied the function of the mitochondrial alterna- Studies and outreach. He has taught first-year Biology to
Alan Noon
tive oxidase. After taking up a faculty position at the over 15 000 students, most of the time with Tom Haffie.
Tom Haffie is a graduate of the University of Guelph to development. He is the founding coordinator of the
and the University of Saskatchewan in the area of biennial Western Conference on Science Education.
microbial genetics. Tom has devoted his 33-year career He holds a University Students’ Council Award for
at Western University to teaching large biology classes Excellence in Teaching, a UWO Edward G. Pleva Award
for Excellence in Teaching, a UWO Fellowship in
Tom Haffie
in lecture, laboratory, and tutorial settings. He led the

development of the innovative core laboratory course Teaching Innovation, a Province of Ontario Award for
in the Biology program; he was an early adopter of Leadership in Faculty Teaching (LIFT), and a Canadian
computer animation in lectures; and, most recently, 3M National Teaching Fellowship for excellence in
has overseen a deep blended redesign of introductory teaching. Tom is currently a Teaching Fellow for Science
biology informed by a students-as-partners approach at Western University.
iv NEL
Bill Milsom (Ph.D., University of British Columbia) is of biodiversity and the physiological costs of habitat selec-
a professor in the Department of Zoology at the University tion. His research has been funded by NSERC, and he has
of British Columbia, where he has taught a variety of received several academic awards and distinctions, includ-
courses, including first-year biology, for almost 40 years. ing the Fry Medal of the Canadian Society of Zoologists,
Bruce Moffat
His research interests include the evolutionary origins of the August Krogh Distinguished Lectorship Award of the
respiratory processes and the adaptive changes in these American Physiological Society, the Bidder Lecture of the
processes that allow animals to exploit diverse environ- Society for Experimental Biology, and the Izaak Walton
ments. He examines respiratory and cardiovascular adapta- Killam Award for Excellence in Mentoring. He has served
tions in vertebrate animals in rest, sleep, exercise, altitude, as the President of the Canadian Society of Zoologists and
dormancy, hibernation, diving, and so on. This research as President of the International Congress of Comparative
contributes to our understanding of the mechanistic basis Physiology and Biochemistry.
Todd Nickle received his Ph.D. from Oklahoma State relate to other disciplines, which inspired him to coauthor
University in 1998, and has been teaching biology at Mount a handbook for first-year science students (Science3).
Royal University ever since. He advocates Active Learning: His interest in promoting best teaching practices among
students come to class prepared to work with material educators had him confirm the Alberta Introductory
rather than just hear about it. Student preparation involves Biology Association as an official Society of Alberta; Todd
reading the text and applying the concepts to online exer- is currently President. His work put him in the first cohort
Penny Nickle
cises, the results of which inform what the next lecture of Full Professors at Mount Royal University in 2012, gar-
will be about. Class time focusses on exploring connec- nered the 2015 ACIFA Innovation in Teaching Award, and
tions between concepts and ideas in biology and how they the Distinguished Faculty Award from MRU in 2016.
Shona Ellis (M.Sc., University of British Columbia) botany courses that have included nonvascular and vas-
is a professor of teaching in the Botany Department and cular plants, economic botany, bryology, and plant sys-
Associate Head of Biology at the University of British tematics, as well as introductory biology. Professor Ellis
Columbia. She developed a keen interest in forests and has taught in a number of settings: large and small lec-
the ocean growing up on the central coast of British tures, laboratories, and fieldtrips. While she feels the best
Columbia. As an undergraduate, Professor Ellis pur- classroom is outdoors, she integrates online technologies
sued her interests in botany and entomology. Her M.Sc. into all her courses; she is an early adopter of online
research incorporated tissue culture, phytochemistry, teaching and learning resources. Professor Ellis has
and plant anatomy. As a teaching assistant, she realized received two Killam Teaching Awards and the Charles
Andy Cotton
a passion for teaching and joined the teaching faculty at Edwin Bessey Teaching Award from the Botanical
the University of British Columbia in 1998. She teaches Society of America.
NEL A B O U T T H E C A N A D I A N AU T H O R S
v
About the U.S. Authors
Peter J. Russell received a B.Sc. in Biology from expression of the genes of the virus; yeast was used as the
the University of Sussex, England in 1968 and a Ph.D. model host. His research has been funded by agencies
in Genetics from Cornell University in 1972. He has including the National Institutes of Health, the National
been a member of the Biology Faculty of Reed College Science Foundation, the American Cancer Society,
since 1972, and is currently a Professor of Biology, the Department of Defense, the Medical Research
Emeritus. Peter taught a section of the introductory Foundation of Oregon, and the Murdoch Foundation.
biology course, a genetics course, and a research litera- He has published his research results in a variety of
ture course on molecular virology. In 1987 he received journals, including Genetics, Journal of Bacteriology,
the Burlington Northern Faculty Achievement Award Molecular and General Genetics, Nucleic Acids Research,
from Reed College in recognition of his excellence in Plasmid, and Molecular and Cellular Biology. Peter has
teaching. Since 1986, he has been the author of a success- a long history of encouraging faculty research involv-
ful genetics textbook: current editions are iGenetics: A ing undergraduates, including cofounding the biology
Molecular Approach, iGenetics: A Mendelian Approach, division of the Council on Undergraduate Research, in
and Essential iGenetics. Peter’s research was in the area Washington, D.C. in 1985. He was Principal Investigator/
of molecular genetics, with a specific interest in char- Program Director of a National Science Foundation
acterizing the role of host genes in the replication of Award for the Integration of Research and Education
the RNA genome of a pathogenic plant virus, and the (NSF-AIRE) to Reed College, 1998–2002.
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
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
UNIT TWO GENES 149 THE CHEMICAL AND PHYSICAL FOUNDATIONS

7 Cell Cycles 149 OF BIOLOGY (PURPLE PAGES) F-1
8 Genetic Recombination 173
9 The Chromosomal Basis of Mendelian Inheritance 199 VOLUME 3: SYSTEMS AND PROCESSES 881
10 Genetic Linkage, Sex Linkage, and Other Non-Mendelian
Inheritance Mechanisms 225 UNIT SEVEN SYSTEMS AND PROCESSES -
PLANTS 885
UNIT THREE DNA AND GENE EXPRESSION 251
33 Organization of the Plant Body 885
11 DNA Structure, Replication, and Repair 251 34 Transport in Plants 913
12 Gene Structure, Expression, and Mutation 277 35 Reproduction and Development in Flowering Plants 933
13 Regulation of Gene Expression 307 36 Plant Nutrition 955
14 DNA Technologies 333 37 Plant Signals and Responses to the Environment 977
15 Genomes 361
UNIT EIGHT SYSTEMS AND PROCESSES -
VOLUME 2: EVOLUTION, ECOLOGY, AND THE ANIMALS 1007
DIVERSITY OF LIFE 389 38 Introduction to Animal Organization and Physiology 1007
39 Animal Nutrition 1029
UNIT FOUR EVOLUTION AND CLASSIFICATION 393 40 Gas Exchange: The Respiratory System 1057
16 Evolution: The Development of the Theory 393 41 Internal Transport: The Circulatory System 1079
17 Microevolution: Changes within Populations 411 42 Regulation of the Internal Environment: Water, Solutes,
18 Speciation and Macroevolution 433 and Temperature 1103
19 Systematics and Phylogenetics: Revealing the Tree of 43 Control of Animal Processes: Endocrine Control 1135
Life 453 44 Animal Reproduction 1161
20 Humans and Evolution 477 45 Control of Animal Processes: Neural Control 1199
46 Muscles, Skeletons, and Body Movements 1255
UNIT FIVE THE DIVERSITY OF LIFE 493
21 Defining Life and Its Origins 493 APPENDIX A: ANSWERS TO SELF-TEST
22 Viruses, Viroids, and Prions: Infectious Biological QUESTIONS A-1
Particles 521 GLOSSARY G-1
23 Bacteria and Archaea 537 INDEX I-1
24 Protists 559
NEL vii
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
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 The Cell: An Overview 25 6 Photosynthesis 125
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
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
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
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
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
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
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
ix
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
Figure 18.16 Observational Research Chromosomal Similarities
15 Genomes 361 and Differences among Humans and the Great Apes 447
15.1 Genomics: An Overview 362 SUMMARY ILLUSTRATION FOR CHAPTER 18 448

15.2 Genome Sequencing 363
19 Systematics and Phylogenetics: Revealing the Tree of
Figure 15.1 Research Method Sanger Sequencing 364
Life 453
Figure 15.2 Research Method Pyrosequencing 368 19.1 Nomenclature and Classification 454
15.3 Annotation Identifies Genes 371 19.2 Phylogenetic Trees 456

19.3 Sources of Data for Phylogenetic Analyses 458
Figure 15.8 Research Method DNA Microarray Analysis of Gene
Expression Levels 376 19.4 Traditional Classification and Paraphyletic Groups 461
19.5 The Cladistic Revolution 462
15.4 Comparative Genomics Can Reveal How Genes and Genomes
Evolved 377 Figure 19.11 Research Method Using Cladistics to Construct a
Phylogenetic Tree 465
Figure 19.13 Research Method Using Genetic Distances to
VOLUME 2: EVOLUTION, ECOLOGY, AND THE Construct a Phylogenetic Tree 468
DIVERSITY OF LIFE 389 19.6 Phylogenetic Trees as Research Tools 469
UNIT FOUR EVOLUTION AND CLASSIFICATION 393 19.7 Molecular Phylogenetic Analyses 469
16 Evolution: The Development of the Theory 393
16.1 The Recognition of Change 394 20 Humans and Evolution 477
16.2 Natural Selection 397 Research in Biology The Cast of Characters: Fossil Hominins 478
16.3 Evolutionary Biology since Darwin 400
20.1 The Fossil Record of Hominins 480
16.4 Evolution Is the Core Theory of Modern Biology but Is Plagued
by Misconceptions 403 20.2 Morphology and Bipedalism 481
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
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
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
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
27.11 Jawed Fishes: Jaws Meant New Feeding Opportunities 690
24 Protists 559 27.12 Early Tetrapods and Modern Amphibians 696
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
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
NEL C O N T E N T S
xi
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
28.12 Taking Action 740 31.6 Scale, Ecosystems, Species 833
SUMMARY ILLUSTRATION FOR CHAPTER 28 742 31.7 Three Sample Ecosystems 834
UNIT SIX ECOLOGY AND BEHAVIOUR 747 32 Animal Behaviour 845

32.1 Genes, Environment, and Behaviour 846
29 Population Ecology 747
32.2 Instinct 848
29.1 Introduction 749
Figure 32.5 Experimental Research The Role of Sign Stimuli in
29.2 Population Characteristics 749
Parent–Offspring Interactions 849
29.3 Demography 752
32.3 Knockouts: Genes and Behaviour 850
29.4 Evolution of Life Histories 753
32.4 Learning 850
29.5 Models of Population Growth 756
32.5 Neurophysiology and Behaviour 852
29.6 Population Regulation 760
32.6 Hormones and Behaviour 853
Figure 29.17 Experimental Research Evaluating Density-Dependent
Interactions between Species 762 32.7 Neural Anatomy and Behaviour 855
32.8 Communication 856
29.7 Human Administered Population Control 765
32.9 Language: Syntax and Symbols 861
29.8 Human Population Growth 766
32.10 Space 862
29.9 The Future: Where Are We Going? 770
32.11 Home Range and Territory 862
29.10 The Pill 770
32.12 Migration 863
Figure 32.32 Experimental Research Experimental Analysis of the
30 Species Interactions and Community Ecology 777
Indigo Bunting’s Star Compass 867
30.1 Introduction 778 32.13 Mates as Resources 868

30.2 Symbiosis: Close Associations 779 32.14 Sexual Selection 869
30.3 Energy Intake 782 32.15 Social Behaviour 870
30.4 Defence 784 32.16 Kin Selection and Altruism 872
30.5 Competition 788 32.17 Eusocial Animals 874
Figure 30.15 Experimental Research Demonstration of Competition 32.18 Human Social Behaviour 876
between Two Species of Barnacles 789 SUMMARY ILLUSTRATION FOR CHAPTER 32 878
Figure 30.16 Experimental Research Gause’s Experiments on

THE CHEMICAL AND PHYSICAL FOUNDATIONS
Interspecific Competition in Paramecium 790
OF BIOLOGY (PURPLE PAGES) F-1
Figure 30.19 Experimental Research The Complex Effects of a What Are the Purple Pages? F-1
Herbivorous Snail on Algal Species Richness 792 Emergent Properties F-1
30.6 The Nature of Ecological Communities 793 The Scientific Basis of Biology F-2
30.7 Community Characteristics 793 Measurement and Scale F-5
30.8 Effects of Population Interactions on Community The Organization of Matter F-7
Structure 798 Atoms Interact to Produce New Properties F-11
30.9 Succession 799 Chemical Bonds F-11
30.10 Variations in Species Richness among Communities 804 Water F-15
SUMMARY ILLUSTRATION FOR CHAPTER 30 808 Carbon Compounds F-21
xii C O N T E N T S NEL
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
History of Earth F-42 37.2 Plant Chemical Defences 989

37.3 Plant Movements 993
VOLUME 3: SYSTEMS AND PROCESSES 881 37.4 Plant Biological Clocks 997
UNIT SEVEN SYSTEMS AND PROCESSES: PLANTS 885
33 Organization of the Plant Body 885

UNIT EIGHT SYSTEMS AND
33.1 Plant Structure and Growth: An Overview 886
PROCESSES: ANIMALS 1007
33.2 The Three Plant Tissue Systems 890
Figure 33.9 Experimental Research Networking the Secondary Cell 38 Introduction to Animal Organization and Physiology 1007
Wall 893 38.1 Organization of the Animal Body 1008
38.2 Animal Tissues 1009
33.3 Primary Shoot Systems 895
38.3 Coordination of Tissues in Organs and Organ Systems 1018
33.4 Root Systems 900
38.4 Homeostasis 1018
33.5 Secondary Growth 903
Figure 38.12 Experimental Research Demonstration of the Use of
the Bill for Thermoregulation in Birds 1021
34 Transport in Plants 913 SUMMARY ILLUSTRATION FOR CHAPTER 38 1024

34.1 Principles of Water and Solute Movement in Plants 914
34.2 Uptake and Transport of Water and Solutes by Roots 917 39 Animal Nutrition 1029
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
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 Plant Nutrition 955 40.5 Transport of Gases in Blood 1069
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
36.2 Soil 960 SUMMARY ILLUSTRATION FOR CHAPTER 40 1074
36.3 Root Adaptations for Obtaining and Absorbing Nutrients 964

41 Internal Transport: The Circulatory System 1079
41.1 Animal Circulatory Systems: An Introduction 1080
37 Plant Signals and Responses to the Environment 977 41.2 Blood and Its Components 1084
37.1 Introduction to Plant Hormones 979 41.3 The Heart 1087
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
xiii
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
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
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
44 Animal Reproduction 1161 Appendix A: Answers to Self-Test Questions A-1

44.1 The Drive to Reproduce 1162 Glossary G-1
44.2 Asexual and Sexual Reproduction 1162 Index I-1
xiv C O N T E N T S NEL
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
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
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-
●●
tional online exercise to further develop student quantitative

standing as they acquire it. Each major section covers one
analysis and mathematical reasoning skills.
broad topic.
The Design an Experiment feature is delivered online as a
Study Break questions follow every major section. These
●●
●●
guided learning activity that takes the student through the
reading comprehension questions encourage students to
process of designing an experiment.
pause at the end of a section and review what they have
Conceptual Learning Activities are repeatable in alternate
learned before going on to the next topic within the chapter.
●●
versions to help students learn the material.

If a student isn’t able to answer a study break question, they
can immediately revisit the previous section to solidify their The Instructor Resource Center provides everything you
understanding. We feel that this is a better learning tool than need for your course in one place. This collection of lecture and
directly providing the answers to these questions. If the class tools is available online for instructors only via www
answer does not come easily, then rereading the material .nelson.com/instructor. There you can access and download
associated with the answer is as important as seeing the PowerPoint presentations, images, the Instructor’s Manual, the
answer itself. Test Bank, videos, animations, and more.
●● Self-Test Questions are found at the end of each chapter. To maximize the chances of producing a useful text that
These chapter review questions are organized according draws in students (and instructors), we sought the advice of col-
to Bloom’s taxonomy into three sections: Recall/Under- leagues who teach biology (members of the MindTap Advisory
stand, Apply/Analyze, and Create/Evaluate. This struc- Board). We also asked students (members of the Student Advi-
ture allows students to review the material in a sequence sory Boards) for their advice and comments. These groups eval-
that moves from the basic knowledge of factual material, uated the effectiveness of important visuals in the textbook,
to more challenging and sophisticated applications evaluated draft chapters, and provided valuable feedback on the
of that knowledge, to novel situations. Answers to the MindTap, but any mistakes are ours.
Self-Test Questions are found in an appendix at the back In summary, we have applied our collective experience as
of the book. teachers, researchers, and writers to create a readable and under-
●● The Chemical and Physical Foundations of Biology, also standable foundation for students who may choose to enrol in
known as The Purple Pages, keep background information more advanced biology courses in the future. Where appro-
out of the main text, allowing students to focus on the bigger priate, we provide straightforward explanations of fundamental
picture. concepts from the evolutionary perspective that binds together
●● Unit 5: The Diversity of Life, also known as The Green Pages, all the biological sciences. Recognizing that students in an intro-
contains readily identifiable chapters that introduce the tre- ductory biology course face a potentially daunting quantity of
mendous variability among living organisms. ideas and information, we strive to provide an appropriate
NEL P R E FA C E
xvii
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
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?”
A streamlined textbook … Emphasizing connections …

In response to feedback from students and instructors across the We recognize that part of the challenge of an introduction to
country, we have made some important changes that have biology course lays in covering a large breadth of knowledge
resulted in a briefer edition. while making meaningful connections across topics, concepts,
and the discipline as a whole. In Biology: Exploring the Diversity
of Life, every chapter begins with a Chapter Roadmap that pro-
Organizational Changes
vides students with a visual overview of the chapter contents,
By combining and reorganizing information, we have reduced while making connections between parts of the chapter and
the number of chapters in the book from 52 to 46. The material other chapters in the book. Within chapters, students will find
on protostomes and deuterostomes has been combined into a cross-references and connections to other chapters where a con-
single super chapter on animals. Using the latest research as our cept is explored further or from a different perspective. Further-
guide, Chapter 27: Animals captures the excitement of how new more, every chapter concludes with a Summary Illustration, a
developments in molecular phylogenetic techniques have two-page spread that synthesizes, integrates, and illustrates con-
resulted in many taxonomic reclassifications as well as changes nections between important concepts covered in the chapter.
to phylogenies. This chapter features a unique research box on
the Tully Monster as a case in point. Clear and thoughtful visuals …
We have also streamlined our coverage of systems and pro-
cesses in animals by combining reproduction and development to Each of the figures in the new edition delivers a clear and
create Chapter 44: Animal Reproduction. The chapters on neural thoughtful message that is tied directly to the discussion it
control and neural integration have been combined to create accompanies. The new edition contains over 200 new and 55
Chapter 45: Control of Animal Processes: Neural Control. revised figures. We have further enhanced this connection
We have also rewritten former Chapter 33: Putting Selec- through the refinement and integration of research figures.
tion to Work and Chapter 52: Conservation and Evolutionary Experimental Research, Observational Research, and Research
Physiology into a collection of case studies and placed them on Methods are further highlighted in a vivid new design, drawing
the MindTap for the book. attention to how biologists formulate and test specific hypoth-
eses by gathering and interpreting data.
Streamlined Pedagogy and Prose
Extensively Revised Unit Four: Evolution and
Our revisions to the fourth Canadian edition were also informed
Classification
by a desire to reduce redundancy across the book, including only
essential, testable information. As a result, students and instruc- The fundamental concepts of evolution are essential for students
tors will find an efficient use of prose across the new edition, as to grasp as they explain the diversity of living organisms as well
well as extensive use of cross-references to other chapters, where as the commonalities that organisms possess. That said, many
necessary. The feature boxes “Molecule behind Biology,” “People first-year students come to university with a poor understanding
NEL xix
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
●●
NEW material on DNA packaging

between evolution and genetics. Chapter 16: Evolution by
●●
NEW figure clarifying replicated versus unreplicated

Natural Selection now includes a section and Research Figure
●●
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
Chapter 11: DNA Structure, Replication, and

Chapter 2: The Cell: An Overview Repair
●● NEW Research Figure about cell fractionalization ●● NEW Canadian Why It Matters about woolly mammoths in
Canada
Chapter 3: Energy and Enzymes ●● Highlighted mechanisms of repair of DNA damage
More precise description of fundamentals of

Chapter 12: Gene Structure, Expression, and
●●
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
Chapter 5: Cellular Respiration Chapter 13: Regulation of Gene Expression

●● Clarified section on chemical basis of cellular respiration to ●● NEW Why It Matters featuring epigenetic regulation of hon-
include stronger connections with Chapter 3 eybee castes
xx N E W TO T H I S E D I T I O N NEL
●● Updated material on lncRNA ●● The concepts of homology and convergent evolution are
●● Updated material on cancer genetics more clearly presented.
Chapter 14: DNA Technologies Chapter 20: Humans and Evolution

●● NEW section on CRISPR and qPCR ●● Unchanged from previous edition
●● Clarified and expanded Health Canada position on geneti-
cally modified foods
●● Added material on knockout mouse protocol
Chapter 21: Defining Life and Its Origins
●● In this edition, this chapter includes a section on the fossil
Chapter 15: Genomics record.
●● Expanded section discussing possible energy sources for
●● Fully updated early life
●● NEW section on comparative genomics ●● More in-depth discussion on LUCA
●● Linked advances in sequencing technologies from Sanger to
early next-gen methods to DNA replication outlined in
Chapter 11 Chapter 22: Viruses, Viroids, and Prions:
●● Enhanced explanation of principles behind BLAST Infectious Biological Particles
●● NEW Canadian example about tracking viral disease
Chapter 16: Evolution: The Development of the ●● NEW Research Figure: A New Discovery for Hepatitis C
Theory Therapy
●● Completely rewritten with a greater focus on fundamental
concepts of evolution and less emphasis on historical Chapter 23: Bacteria and Archaea
development
●● Phylogenetic tree updated to reflect latest research
●● NEW Why It Matters about antibiotic resistance
●● NEW Research Figure: Genetic Recombination in Bacteria
●● Section and Experimental Research Figure focused on
NEW discussion of a recent finding of a group within
experimental evolution in E. coli
●●
Archaea (Lokiarchaeota) that has a number of genes in

●● Concluding figure explains the major misconceptions stu-
common with eukaryotes
dents have with evolution and natural selection
Chapter 17: Microevolution: Changes within Chapter 24: Protists

Populations ●● Phylogenetic tree updated to reflect latest research
Incorporated recent research on Diplonemids that had been
Completely rewritten with greater clarity on the processes
●●
●●
previously known from only a single environmental gene
that cause evolutionary change
from marine planktonic samples
●● Stronger connections with genetics by fully explaining terms
NEW Research Figure: Isolation and Identification of Marine
such as allele, gene, gene pool, locus
●●
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
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 discussion of the impact of wolf predation on popula-

tions of caribou Environment
●● Updated section on Darwin’s experiments using light and
Chapter 29: Population Ecology oat coleoptiles
●● NEW Why It Matters about the other malaria
●● NEW Research Figure: Evaluating Density-Dependent Chapter 38: Introduction to Animal
Interactions between Species Organization and Physiology
●● NEW section on Human Administered Population Control
●● NEW Research Figure: Demonstration of the Use of the Bill
for Thermoregulation in Birds
Chapter 30: Species Interactions and
Community Ecology
Chapter 39: Animal Nutrition
●● NEW Why It Matters about oxpeckers and their hosts
●● NEW coverage of blood feeders
●● NEW Research Figure: Association of Bacterial Populations
●● NEW material on venoms, how animals use them and how in the Gut Microbiome with Obesity in Humans
they work
●● NEW definition of essential nutrients, malnutrition, and
undernutrition
NEW figure illustrating intracellular digestion
Chapter 31: Ecosystems
●●
●● NEW Why It Matters about cave ecosystems Chapter 40: Gas Exchange: The Respiratory
Updated discussion of mass mortality
System
●●
●● Enhanced discussion of urban ecosystems

●● NEW Why It Matters, featuring a discussion of adaptations
Chapter 32: Animal Behaviour that allow animals to live in oxygen-limited environments
(burrows, during diving, at altitude)
●● NEW Why It Matters about bird migration ●● NEW Research Figure: Demonstration of a Molecular Basis
●● NEW discussion about changing behaviour, featuring for High-Altitude Adaptation in Deer Mice
moose, salt, and cars
NEW coverage of echolocation
Chapter 41: Internal Transport: The Circulatory
●●
●● NEW material on the evolution of human language

System
Chapter 33: Organization of the Plant Body ●● NEW Why It Matters, featuring the effects of animal body
●● Incorporation of current research that demonstrates, with size on resting heart rate (but not longevity)
the discovery of new transcription factors, that there is an
●● NEW Research Figure: Demonstration of a Vasodilatory
unexpectedly complex regulatory network governing sec- Signalling Molecule
ondary wall development
●● NEW Research Figure: Networking the Secondary Cell Wall Chapter 42: Regulation of the Internal
Environment: Water, Solutes, and Temperature
Chapter 34: Transport in Plants ●● NEW Research Figure: ADH-Stimulated Water Reabsorp-
●● NEW Research Figure: Translocation Pressure tion in the Kidney Collecting Duct
xxii N E W TO T H I S E D I T I O N NEL
●● 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:
●●
and the axon hillock

Endocrine Control ●● Clarified resting and membrane potential
●● NEW Why It Matters, featuring endocrine control of mating ●● NEW Concept Fix addressing passive versus gated channels
behaviour in elk ●● Enhanced discussion of the refractory period and why it is
●● NEW Research Figure: Demonstration That Epinephrine important for nerve conduction
Acts by Binding to a Plasma Membrane Receptor ●● NEW Research Figure: Demonstration of Chemical Trans-
mission of Nerve Impulses at Synapses
Chapter 44: Animal Reproduction
Chapter 46: Muscles, Skeletons, and Body
NEW chapter created by combining reproduction and
Movements
●●
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
Welcome to Biology:
Exploring the Diversity of Life, 4Ce
Biology: Exploring the Diversity of Life and MindTap engage students so they learn not only WHAT
scientists know, but HOW they know it and what they still need to learn.
b Engage, Adapt, and Master! Stay

organized and efficient with MindTap—a
single destination with all the course
material and study aids you need to succeed.
Built-in apps leverage social media and the
latest learning technology to help you
succeed. Our customized learning path is
designed to help you engage with biological
concepts, identify gaps in your knowledge,
and master the material!
. Adapt! Reinforce your knowledge of concepts by

working through our Biology MindTap Study Guide,
which includes
• animations and videos
• topic maps, learning outcomes, and study
strategies
• multimodal quiz questions with instant feedback
• vocabulary flashcards
• chapter summaries
m Engage! The learning path for each chapter begins

with an engaging video designed to pique your
interest in the chapter contents. Take the tutorial quiz
to assess gaps in your knowledge, and strengthen your
knowledge of concepts by reviewing the ebook.
xxiv NEL
b Master! Think and engage like
a scientist by taking gradable
short-answer quizzes:
• Apply Evolutionary Thinking
questions ask you to interpret
a relevant topic in relation to
the principles of evolutionary
biology.
• Design an Experiment
challenges your understanding
of the chapter and helps you
deepen your understanding of
the scientific method as you
consider how to develop and
test hypotheses about a
situation that relates to a main
chapter topic.
• Interpret the Data questions
help you develop analytical
and quantitative skills by
asking you to interpret
graphical or tabular results of
experimental or observational
research experiments for which
the hypotheses and methods
of analysis are presented.
Test your mastery of concepts with

Aplia for Biology, a series of
Conceptual Problem Sets that
complement the text and help you
learn and understand key concepts
through focused assignments, an
engaging variety of problem types,
exceptional text/art integration,
and immediate feedback.
Assess your knowledge of chapter

concepts by taking the Post-
Learning Assessment, a set of
higher-level quiz questions
designed to test the depth of your
understanding.
NEL xxv
ism that can grow as a diploid (2n = 32) and as a haploid (n =
Next Mitosis
16). Still others, such as many plant species, have three, four, or If we set the formation of a new daughter cell as the beginning of
even more complete sets of chromosomes in each cell. The the mitotic cell cycle, then the first and longest stage is interphase
Active Learning
number of chromosome sets is called the ploidy of a cell or spe-
cies. See Chapter 18 for a look at the role of ploidy in the forma-
(Figure 7.5). Interphase comprises three phases of the cell cycle:
1. G1 phase, in which the cell carries out its function and, in
tion of new species.
Before a cell divides in mitosis, replication of the DNA in some cases, grows.
Visually stunning features that
a given chromosome produces two identical copies of that
engage your students 2. S in phase, theinprocess which DNA ofreplication and chromosome du-
learning
chromosome because
called sister an engaged
chromatids. student Newly formed is a successful
sister student.
plication occur ( Figure 7.6).
chromatids are held together along their length by proteins 3. G 2 phase, a brief gap in the cell cycle during which cell
called cohesins. During mitosis, the cohesins are removed growth continues and the cell prepares for mitosis (the
Chapter
and Roadmap
the sister chromatids are separated, with one of each pair fourth phase of the cell cycle; also called M phase) and
going to each of the two daughter nuclei. As a result of this cytokinesis.
precise division,Non-Mendelian
each Inheritancedaughter
Genetic Linkage, Sex Linkage, and Other
Mechanisms
nucleus receives exactly the same G2 refers to the second gap in which
There are many examples of traits that are not inherited in the fashion
discovered by Mendel.
there is no DNA synthesis. During

number and types of chromosomes G2, the cell continues to synthesize
and contains the same genetic infor-
10.1 Genetic Linkage and Recombination
From Chapter 8 RNAs and proteins, including those
Regents of the University of California 2005/Dr. Uli Weier/Science Source

If several genes of interest are present on the same chromosome, their traits
will not be inherited independently as shown by Mendel.
mation as the parent cell that entered for mitosis, and it continues to
grow. The end of G2 marks the end
the division. The equal distribution of a b c
of interphase; mitosis then begins.
daughter chromosomes into each of
Martin Shields/Science Source
the two daughter cells that result from 10.2 Sex-Linked Genes
cell division is called chromosome Mitosis

Some genes are carried on sex chromosomes; these traits have characteristic
Fluorescent probes show the relative location of several genes linked together along a chromosome in moths (light micrograph).
New ways of mapping chromosome structure yield insights into the inheritance of normal and abnormal traits.
inheritance patterns.
(M p Cytokinesis
10
has
segregation. 5.5c Chemiosmosis Powers ATP Synthesis by phosphorylation relies e) on the action of a large multiprotein
The precision of chromosome repli- a Proton Gradient
Genetic complex Linkage,
that spans the inner Sexmitochondrial
Linkage, membrane, called
Prophase
e
and Other Non-Mendelian
has
(Interphase ends
Martin Shields/Science Source
ATP synthase (Figure 5.16).

cation and segregation in the mitotic cellgoal of cellular respiration is the synthesis of ATP,
se
in parent cell.)
Although the
etap
a
Inheritance 5.5d ATPMechanisms
se
aph
cycle creates a group of cells called
electron a from NADH (or FADH2) to O2 does not actu-
transport 10.3 Chromosomal Mutations that Affect Inheritance
From Chapter 8
Synthase Is a Molecular Motor
ha
Changes in chromosome structure or number affect the inheritance of traits.
M
ally produce any ATP. Electrons are simply passed along a chain
An
lop
clone. Rare mutations arising during
lls s
of electron carriers until they are donatedG to oxygen, producing ATP synthase Sometime is a lollipop-shaped complex consisting of a basal
ce gin
Why it matters...
e
during the summer after his first year in music at Univer-
T
2
.)
replication make the cells of a Toclone
er be
water. understand how ATP is formed let’s go back and sité take
in
du Québec à Montréal, Julien noticed some blurry vision in one eye. By the start of classes
unit that is embedded in the inner mitochondrial membrane
the fall he was relying almost entirely on his one good eye to read music during ensemble
ht e
another look at Figure 5.14. As we have already mentioned, and connected to a headpiece by a stalk (see Figure 5.16). The
ug as
practice.
genetically very similar but not strictly
da rph
Walking out of his appointment with an ophthalmologist, Julien texted his sister, Madeleine.
NADH has more potential energy than O2 has, so one can ask“We headpiece
theneed to talk.” Madeleine was extends intoandthe
a veterinarian hadmitochondrial
always been the family matrix.
biologist. The basal unit
in nte
identical. Since all the diversequestion: cell types 10.4 Human Genetic Traits, Pedigree Analysis, and Genetic Counselling
Where does this energy go during electron transport? forms a channel through which H+ can
“They said I have Leber’s hereditary optic neuropathy,” he explained
pass freely. The proton-
to Madeleine over coffee
in his apartment. “The bad news is that it will also likely affect my other eye. I may start twitching
Inheritance patterns can reveal the underlying mechanisms of disease.
(I
tomusclesmotive force is what The propels protons inmutation
the intermembrane space
of a complex multicellular The energy that is released during electron transport is used
organism in my and there is no treatment. only good news is that the I have—here
look in the report; it’s called T14484C—is the best one to have since it often shows some improve-
theover time.through
do work, specifically the work of transporting protons acrossment They said itthe channel
is unusually common ininthe enzyme’s
Quebec. ” basal unit, G1 down
phasetheir is a con-
period of
arise by mitosis from a single cell (zygote),
© Cengage Learning 2017
inner mitochondrial membrane, from the matrix to the inter- Madeleinecentration

flipped through gradient, and and
screens on her phone into the “Wikipedia
reported, matrix. says Evidence
that T14484C indicates that
is a single base-pair change of a T to a C. The change reduces energy production growth by electron before the DNA
transport, so Ithe
guessbinding
that is what of individual protons
nerves and to sitesTheinmutation
the headpiece causes
they all contain the same genetic membrane infor- space. As a consequence of this proton pumping causes trouble in optic
G1 thatchromosome
on regular chromosomes but on your mitochondrial
muscles.
instead.”
replicates.
is not
The cell makes
across the inner membrane, which is essentially impermeable to it to rotate in a way catalyzes the formation of ATP from
mation, with only rare variation various RNAs, proteins,
protons, the in
10.5 Additional Non-Mendelian Patterns of Inheritance
H+ concentration becomes much higher (and the ADP and Pi. The spinning of the headpiece of ATP synthase rep-
Some genes are carried on organelle chromosomes; some genes are To Chapter 12
and other types of cellular
sequence. Forensic scientists rely on this
pH lower) in the intermembrane space than in the matrix.
imprinted.
resents the smallest molecular rotary motor known in nature.
molecules but use
not DNA
Proton translocation occurs at distinct sitesSalong the In Chapter 4, we described active transport pumps that
common genetic information 224
ETC (seewithin Figure 5.14). Within complexes I and IV, specific

NEL
energy from ATP to transport ions across membranes (the G in Gagainst

NEL
1 stands for gap,

225
organisms when, for instance, they

protein components use the energy released from electron their concentration gradients (see Figure 4.17). referring
An active to the absence
trans-
of DNA synthesis).
Chapter
mmatch the Roadmaps
genetic profileThe transport
of Chapter for proton pumping.
a smallRoadmaps provide aI and
molecules accept electrons from complexes
In addition, as ubiquinone
visual overview
II, they pick
port pump is, in fact, an ATP synthase that
of the major sections in the chapter and show
reverse. It doesn’t synthesize ATP but rather uses the free
is operating in
In
18886_ch10_rev02.indd 224 1/13/2018 3:13:00 AM
18886_ch10_rev02.indd 225 1/15/2018 2:35:36 PM
the connections
amount of tissuebetween
(e.g., cellsthe topics
inupdog saliva in the chapter and t
protons from the matrix. After emigrating other
rp
chapters
through the in theenergy book. from the hydrolysis of ATP to provide the energy neces-
recovered from a bite victim) with membrane thatand of donating electrons to complex ha III, ubiquinone sary to pump ions (such as protons) across a membrane.
se
retains a neutral charge by releasing protons into the inter- Harnessing the potential energy that is present in a proton
Why It Matters
a blood sample … fromWhythe It Matters
suspected
membrane
draws students in with an engaging vignette
space. gradient to synthesize that is ATP linked Gto
is fundamental
0
the concepts
to almost all forms discussed
of in
the chapter.
animal. The situation in which one side of the inner mitochondrial life and developed early in the
Cell evolution
cycle of life. This is shown by
membrane has a higher concentration of protons than the other the fact that the ATP synthasearrest complex found in mitochondria is
side represents potential energy Ifthat thecan
cellbe going Concept
is harnessedto divide,
to do DNA
Fix Icons
structurally veryConcept
similar to the Fixes ATP draw synthaseon the extensive
complexes found
work. The situation is somewhatreplicationanalogous to water research
begins. DuringaS phase,
behind literature
in the thylakoid dealing
membrane with of themisconceptions
chloroplast and in the commonly
plasma held
the cell
by duplicates each chromo- membrane of many bacteria and archaea.
dam. The potential energy possessed by
a proton gradient biologyis students. Strategically placed throughout the text,
STUDY BREAK QUESTIONS some, including
derived from two factors: First, a chemical both across
gradient exists the DNA
1. What are the three interrelated the membrane because the concentration protons is these
and theofchromosomal not equal short segments help students identify—and correct—a
proteins,
5.5e Electron Transport and Chemiosmosis
on both sides. Second, because protons and it are
alsocharged,
continues wide
there range of misunderstandings. .
synthesis
is an
systems that contribute to the
electrical difference, with the intermembrane spacemolecules.
of other cellular more posi- Can Be Uncoupled
eukaryotic cell cycle? tively charged than the matrix. The combination of a concen- Coming out of high school, many students think that
tration gradient and a voltage difference across the membrane ATP is a product of the respiratory ETC. This is a misconcep-
2. What is a chromosome composed of? FIGURE
produces stored energy as theThe
known 7.5 cell cycle.force.
proton-motive The lengthtion
of Gthat
1 varies, but for a given cell type, the timing of S phase, G2
we need to fix. The generation of ATP by the ATP
3. When is a chromosome composed of
Harnessing thephase, and mitosis
proton-motive force tois do
usually
work isrelatively
referred uniform.
synthaseCytokinesis (red segment)
complex is linked, usually
or coupled, begins
to electron while mitosis is in
transport
two chromatids? to as chemiosmosis. It was
progress first
and proposed
reaches as a mechanism
completion to
as mitosis
by ends.
the Cells
proton in a
gradientstate of division
established acrossarrest
the are
inner considered
mitochon- to enter a
generate ATP by the British biochemist Peter Mitchell, who was drial”membrane. But electron transport and the chemiosmotic
side loop (or shunt) from G1 called “G0 phase.
later awarded a Nobel Prize for Chemistry for his work in this generation of ATP are separate and distinct processes and are
area. Whereas in mitochondria, the energy for chemiosmosis not always completely coupled (Figure 5.17). For example, it
CHAPTER 7 CEll CyClEs 153
mNEL Study Breaks
The Studycomes Breaks fromfall
the at the end
oxidation of
of energy-rich molecules such as is possible to have high rates of electron transport (and thus
NADH by the ETC, chemiosmosis also applies to the genera- high rates of oxygen consumption) and yet no ATP generated
each major section and encourage students
tion of ATP to pause
by photosynthesis energy. The utility of chemios- by chemiosmosis. This uncoupling of the two processes occurs
and review what they have learned before
mosis is shown going
by the onit is not used only for ATP when mechanisms prevent the formation of a proton-motive
fact that
to the next topic within the chapter.synthesis, the proton-motive force is also used, for example, to force.
pump substances across membranes (see Chapter 4) and to A class of chemicals called ionophores form channels across
drive the rotation of flagella in bacteria. membranes through which ions, including protons, can pass
The mode of ATP synthesis that is linked to the oxidation freely. As a consequence, in the presence of ionophores, proton
xxvi NEL
h07_rev02.indd 153 of energy-rich molecules by an ETC is called oxidative phos- pumping during electron transport is followed by the protons 1/15/2018 2:28:41 PM
phorylation. Compared to the substrate-level phosphorylation simply flowing back into the matrix through the ionophore
that occurs during glycolysis and the citric acid cycle, oxidative channels. A proton gradient is prevented from becoming
NEL CHAPTER 5 CELLULAR RESPIRATION 113
b Experimental Research Figures Experimental
FIGURE 10.2 Experimental Research Research figures describe specific studies in which
Evidence for Gene Linkage research used both experimental and control
Question: Do the purple-eye and vestigial-wing genes of Drosophila assort independently?
treatments—either in the laboratory or in the field—to
Experiment: Morgan crossed true-breeding, wild-type flies having red eyes and normal wings with purple-eyed, vestigial-winged flies. The F1
dihybrids were all wild type in phenotype. Next he crossed the F1 dihybrid flies with purple-eyed, vestigial-winged flies (this is a testcross) and
test hypotheses or answer research questions by
analyzed the phenotypes of the progeny. manipulating the system they studied.
1. P generation Wild-type female Double mutant male
(red eyes, normal wings) (purple eyes, vestigial wings) The wild-type parent is
homozygous for the
wild-type allele of each
gene: pr+pr+ vg+vg+. The
purple-eyed,
vestigial-winged parent
is homozygous for the
recessive allele of each
pr + pr + vg + vg + pr pr vg vg gene: prpr vgvg.
2. F1 generation F1 dihybrid F1 dihybrid
(red eyes, normal wings) (red eyes, normal wings)
All F1 flies were wild-

type in phenotype and
were heterozygous for
both pairs of alleles:
pr+pr vg+vg.
pr + pr vg + vg pr + pr vg + vg
3. Testcross F1 dihybrid Double mutant male

(red eyes, normal wings) (purple eyes, vestigial wings)
Morgan next performed

a testcross of F1 wild-type
phenotype females with
purple-eyed, vestigial-
winged males; this is a
testcross.
pr + pr vg + vg pr pr vg vg
4. Progeny of testcross Gametes from female parent
pr+ vg+ pr vg pr+ vg pr vg+
wild type purple, red, purple,

(red, normal) vestigial vestigial normal
Gamete from
Fusion of the one type
male parent of sperm produced by
the male, and the four
pr vg types of eggs produced
by the female,
generates the progeny
of the testcross.
pr + pr vg + vg prpr vg vg pr + pr vg vg pr pr vg+ vg
Observed 1339 1195 151 154 = 2839 total progeny
Expected, Parental phenotypes Recombinant phenotypes

if assort
independently ~710 ~710 ~710 ~710 (709.75 each) = 2839 total progeny
Results: 2534 of the testcross progeny flies were parental—wild-type or purple, vestigial—while 305 of the progeny were recombinant—red,
vestigial or purple, normal. If the genes assorted independently, the expectation is for a 1:1:1:1 ratio for testcross progeny: approximately 1420 of
Research Method Figures Research Method Figures
both parental and recombinant progeny.
provide examples of important techniques, lead
Conclusion: The purple-eye and vestigial-wing genes do not assort independently. The simplest alternative is that the two genes are linked on
the same chromosome. The small number of flies with recombinant phenotypes is explained by crossing-over. students through the purpose of the technique and
228 UNIT T WO GENES NEL protocol, and describe how scientists interpret the data
generated. .
18886_ch10_rev02.indd 228 1/13/2018 3:13:02 AM

FIGURE 14.3 Research Method
Identifying a Recombinant Plasmid Containing a Gene of Interest
Purpose: To identify a recombinant plasmid containing a gene of

FIGURE 18.16 Observational Research Protocol: interest from a ligation reaction mixture containing a bacterial
Inserted DNA fragment Resealed plasmid cloning vector cloning vector and a DNA fragment containing the gene of
with gene of interest (red) with no inserted DNA fragment interest, each digested with the same restriction enzyme
Chromosomal Similarities and Differences among Humans and the Great Apes
Question: Does chromosome structure differ between humans and their closest relatives among the apes?
1. The ligation reaction produces recombinant plasmids (the only
Hypothesis: Large-scale chromosome rearrangements contributed to the development of reproductive isolation between species within the evolutionary products that might contain the gene of interest), nonrecombinant
lineage that includes humans and apes.
plasmids, and joined pieces of genomic DNA (not shown).
Prediction: Chromosome structure differs markedly between humans and their close relatives among the great apes: chimpanzees, gorillas, and orangutans.
Recombinant plasmid Nonrecombinant plasmid
Method: Jorge J. Yunis and Om Prakash, of the University of Minnesota Medical School, used Giemsa stain to visualize the banding patterns on metaphase
chromosome preparations from humans, chimpanzees, gorillas, and orangutans. They identified about 1000 bands that are present in humans and in the
three ape species. By matching the banding patterns on the chromosomes, the researchers verified that they were comparing the same segments of the
genomes in the four species. They then searched for similarities and differences in the structure of the chromosomes. 2. Transform ampicillin-sensitive, lacZ–
E. coli (which cannot make
Results: Analysis of human chromosome 2 reveals that it was produced by the fusion of two smaller chromosomes that are still present in the other three
species. Although the position of the centromere in human chromosome 2 matches that of the centromere in one of the chimpanzee chromosomes, in β-galactosidase) with a sample of the
gorillas and orangutans it falls within an inverted segment of the chromosome. ligation reaction. In this step, some
bacteria will take up DNA, whereas
Human others will not.
Bacteria transformed with plasmids Bacteria not transformed with
a plasmid or transformed
Centromere position is similar
in humans and chimpanzees. Selection: with gene fragments
Chimpanzee Bacteria transformed with plasmids
grow on medium containing ampicillin
because of ampR gene on plasmid.
Untransformed bacteria 3. Spread the bacterial cells on a plate of growth medium
Matching bands or bacteria transformed
Screening: with gene fragments
containing ampicillin and X-gal, and incubate the plate
Gorilla Blue colony contains bacteria until colonies appear.
cannot grow on medium
with a nonrecombinant containing ampicillin.
plasmid; that is, the lacZ +
Compared to the chromosomes of humans and chimpanzees,
gene is intact.
the region that includes the centromere is inverted (its position
is reversed) in both gorillas and orangutans.
Orangutan White colony contains bacteria
with a recombinant plasmid, that is, Plate of growth
the vector with an inserted DNA fragment, medium containing KEY
in this case the gene of interest. ampicillin and X-gal
Restriction lacZ+ gene
Conclusion: Differences in chromosome structure between humans and both gorillas and orangutans are more pronounced than they are between humans © Cengage Learning 2017
site Plasmid
and chimpanzees. Structural differences in the chromosomes of these four species may contribute to their reproductive isolation.
cloning
Source: © Cengage Learning 2017. Based on J. J. Yunis and O. Prakash. 1982. The origin of man: A chromosomal pictorial legacy. Science 215:1525–1530.
ampR Origin of vector
gene replication (ori)
m Observational Research Figures Observational Interpreting the Results: All the colonies on the plate contain plasmids because the bacteria that form the colonies are resistant to the ampicillin
present in the growth medium. Blue–white screening distinguishes bacterial colonies with nonrecombinant plasmids from those with
Research Figures describe specific studies in which recombinant plasmids. Bacteria making up blue colonies contain nonrecombinant plasmids. These plasmids have intact lacZ + genes and produce
β-galactosidase, which changes X-gal to a blue product. Bacteria that form the white colonies contain recombinant plasmids. Each recombinant
biologists have tested hypotheses by comparing plasmid has a DNA fragment (in this example, the gene of interest) inserted into the lacZ + gene, so β-galactosidase cannot be produced. As a result,
bacteria with recombinant plasmids cannot convert X-gal to the blue product and the colonies are white. Culturing a white colony produces large
systems under varying natural circumstances. quantities of the recombinant plasmid that can be isolated and purified for analysis and/or manipulation of the gene = “gene of interest”.
NEL xxvii
NEL CHAPTER 18 SPECIATION AND MACROEVOLUTION 447
NEL CHAPTER 14 DNA TECHNOlOGIES 337

18886_ch18_rev03.indd 447 1/12/2018 6:47:37 PM
Summary Illustrations Vivid, engaging, and carefully developed Summary Illustrations appear at the end of each chapter
and help students visualize the main concepts covered in the chapter. .
Summary Illustration
An animal cell contains a nucleus and cytoplasmic organelles,
such as mitochondria, lysosomes, the endoplasmic reticulum,
In most animals, individual cells are and the Golgi complex. It also has a highly developed
cytoskeleton. Most animal cells secrete extracellular material and
organized into tissues, which are classified have structures at their surfaces that play vital roles in the support
as epithelial, connective, muscle, or nervous. All animal cells and organization of animal body structures.
Tissues are organized into organs, and contain the same
organs into organ systems. The different basic organelles.
physiological systems operate to
homeostatically regulate body functions.
In most animals
cells
Individual cells divide

divide and
differentiate into
and differentiate into Epithelial tissue: Connective tissue: Muscle tissue: Nervous tissue:
cells of 4 different Protection, transport, Structural support Movement Communication,
secretion, and absorption coordination, and control
tissues. of nutrients released by
different tissues There are 4 types: digestion of food
that combine
and are organized
to form
organs / systems In vertebrates, there are 11 systems:
that work together Different tissues

to maintain combine and are
all body functions.
organized to form up
to 11 different organ
systems. Nervous Endocrine Muscular Skeletal Integumentary Circulatory Immune
System System System System System System System
that send
that activate
information to
effectors
integrating sites Homeostasis by
negative feedback to produce changes Respiratory Digestive Excretory Reproductive
are sensed by System System System System
that return things
receptors
to normal
Environmental changes
(internal or external) Organ systems work together to maintain homeostasis by negative feedback.
2 NEL NEL 3
c38_css_bio4ce.indd All Pages 2017-11-09 11:22 AM
Self-Test Questions These chapter review questions are organized according to Bloom’s taxonomy into three sections:
Recall/Understand, Apply/Analyze, and Create/Evaluate. This structure allows students to review the material in a sequence
that moves from the basic knowledge of factual material, to more challenging and sophisticated applications of that
knowledge to novel situations. Answers to the Self-Test Questions are found in an appendix at the back of the book. .
Appendix A: Answers to Self-Test Questions
self - test questions
Chapter 1 13. In an exergonic reaction, reactants his cells dissipated a lot of heat (the
1. a 2. a 3. d 4. a 5. c 6. d 7. c contain more free energy than the patient was hot all the time).
Recall/Understand 8. Suppose you are a doctor and your patient complains of feeling 8. a 9. b 10. a 11. c 12. b 13. d products; energy is released and the 9. b 10. c 11. d
hot all the time, even on the coldest winter days. The young 14. Eyes are usually not exposed to full reaction is spontaneous. In an ender- 12. Direct burning of glucose is an uncon-
1. Which of these factors is found in organic molecules that are man perspires constantly and his skin is always flushed. He also sunlight for a very long period of gonic reaction, reactants contain less trolled process; cellular respiration
considered good fuels? eats a lot but is rather thin. You perform some laboratory tests, time, such as the photosynthetic free energy than the products; energy occurs in a series of steps and is there-
a. many C—H bonds and find that the patient consumes lots of oxygen in his meta- apparatus is. Damage due to exposure is required and the reaction is not fore a form of controlled
b. many C=C double bonds bolic pathways. What would you suspect this patient suffers of photosystems can be repaired by spontaneous. combustion.
c. an abundance of oxygen from and why? removing damaged proteins and 14. At any time in a cell, there must be 13. Reduction is the acceptance of elec-
d. a high molecular weight replacing them with newly synthe- exergonic reactions happening to trons during a redox reaction. Oxida-
2. What is one of the places in a cell where cellular respiration Create/Evaluate sized ones, which is not possible in a provide enough energy for ender- tion is the loss of electrons during a
occurs? damaged eye. gonic reactions. In addition, the redox reaction.
a. in plant mitochondria, but not in animal mitochondria 9. In cellular respiration, which of the following does the term un- 15. Melanin protects skin cells because it energy released by exergonic reac- 14. Hydrogen and its electrons move
b. in plant chloroplasts coupled refer to specifically? absorbs ultraviolet light, and it is tions must be higher than the energy from sugar to oxygen, forming water.
c. in the mitochondria of both animals and plants a. The two parts of glycolysis are running independently of increasingly synthesized upon expo- needed for endergonic reactions 15. The process of oxidative phosphoryla-
d. in animal mitochondria, but not in plant mitochondria each other. sure to the Sun, which results in the because some energy is always trans- tion produces the large number of
b. Respiratory electron transport is operating, but chemiosmo- darker shade of her skin. ferred to heat (second law of ATP molecules needed for the ender-
3. Which of these processes occurs during glycolysis? sis is inhibited.
a. oxidation of pyruvate thermodynamics). gonic reactions in the cell that we are
c. Respiratory electron transport is operating, but proton so dependent on. One of the major
b. reduction of glucose pumping is inhibited. Chapter 2 sequences of proteins embedded in
c. oxidative phosphorylation d. Oxidative phosphorylation is occurring, but the proton- 1. d 2. c 3. c 4. a 5. a 6. b 7. b Chapter 4 the mitochondrial membrane—
d. substrate-level phosphorylation motive force remains high. 8. a 9. d 10. a 11. d 12. d 13. c 1. a 2. c 3. b 4. c
called the electron transport chain—
4. Which of these processes links glycolysis and the citric acid cycle? 14. ribosomes, rough ER, transport ves- 5. Some proteins perform transport;
10. Phosphofructokinase (PFK) is regulated by a number of metabo- can accept electrons rich in energy. As
a. chemiosmosis icle, Golgi complex, secretory vesicle, others have enzymatic activities;
lites. In addition to those mentioned in the text, which one of the the energized electrons fall from pro-
b. formation of G3P plasma membrane some are a part of signal transduction
following would also make sense? tein to protein in the ETC, they
c. reduction of NAD+ a. Pyruvate could function as an activator of PFK. 15. Anchoring junctions function to rein- process; and others are involved in
deposit energy that they carry. At the
d. pyruvate oxidation b. Glucose could function as an inhibitor of PFK. force cell-to-cell connections made by attachment and/or recognition.
end of the ETC, there must be oxygen
adhesion molecules. Tight junctions 6. b 7. c 8. b 9. c 10. c 11. b
c. ADP could function as an activator of PFK. ions present to accept these energeti-
Apply/Analyze seal the spaces between cells. Gap 12. a 13. d
d. Acetyl-CoA could act as an activator of PFK. cally depleted electrons. If these
junctions create direct channels for 14. Passive transport occurs down the
5. The breakdown of fats releases fatty acids. In what form do the 11. Which of these statements best describes the “paradox of aerobic energy-depleted electrons are not car-
communicating between adjacent concentration gradient of the solute,
carbon molecules enter the respiratory pathway? life”? ried away by the oxygen ions, ATP
cells. and active transport occurs against
a. as NADH a. Humans are completely protected from the toxic effects of production would stop. Cyanide
the gradient of the transported solute.
b. as acetyl-CoA oxygen. exerts its deadly effects by reacting
Active transport therefore requires a
c. a glucose b. Hydrogen peroxide is formed when a single electron is Chapter 3 protein and energy to perform.
with the final protein in ETC, blocking
d. as pyruvate donated to O2. 1. c 2. d 3. b 4. a 5. b 6. c 7. c 8. d oxygen from accepting electrons from
15. They are both a form of passive trans-
c. Cytochrome oxidase is a major source of reactive oxygen 9. c 10. d this protein.
6. You are reading this text while breathing in oxygen and breathing port, but facilitated diffusion utilizes
out carbon dioxide. Which two processes are the sources of the species. 11. As they dissolve, the sugar molecules proteins to speed up the transport of
carbon dioxide? d. Strict anaerobes often lack the enzyme(s) superoxide dis- raise their entropy. However, the crys- solute across the membrane. Chapter 6
a. glycolysis and pyruvate oxidation mutase and/or catalase. tals re-form because the water 1. d 2. c 3. c 4. a 5. c
b. glycolysis and oxidative phosphorylation 12. Compare direct burning of glucose and cellular respiration with decreases in its order, changing from 6. A group of pigment proteins form an
c. pyruvate oxidation and the citric acid cycle reference to their progression. compact liquid to disordered vapour. Chapter 5 antenna complex that surrounds
12. Any substance in ordered state (min- 1. a 2. c 3. d 4. d 5. b 6. c 7. a a reaction centre. Light energy
d. the citric acid cycle and oxidative phosphorylation 13. Distinguish between reduction and oxidation during redox imum entropy) will contain mole- 8. This patient might have defective absorbed anywhere in the antenna
7. Under conditions of low oxygen, what key role is played by fer- reactions. cules with maximum free energy. On mitochondria in his cells. This condi- complex is transferred to a special
mentation in overall metabolism? 14. Explain what happens with hydrogen and its bonding electrons the contrary, any substance in disor- tion is common in a number of dis- chlorophyll a molecule in the reaction
a. It regenerates the NAD+ required for glycolysis. during cellular respiration. dered state (maximum entropy) will eases. The reason why it was suspected centre. The absorbed light is con-
b. It synthesizes additional NADH for the citric acid cycle. contain molecules with minimum is that, based on his symptoms, prob- verted to chemical energy when an
c. It allows for pyruvate to be oxidized in mitochondria. 15. Cyanide is a strong toxin that reacts with the final protein in the
free energy. The relationship is ably little ATP is synthesized, in spite excited electron from the chloro-
d. By activating oxidative phosphorylation, it allows for the electron transport chain (ETC). Explain why it can kill a human
reversed. of high oxygen consumption, since phyll a is transferred to a primary
synthesis of extra ATP. within a few minutes.
NEL A-1
xxviii NEL
18886_app_rev00.indd 1 1/15/2018 8:14:16 AM
b The Purple Pages: The Chemical and
Secondary Structure
The amino acid chain of a protein, rather than being stretched
The α-Helix
A model of the α-helix (below, left), a coil shape formed when
Physical Foundations of Biology While
out in linear form, is folded into arrangements that form the
protein’s secondary structure. Secondary structure is based on
hydrogen bonds form between every N—H group of the back-
bone and the C5O group of the amino acid four residues ear-
many textbooks use the first few chapters to
hydrogen bonds between atoms of the backbone. More pre-
cisely, the hydrogen bonds form between the hydrogen atom
lier. In protein diagrams (below, right), the α-helix is depicted introduce and/or review, we believe that the
as a cylinder or barrel.
attached to the nitrogen of the backbone and the oxygen
attached to one of the carbon atoms of the backbone. Two
first chapters should convey the excitement
highly regular secondary structures are the alpha helix and the The β-Sheet and interest of biology itself. We therefore
beta sheet. In the alpha helix, side chains project outward, A β-sheet is formed by side-by-side alignment of β-strands
supporting the tertiary level of structure. Beta sheets have the (picture below shows two strands). The sheet is formed by placed important background information
side chains sticking out from the plane of the sheet alternating hydrogen bonds between atoms of each strand. In protein dia-
to either side, again supporting the overall structure. A third, grams, the β-strands are depicted as ribbons with arrowheads about biology and chemistry in the reference
less regular arrangement, the coil or loop, imparts flexibility to
certain regions of the protein. Most proteins have segments
pointing toward the C-terminal.
section entitled The Chemical and Physical
of all three arrangements.
Foundations of Biology, in the centre of the
Ball-and-stick model of α helix Cylinder representation
of α helix
book. With their purple borders, these pages
H
H H N C O are distinct and easy to find and have become
R
C
N
H
Amino acid
N
R C H H C R affectionately known as The Purple Pages.
side group
H
N R
C
O
N
C N
H O C N H References to material covered in The Purple
Pages are set in purple throughout the text.
C
HO N H O C
C H
O H N C H C R R C H
Hydrogen N O
bond C R Hydrogen
H C N H bond C O H N
C
O H O C N
N Peptide H N C O
R C HO
bond
C H N C R C H H C R
H O O
N R O C
C C N H N H
C H O C N
O N H O C
O
N
C R
C H C R R C H
H C O
O H C O H N
N
R C
H N C O
C
H O
R R C H
N H C R
C
C H O C N H
H O
N N H O C
C R
C H C R R C H
O H
C C O H N
The Green Pages: Unit 5: The Diversity

Hydrogen
bond
of Life We emphasize the richness and

F-32 T H E C H E M I C A L A N D P H Y S I C A L F O U N D AT I O N S O F B I O LO G Y NEL tremendous variability among living organisms
in The Green Pages. With their green borders,
these pages identify chapters that introduce
18886_ch32a_rev02.indd 32
and explore the tree of life. .
1/12/2018 7:42:01 PM
a. Multicellular fungus grows outward very quickly. Together, this apical growth and Fungi reproduce by spores, and this spore production can
Fruiting Body
absorptive nutrition account for much of the success of fungi. be amazingly prolific, with some species of fungi producing
As the hyphal tips extend, they exert a mechanical force, billions of spores per day (Figure 25.4). These spores are micro-
allowing them to push through their substrate, releasing scopic, featherlight, and able to survive in the environment for
enzymes and absorbing nutrients as they go. Fungal species extended periods after they are released. Reproducing via such
differ in the particular digestive enzymes they synthesize, so a spores allows fungi to be opportunists, germinating only when
Mirek Kijewski/Shutterstock.com
substrate that is a suitable food source for one species may be favourable conditions exist and quickly exploiting food sources
unavailable to another. Although there are exceptions, fungi that occur unpredictably in the environment. Releasing vast
typically thrive only in moist environments, where they can numbers of spores, as some fungi do, improves the odds that
directly absorb water, dissolved ions, simple sugars, amino the spores will germinate and produce a new individual.
Nancy Kennedy/Shutterstock.com
Honey mushroom
(Armillaria ostoyae) acids, and other small molecules. When some of a mycelium’s Spores can be produced asexually or sexually; some fungi
hyphal filaments contact a source of food, growth is channelled produce both asexual and sexual spores at different stages of
in the direction of the food source. their lives. Sexual reproduction in fungi is complex. In all
Nutrients are absorbed at the porous tips of hyphae; small organisms, sexual reproduction involves three stages: the fusion
atoms and molecules pass readily through these tips, and then of two haploid cells (plasmogamy), bringing together their two
© Cengage Learning 2017.
FIGURE 25.1 Example of a wood decay fungus: sulfur shelf fungus

Gary T. Cole, University of Texas, Austin/BPS
transport mechanisms move them through the underlying nuclei in one common cytoplasm. Cytoplasmic fusion in most
(Polyporus)
Mycelium plasma membrane. Some hyphae have regular cross-walls, or organisms is quickly followed by nuclear fusion (karyogamy),
septa (“fences” or “walls”; singular, septum), that separate a but in fungi the cells can remain bi-nucleate as the organism
hypha into compartments (Figure 25.3), whereas others lack grows. Nuclear fusion is often followed by meiosis to produce
essential components in all ecosystems and Earth’s premier b. Rhizomorphs
septa and are effectively one large cell. But even septate hyphae genetically distinct haploid spores. As we will see, fungi are
decomposers (Figure 25.1). should be thought of as interconnected compartments rather unique in that plasmogamy and karyogamy can be separated in
Despite their profound impact on ecosystems and other Fungal hyphae
than separate cells, as all septa have pores that allow cytoplasm time for durations ranging from seconds to many years.
life forms, most of us have only a passing acquaintance with and, in some fungi, even nuclei and other large organelles to
fungi, perhaps limited to the mushrooms on our pizza or the flow through the mycelium. By a mechanism called cyto-
invisible but annoying types that cause skin infections, such as
Gregory M. Filip/USDA
plasmic streaming (flow of cytoplasm and organelles around a

athlete’s foot. This chapter provides you with an overview of cell or, in this case, a mycelium), nutrients obtained by one part
fungal biology. We begin with the features that set fungi apart of a mycelium can be translocated to other non-absorptive
from all other organisms, and discuss the diversity of fungi regions, such as reproductive structures.
FIGURE 25.2 Fungal structure: mycelia, hyphae. (a) Illustration of
existing today before revisiting associations between fungi and When a fungus releases enzymes into its substrate, it faces
themycelium of a mushroom-forming fungus, which consists of branching
other organisms. septate hyphae. Inset: Micrograph of fungal hyphae (b) Rhizomorph – a competition from bacteria and other organisms for the nutri-
cordlike aggregation of hyphae formed by some basidiomycete fungi. ents that are now available. How can a fungus prevent these
competitors from stealing the nutrients that it has just expended
25.1 General Characteristics energy and resources to obtain? Many fungi produce antibacte-
Regardless of their nutrient source, fungi feed by absorp-
of Fungi tive nutrition: they secrete enzymes into their environment,
rial compounds and toxins that inhibit the growth of com-
peting organisms. Many of these compounds are secondary
We begin our survey of fungi by examining the features that breaking down large molecules into smaller soluble molecules metabolites, which are not required for day-to-day survival but
distinguish fungi from other forms of life, how fungi obtain that can then be absorbed into their cells. This mode of nutri- are beneficial to the fungus. As we will see, many of these com-
nutrients, and adaptations for reproduction and growth that tion means that fungi cannot be stationary, as they would then pounds not only are important in the life of a fungus but also
enable fungi to spread far and wide through the environment. deplete all the food in their immediate environment. Instead, benefit organisms associated with the fungus. Many are also of
Fungi are heterotrophic eukaryotes that obtain carbon by fungi have evolved the ability to proliferate quickly through commercial or medical importance to humans; for example,
breaking down organic molecules synthesized by other organ- their environment, digesting nutrients as they grow. How can the antibiotic penicillin is a secondary metabolite produced by
isms. Although all fungi are heterotrophs, fungi can be divided fungi proliferate so quickly? Although some fungi grow as uni- a species of Penicillium.
into two broad groups based on how they obtain carbon. If a cellular yeasts, which reproduce asexually by budding or
fungus obtains carbon from non-living material, it is a sapro- binary fission (see Figure 25.13), most are composed of hyphae
troph. Fungi that decompose dead plant and animal tissues, for (“web”; singular, hypha; Figure 25.2), fine filaments that spread
Hyphae
example, are saprotrophs. If a fungus obtains carbon from through whatever substrate the fungus is growing in—soil,
living organisms, it is a symbiont. Symbiosis is the living decomposing wood, your skin—forming a network, or
Andrew Darrington/Alamy Stock Photo
together of two (or sometimes more) organisms for extended mycelium (Figure 25.2). Hyphae are essentially tubes of Apical growth
periods; symbiotic relationships range along a continuum from cytoplasm surrounded by cell walls made of chitin, a polysac-
parasitism, in which one organism benefits at the expense of charide also found in the exoskeletons of insects and other
the other, to mutualism, in which both organisms benefit. arthropods. Septum
Although we often think of fungi as decomposers, fully half of Hyphae grow only at their tips, but because a single
all identified fungi live as symbionts with another organism. mycelium contains many, many tips, the entire mycelium FIGURE 25.3 Septa. In some fungi, septa divide each hypha into separate FIGURE 25.4 Spore production by fungal fruiting bodies. Some
compartments. fruiting bodies can release billions of spores per day.
588 UNIT FIVE THE DIVERSIT Y OF LIFE NEL NEL CHAPTER 25 FUNgI 589
NEL xxix
18886_ch25_rev03.indd 588 1/16/2018 2:16:12 AM 18886_ch25_rev03.indd 589 1/13/2018 6:18:37 PM
MindTap
Stay organized and efficient with
MindTap—a single destination with all InSTRUCToR R
Student and Instructor Resources the course material and study aids you need to succeed.
Built-in apps leverage social media and the latest
learning technology. MindTap To ensure the high quali
ancillaries have been pro
For example: Engage, adapt, and master!
Be sure to visit
• ReadSpeaker will read
Engage yourthe text towith
students you.the person-
Instruction website at
Succeed in the course with these dynamic resources! • Flashcards
alized are prepopulated
teaching experience of MindTap.to provide Withyou with a
relevant
find out more about NET
assignments that guide students to analyze, apply, own.
jump start for review—or you can create your and
of instructors who have
• You
elevate can highlight
thinking, MindTap textallows
and make notes toinmea-
instructors your
MindTap sure MindTap
skills andReader. promote Yourbetter
notes will flow into
outcomes withEvernote,
ease.
seen student engageme
the electronic notebook appthis thatonline
you can accessand any- NETA Test Bank:
With relevant assignments that guide Including interactive quizzing, tutorial
wheretool when it’s timeeachto study for theunique
exam. needs Ivona Mladenovic of
students to analyze, apply, and elevate diagnostic identifies student’s
includes over 2500 mu
thinking, MindTap allows instructors to measure skills and pro- with Self-quizzing
a pretest. The learning allows path youthen to helps
assess your
students
understanding. according to NETA gui
mote better outcomes with ease. Including interactive quizzing, focus on concepts they’re having the most difficulty
tion and development o
this online tutorial and diagnostic tool identifies each student’s Also available
mastering. It refers into the the MindTap
accompanying for BiologyMindTap are
engaging andand informative Test Bank was copy-ed
unique needs with a pre-test. The learning path then helps stu- Reader eBook provides avideosvarietythat accompany
of learning the
activi-
Purple Pages. From matter and reviewed by David
dents focus on concepts they’re having the most difficulty mas- ties designed to appeal to to polypeptides,
diverse ways ofTodd Nickle
learning.
of Mount Royal University (pictured) willtake walk you NETA best practices.
tering. It refers to the accompanying MindTap Reader eBook After completing the study plan, students Aplia
through sets these essay, short-answer, ma
and provides a variety of learning activities designed to appeal problem andfoundational
then take a posttestconcepts, strengthening
to measure their
your understanding tions. Test Bank files ar
to diverse ways of learning. After completing the study plan, understanding of theand helping
material. you buildhave
Instructors a strong
the
base oftoknowledge format for easy editing
students take Aplia problem sets and then take a post-test to ability customizeand the understanding
learning path, for addbiology.
their own
Visitand www.nelson.com/student to progress
start using prin
measure their understanding of the material. Instructors have content, track and monitor student by
MindTap.
using Enter the Gradebook
the instructor Online Access andCodeProgressfromapp. the The
card
the ability to customize the learning path, add their own con- able
included with your textbook.
MindTap resources have been developed by DoraIf a code card is not provided,
tent, and track and monitor student progress by using the form
instructor Gradebook and Progress app.
Aplia
you canfor
Cavallo-Medved Biology
purchase instant
of the access
University at NELSONbrain.com.
of Windor and
Cognero® is a secure on
Students stay organized and efficient with MindTap, a single Reehan MirzaAplia ofStrengthen
Nipissing
for BiologyUniversity.
your understanding of biologyyou withto author, edit, an
destination with all the course material and study aids students Aplia™! from any place you ha
Strengthen your understanding of
need to succeed. Built-in apps leverage social media and the Aplia’s focused assignments and active
installations or down
Aplia
biology forwith
Biology
Aplia™!
latest learning technology. For example, learning opportunities
This innovative, easy-to-use, interactive help students learn key
desktop-inspired interf
Get your students engaged andautomatically
motivated
ReadSpeaker will read the text to you. technology givesconcepts you more bypractice,
randomized, with detailed feed- graded and familiar, intuitive t
with Aplia for Biology.
●●
questions,
back to help exceptional
you learn with text/art
every integration,
question! and immediate manage feed- tests with ease
Flashcards are prepopulated to provide you with a jump start
Aplia Help your students learn key concepts via
●●
back.Aplia’s has a full course management
focused assignments and active learning system that can be
versions in an instant
for review, or you can create your own. Aplia’s focused assignments and active
used independently
opportunities (including or inrandomized
conjunctionquestions, with other course into
excep- man-other systems. Tes
You can highlight text and make notes in your MindTap learning opportunities that include randomized, WebCT.auto-
●●
agement systems
tional text/art such as
integration, Blackboard
and immediate and feedback) get learning management
Reader. Your notes will flow into Evernote, the electronic matically graded questions, exceptional text/art inte-
This innovative,
you involved with biology easy-to-use,
and help you interactive
think liketechnology
a scien- gives
wherever you want. Ne
notebook app that you can access anywhere when it’s time to gration, and
students moreimmediate
tist. For more practice, with
information, feedback.
visitdetailed Aplia
feedbackhas to
www.aplia.com/biology. a full
help students
nero can be accessed
study for the exam. course management system
Self-quizzing allows you to assess your understanding.
learnInteractive
with everyproblems
question! andthat can be
figures helpused
youindepen-
visualize instructor.
●●
dently or
Aplia’s in conjunction
focused with
assignments other
dynamic biological processes and integrate concepts, and course
active manage-
learning opportuni- NETA PowerPoint:
The MindTap resources were developed by Dora Cavallo- ment
ties systems
(including such as
randomized
art, media, and homework practice. Blackboard
questions, and WebCT.
exceptional Visit
text/art inte-
slides for every chapter h
Medved of the University of Windsor, Reehan Mirza of Nipissing www.aplia.com/biology.
gration, and immediate feedback) get students involvedofwith the University of Nor
University, Roy Rea of the University of Northern British StudyThe guide
biology Apliahelp
and coursethem forthink
Biology:likeExploring
scientists.the Diversity is an average of 80 slid
Columbia, and Miranda Meents. of Life, Third Canadian Edition, was prepared by Anna
The Interactive
Study Guideproblems and figures help from
(ISBN: 978-0-17-660144-7) studentsthe visualize
key figures, tables, an
Also available in MindTap for Biology are engaging and Rissanen
dynamic of Memorial
biological University,
processes and and Todd Nickle
integrate concepts, andart, media,
Exploring the Diversity o
second Canadian edition has been adapted for the third
informative videos that accompany The Purple Pages. From Alexandria
and homework
Canadian Farmer
edition byofDora
practice.Mount Royal University.
Cavallo-Medved of the Univer- The PowerPoint slides
matter to polypeptides, author Todd Nickle, of Mount Royal For
Instructor’s
sity of moreResource
Windsor, information,
Roy V. Rea DVD: ofvisit
theThe www.aplia.com/biology.
ultimate
University tool for
of Northern selected illustrations wit
University (pictured), will walk you through these foundational customizing
TheColumbia,
British Aplia lectures
course andand
for presentations.
Biology:
Julie Smit,Exploring the Diversity
of the University of ofbeenLife,reworked to allow
concepts, strengthening your understanding and helping Select
Windsor.
Fourth instructor
The
Canadian Study ancillaries
Guide
Edition, helps are provided
students
was prepared integrate
by Anna on thethe NETA
Rissanen of principles of clea
you build a strong base of knowledge and understanding for Instructor’s
concepts within
Memorial Resource
University DVD
the text and
and (ISBNTodd978-0-17-660143-0)
provides study strategies,
Nickle of Mount Royal have been incorporated
biology. interactive
and exercises,
the Instructor
University. self-test questions,
Companion site. and more. for instructors to custom
Visit www.nelson.com/student to start using MindTap. The Nelson Education Teaching AdvantageImage Library: Th
The Nelson Education Teaching
xxxii
Enter the Online Access Code from the card included stUdent
with your resOUrces Advantage
(NETA) program (NETA) program delivers delivers copies of figures, short
research-based
textbook. If a code card is not provided, you can purchase instructor resources that promote student engagementinand
research-based instructor resources that the book. Instructor
instant access at NELSONbrain.com. higher-order thinking to enable the success of Canadian students the NETA Power
promote student engagement and higher-order thinking tomize
to enable the success of Canadian students and educators. PowerPoint presentatio
32137_00_fm.indd 32
xxx NEL NEL
ancillaries have been professionally copy-edited.
er!
Be sure to visit Nelson Education’s Inspired
h the person-
Instruction website at www.nelson.com/inspired/ to
With relevant and out
educators. To ensure
find more about NETA.the high
Don’t quality
miss of these materials, all
the testimonials ciples of clear design and engaging content have been incorpo-
ze, apply, and Nelson ancillaries have been professionally copy-edited. rated throughout, making it simple for instructors to customize
of instructors who have used NETA supplements and
ctors to mea- Be sure to visit Nelson Education’s Inspired Instruction the deck for their courses.
seen student engagement increase!
es with ease. website at www.nelson.com/inspired/ to find out more about Image Library: This resource consists of digital copies of
NETA Test Bank: This resource was written by
e tutorial and
Ivona Mladenovic of Simon Fraser University. It have used
NETA. Don’t miss the testimonials of instructors who figures, short tables, and photographs used in the book. Instruc-
unique needs NETA supplements and have seen student engagement
includes over 2500 multiple-choice questions written increase! tors may use these jpegs to customize the NETA PowerPoint
elps students NETA Test Bank: This resource was written by Ivona Mlade- slides or create their own PowerPoint presentations.
according to NETA guidelines for effective construc-
ost difficulty NETA Instructor’s Manual: This resource was written by
tion and development of higher-order questions. 2500
novic of Simon Fraser University. It includes over The multiple-
ng MindTap choice questions written according to NETA guidelines for Tamara Kelly of York University and Tanya Noel of the Univer-
Test Bank was copy-edited by a NETA-trained editor
arning activi- effective construction and development of higher-order questions. sity of Windsor. It is organized according to the textbook chap-
and reviewed by David DiBattista for adherence to
of learning. The Test Bank was copy-edited by a NETA-trained
NETA best practices. Also included are true/false, editor for ters and addresses key educational concerns, such as typical
ts take Aplia adherence to NETA matching,
best practices.
essay, short-answer, and Also includedques-
completion are true/false, stumbling blocks students face and how to address them. Other
measure their essay, Test
short-answer,
tions. Bank filesmatching,
are provided andincompletion
Microsoft Word questions. Test features include tips on teaching using cases as well as sugges-
ors have the Bank files
format for areeasyavailable
editing in andMicrosoft
in PDF for Word format from your
convenient tions on how to present material and use technology and other
dd their own Nelson publishing representative. resources effectively, integrating the other supplements available
printing, whatever your system.
progress by NETA
The The TestTestBank is available
NETA Bank is avail- in a new, to both students and instructors. This manual doesn’t simply
ress app. The cloud-based platform. Nelson Testing Pow-
able in a new, cloud-based plat- reinvent what’s currently in the text, it helps the instructor make
ped by Dora
ered by Cognero® is a secure
form. Nelsononline
Testingtesting systembythat allows
Powered the material relevant and engaging to students.
Windor and
you to author,
Cognero ® edit, and
is a secure manage
online test system
testing bank content from any place
that allows TurningPoint®: Another valuable resource for instructors
you to
you have Internet
author, access.
edit, and No specialtest
manage installations
bank content or downloads is TurningPoint® classroom response software customized for
are needed,
from any place andyou thehave
desktop-inspired
Internet access. interface, with its drop-
No special Biology: Exploring the Diversity of Life, Fourt Canadian Edition,
down menus or
installations anddownloads
familiar, intuitive tools, allows
are needed, and you the to create by Jane Young at the University of Northern British Columbia.
and manage tests interface,
desktop-inspired with ease. with
You can create multiple
its drop-down menus test versions Now you can author, deliver, show, access, and grade, all in
nd motivated in anfamiliar,
and instant intuitive
and import or export
tools, allows you content into other
to create and systems. PowerPoint, with no toggling back and forth between screens!
Tests cantests
manage be delivered
with ease. from
Youyour learning
can create management
multiple test system, JoinIn on TurningPoint is the only classroom response software
concepts via your classroom, or wherever you want. Nelson Testing Powered
versions in an instant and import or export content tool that gives you true PowerPoint integration. With JoinIn,
s and active into other systems.
by Cognero Tests can
can be accessed be delivered
through from your
www.nelson.com/instructor. you are no longer tied to your computer. You can walk about
omized, auto- NETA PowerPoint: Microsoft PowerPoint® lecture
learning management system, your classroom, or slides your classroom as you lecture, showing slides and collecting
text/art inte- wherever
for every you chapterwant. Nelson
were Testing
created by Jane Powered
Young by of Cog-
the University and displaying responses with ease. There is simply no easier or
a has a full nero can beBritish accessed through www.nelson.com/
of Northern Columbia. There is an average of 80 slides more effective way to turn your lecture hall into a personal,
sed indepen- instructor.
per chapter, many featuring key figures, tables, and photographs fully interactive experience for your students. If you can use
urse manage-
fromNETA PowerPoint:
ExploringMicrosoft PowerPoint lecture Canadian
®
Biology: the Diversity of Life, Fourth PowerPoint, you can use JoinIn on TurningPoint! (Contact
WebCT. Visit slides for every chapter have slides
been created by Jane “build
Young slides”—
Edition. The PowerPoint also feature your Nelson publishing representative for details.) These con-
of
selected illustrations with labels from the bookThere
the University of Northern British Columbia. that have been tain poll slides and pre- and post-test slides for each chapter in
g the Diversity is an average of 80 optimal
slides perdisplay
chapter,
reworked to allow in many featuring
PowerPoint. NETA prin- the text.
ared by Anna key figures, tables, and photographs from Biology:
dd Nickle and Exploring the Diversity of Life, Third Canadian Edition.
ersity. The PowerPoint slides also feature “build slides”—
mate tool for selected illustrations with labels from the book that have
been reworked to allow optimal display in PowerPoint.
vided on the NETA principles of clear design and engaging content
17-660143-0) have been incorporated throughout, making it simple
for instructors to customize the deck for their courses.
on Teaching Image Library: This resource consists of digital
ram delivers copies of figures, short tables, and photographs used
resources that in the book. Instructors may use these jpegs to cus-
rder thinking tomize the NETA PowerPoint slides or create their own
and educators. PowerPoint presentations.
instrUctOr resOUrces xxxiii
21/01/15 2:05 AM
NEL S T U D E N T A N D I N S T R U C TO R R E S O U R C E S
xxxi
Acknowledgements
We thank the many people who have worked with us on the It is never easy to be in the family of an academic scientist.
production of this text, particularly Paul Fam, Senior Publisher, We are especially grateful to our families for their sustained
whose foresight brought the idea to us and whose persistence support over the course of our careers, particularly during
saw the project through. those times when our attentions were fully captivated by
We are also grateful to the members of the MindTap bacteria, algae, fungi, parasites, snakes, geese, or bats. Saying
Advisory Board and the Student Advisory Boards for the fourth “yes” to a textbook project means saying “no” to a variety of
Canadian edition, who provided us with valuable feedback and other pursuits. We appreciate the patience and understanding
alternative perspectives (special acknowledgments to these of those closest to us that enabled the temporary reallocation of
individuals are listed below). considerable time from other endeavours and relationships.
We also thank Richard Walker at the University of Calgary Many of our colleagues have contributed to our develop-
and Ken Davey at York University, who began this journey with ment as teachers and scholars by acting as mentors, collabora-
us but were unable to continue. We are very grateful to Heather tors, and, on occasion, “worthy opponents.” Like all teachers,
Addy of the University of Calgary for her significant contribu- we owe particular gratitude to our students. They have gath-
tions to the first three editions. ered with us around the discipline of biology, sharing their
We are especially grateful to Toni Chahley, Content potent blend of enthusiasm and curiosity, and leaving us ener-
Manager, who kept us moving through the chapters at an effi- gized and optimistic for the future.
cient pace, along with Charu Verma, Project Manager, and
Imoinda Romain, Senior Production Project Manager. A very Editorial and Student Advisory Boards
special thanks to Kathy Hamilton, for her guidance and helpful We were very fortunate to have the assistance of some extraor-
suggestions with the summary illustrations, and we are very dinary students and instructors of biology across Canada who
grateful to Roy Rea of the University of Northern British provided us with feedback that helped shape this textbook into
Columbia for his critical read of Chapter 27, and his significant what you see before you. As such, we would like to say a very
contribution to the summary illustrations for Chapters 27–32. special thank you to the following people:
We thank Kristiina Paul, our photo researcher, for her hard work
MindTap Advisory Board
with the numerous photos in the book, and Frances Robinson
for her careful and thoughtful copy-editing. Finally, we thank Brett Couch, University of British Columbia
Kim Carruthers, Marketing Manager, for making us look good. Stewart Daly, Marianopolis College
Brock Fenton thanks Allan Noon for offering advice about Jon Houseman, University of Ottawa
taking pictures; Laura Barclay, Jeremy McNeil, Tony Percival- William Huddleston, University of Calgary
Smith, C. S. (Rufus) Churcher, and David and Meg Cumming Ivona Mladenovic, Simon Fraser University
for the use of their images; Karen Campbell for providing a Ken Otter, University of Northern British Columbia
critical read of Putting Selection to Work; and Michael Owen Lisa Prichard, MacEwan University
for his outstanding contribution to Chapter 27: Animals. Roy Rea, University of Northern British Columbia
Tom Haffie would like to acknowledge the cheerful and Frieder Schoeck, McGill University
insightful editorial work of Jennifer Waugh on Chapter 16 and Marina Silva-Opps, University of Prince Edward Island
the conscientious research assistance of Dr. Aniruddho Matt Smith, Wilfrid Laurier University
Chokroborty-Hoque. Chris Todd, University of Saskatchewan
Denis Maxwell would like to thank David Brock for helping Paula Wilson, York University
fine-tune the coverage of thermodynamics in Chapter 3. Ken Wilson, University of Saskatchewan
Todd Nickle thanks his family, students, and colleagues
for humouring his mad exploits and for their understanding
when he doesn’t make things easy on himself—or others—as he
explores insane and sometimes creative ways to repurpose old
ways of doing things.
The authors are all indebted to Ivona Mladenovic of Simon
Fraser University for her excellent work on the Self-Test
Questions, and Johnston Miller, whose extensive background
research anchored our Concept Fixes in the education
literature.
xxxii NEL
M.B. Fenton
(Top) Ivona Mladenovic, Ken Wilson, Jon Houseman, Stewart Daly, Frieder
Schoeck; (bottom) William Huddleston, Chris Todd, Lisa Prichard
Paula Wilson
UNBC
Paula Wilson, York University Ken Otter, University of Northern British Columbia
John Ternan
Roy V. Rea
Roy Rea, University of Northern British Columbia Matt Smith, Wilfrid Laurier University
NEL A C K N O W L E D G E M E N T S
xxxiii
Student Advisory Boards Mount Royal University
Alexis Andres / Nelson

Depicted above are members of the Student Advisory Board at Mount Royal
University. From left to right, back row: Evan Olar, Moroni Lopez, Taelor Evans,
Todd Nickle, Jonathan Roveredo, Darlene Skagen, Andrew Roberts, Surafel Girma,
Danielle Schmidt, Laura Villarraga Ulloa. Front row: Kyle Poffenroth, Heaven Berhe
Paul Fam / Nelson
Sium, Aderinsoye Ademoye, Ravneet Gill, Meena Kanthimathinathan, Alexandra

Presbitero, Anastasia Socolnicova. Not pictured: Ashley Chicote, Cassidy Fleming
Shown above are members of the University of British Columbia Student Advisory
Board. From left to right: Kristina Balce, Amy Dhillon, Humaam Hamado, Ebi Oliya,
Laura Fash, Sina Soleimani, Adam Book, Ashley Pinter, Garrett Huwyler, Quinn Nelson would like to thank some of the current student users of
Stewart, Gavindeep Shinger. Not pictured: Kendrix Kek, Hassan Ali Biology: Exploring the Diversity of Life, in particular the students
at Western University’s Biology 1001A course who shared their
feedback via their participation in a student focus group early in
University of Calgary the development process.
Alexis Andres / Nelson
Shown above are members of the University of Calgary Student Advisory Board.
From left to right: Gina Mannella, Jesse Provick, Amanda Bennett, Leesa Le, Vivian
Nguyen
xxxiv A C K N O W L E D G E M E N T S NEL
VOLUME 1
BIOLOGY OF THE CELL
Lebendkulturen.de/Shutterstock.com
Volvox is a genus of green algae. As photosynthetic eukaryotes, Volvox cells exist in colonies of
thousands of cells. Some of the cells are vegetative (non-reproductive); a smaller number of much
larger reproductive cells are found in the interior of the colony.
There is a huge diversity of life on Earth; some estimates peg the total number of species at over
1 billion (with most yet to be described!). Yet, what this opening volume of the textbook should
covey to you is that underlying that diversity is a remarkable level of similarity. From monkeys to
mycoplasma to monocots, everything that is alive on Earth employs a variation on that remarkable
innovation: the cell. No matter if that cell is communicating with other cells in the brain of a fruit
fly, or capturing sunlight in a spruce needle, or driving the muscles of a sprinting cheetah, or
thriving in the mineral-rich water of deep-sea vents, no matter what their role or activity, all cells
share a remarkably long list of common features. Volume 1 explores these common features
in detail.
NEL
The invention of the microscope allowed scientists to finally cells contain ribosomes where some kinds of RNAs get
understand how living organisms were built from cells, which translated into proteins, the fundamental structural,
to early scientists were astonishingly small. Further work clearly f unctional, and r egulatory molecule of the cell.
delineated two major divisions in cell types: those without a Genes are stretches of DNA sequence in an organism that
nucleus (prokaryotic) and those with a nucleus (eukaryotic). collectively comprise a kind of library of information about
And within both these groups there are clear subdivisions: for how a cell functions. Recent advances in technology have made
example, plants, fungi, and animal cells, in eukaryotes. it relatively easy to determine the entire DNA sequence of an
The chapters of Unit 1 talk a lot about energy because organism, including individual humans. As a result, modern
without it living cells would die. Like the non-living world, all biology is awash in the As, Ts, Gs, and Cs of DNA sequence
forms of life abide by the foundational laws of thermodynamics revealed by thousands of sequencing projects. New insights
and need to bring in energy and matter from the environment into evolutionary history as well as gene structure and func-
to maintain their highly ordered state. In part, energy is tion are arising from bioinformatic analysis of such extensive
required to build complex things (e.g., proteins) out of simpler data sets.
things (amino acids). In addition to energy, the evolution of life The elegant double-strandedness of DNA, whereby two
is tied to the development of a remarkable group of proteins long strands of nucleotides are held together by hydrogen bonds
called enzymes, which when they get tied to a biochemical reac- formed between complementary base pairs, affords a straight-
tion can increase its rate by 1010 times! forward mechanism for replication that was recognized early
Another remarkable feature of cells that we dedicate a on by Watson and Crick. Although conceptually simple, the
chapter to in Unit 1 is membranes. These self-forming lipid mechanism for unwinding the DNA double helix and polym-
bilayers act as the gatekeepers of the cell: they allow certain erizing new complementary bases is rather complicated and
things in but keep other things out. How they do this is by managed by a suite of interacting enzymes. Again, we see that
acting in concert with membrane-specific proteins that shuttle all DNA on the planet is replicated using variations on one
molecules from one side to another. Membrane proteins also underlying strategy.
play a remarkable role in transducing signals from outside the DNA genes provide the cell with needed RNA by tran-
cells and compartments to the inside. As we will see, the trans- scription. One remarkable feature of all protein-coding genes is
duction of signals associated with hormones, for example, can that, with minor exceptions, the information they carry is
profoundly affect cell function. specified by a universal code. That is, a gene from one organism
What you may not realize is that virtually all the energy can be “understood” by any other organism, even if only dis-
used by living systems comes ultimately from sunlight being tantly related: a gene from a spider can be expressed by a goat;
harvested and converted into a useable chemical form through a gene from a jellyfish can be expressed in a flower. The field of
photosynthesis. This process evolved perhaps as early as genetic engineering is devoted to developing the tools and
3.5 billion years ago and used photons of light energy to extract applications of this technology for moving genes from one
electrons from water, releasing oxygen as a by-product. The rise organism to another.
in oxygen in an atmosphere that previously had none led to an In a story that is about to come full circle, synthetic biolo-
explosion of life as the mechanism of cellular respiration gists have extensively customized naturally occurring cells and
evolved that could use that oxygen, and enabled cells to pro- have made important advances toward their ultimate goal of
duce huge amounts of energy. creating novel life forms artificially in the lab. As students of
The chapters of Units 2 and 3 are dedicated to molecular biology in the early twenty-first century, you can well expect to
biology and genetics, the central player being the gene. All cells witness a momentous event in Earth’s history, the creation of
possess genes that are coded by the molecule DNA that, one life form by another.
through the process of transcription, get copied into RNA. All
2 V O LU M E O N E B i olo g y o f t h e C ell NEL
Chapter Roadmap
Light and Life

Light serves two important functions for life on Earth: First, it is the ultimate
source of energy for almost all organisms. Second, light provides many
organisms with information about the physical world that surrounds them.
1.1 The Physical Nature of Light

To be used as a source of energy or information, photons of light must be To Purple Pages
absorbed by a molecule.
1.2 Light as a Source of Energy

The energy of light can be used in photosynthesis to convert carbon dioxide To Chapter 6
into carbohydrates.
1.3 Light as a Source of Information To Chapter 37

The photoreceptor, a pigment molecule bound to a protein, is the common
molecule used in sensing light in the environment. To Chapter 45
1.4 The Uniqueness of Light

Light possesses an ideal amount of energy: enough energy to excite
electrons, but not enough to directly destroy biological molecules.
1.5 Light Can Damage Biological Molecules

Organisms possess mechanisms to both protect themselves from the
damaging effects of light and repair the inevitable damage that light causes.
1.6 Using Light to Tell Time

Organisms have evolved circadian clocks, which allow them to keep track of
time on a daily and seasonal scale.
Anita Patterson Peppers / Shutterstock.com
1.7 The Role of Light in Behaviour and Ecology To Chapter 30

The light environment of a particular habitat plays a central role in
adaptation. It leads to unique colouration that may serve to attract a mate
or hide from predators. To Chapter 32
1.8 Organisms Making Their Own Light

Some organisms generate their own light to attract a mate or prey, or for
communication.
4 NEL
(Left) The Japanese Bridge (The Water-Lily Pond), Claude Monet, 1899; (Right) The Japanese Bridge at Giverny, Claude Monet, 1926
Paintings by Claude Monet (1840–1926). Compared to his early works, including The Water Lily Pond (a), his later paintings, including The
Japanese Footbridge (b), bordered on the abstract, with almost complete loss of light blue. Monet suffered from cataracts, a degenerative
vision disease, diagnosed in 1912.
Light and Life

Why it matters . . . Claude Monet (1840–1926), a French painter, is considered by many
to be the master of the impressionist form that rose to prominence in the late nineteenth century.
Impressionism as an art movement was characterized by the use of small visible brush strokes that
1
emphasized light and colour, rather than lines, to define an object. The artists used pure, unmixed
colour, not smoothly blended, as was the custom at the time. For example, instead of physically
mixing yellow and blue paint, they placed unmixed yellow paint on the canvas next to unmixed
blue paint so that the colours would mingle in the eye of the viewer to create the impression of
green. The impressionists found that they could capture the momentary and transient effects of
sunlight and the changing colour of a scene by painting en plein air (in the open air), outside the
studio, where they could more accurately paint the reflected light of an immediate scene.
Interestingly, compared with his early works, which included The Water Lily Pond (1899),
Monet’s later paintings verge on the abstract, with colours bleeding into each other and with a
lack of rational shape and perspective. For example, The Japanese Footbridge is an explosion of
orange, yellow, and red hues, with heavy, broad brush strokes, leaving the viewer barely able to
discern the vague shape of the arched bridge. In many of Monet’s later works, the colours in his
paintings became more muted, far less vibrant and bright, with a pronounced colour shift from
blue–green to red–yellow and an almost total absence of light blues. The sense of atmosphere and
light that he was famous for in his earlier works disappeared.
NEL 5
Although the change in Monet’s paintings could easily be the process of photosynthesis. In addition, each cell contains a
explained by an intentional change in style or perhaps an age- light sensor called an eyespot that allows Chlamydomonas to
related change in manual dexterity, Monet himself realized gather information about the direction and intensity of the light
that it was not his style or dexterity that had changed but, in its environment. With this information, cells can move
rather, his ability to see. Monet suffered from cataracts, a toward or away from the light source using a pair of flagella.
vision-deteriorating disease diagnosed in both eyes when he Regardless of whether the light is used as a source of energy or
was 72. A cataract is a change in the lens of the eye, making it as a source of information about the environment, both rely on
more opaque, which changes the ability to see different colours the same fundamental properties of light.
of light.
In this chapter, the first of the 46 of this textbook, we intro- 1.1a What Is Light?
duce you to the science of biology by using light as a central
connecting theme. Light is arguably the most fundamental of Through the process of nuclear fusion, the Sun transforms a
natural phenomena, and foundational experiments into the staggering 3.4 × 1038 hydrogen nuclei into helium each second
nature of light were a key part of the scientific revolution that (Figure 1.2). In the process, about 4 million tonnes of matter are
took place in the sixteenth and seventeenth centuries. Beyond converted into energy. This energy is given off by the Sun as
formally defining light and discussing its properties, in this electromagnetic radiation, which travels in the form of a wave at
chapter we explore the huge diversity of areas of biology that a speed of 1.1 × 109 km/h (the speed of light) and reaches Earth
light influences, from the molecular to the ecological. This in just over 8 minutes. Electromagnetic radiation is generated at
introductory tour is not intended to be complete or exhaustive a range of wavelengths (Figure 1.3): cosmic rays have a wave-
but to simply set the stage for the topics that come in subse- length of less than one picometre (10–12 m); radio waves have a
quent chapters. wavelength longer than one kilometre (106 m). The complete
range of wavelengths of electromagnetic radiation is referred to
as the electromagnetic spectrum.
1.1 The Physical Nature of Light So what is light? Light is most commonly defined as the
portion of the electromagnetic spectrum that we can detect
Light serves two important functions for life on Earth: First, it is with our eyes. As shown in Figure 1.3, this is a very narrow
the ultimate source of energy that sustains virtually all organ- portion of the total electromagnetic spectrum, spanning only
isms. Second, light provides many organisms with information the wavelengths from about 400 to 700 nm.
about the physical environment in which they live. These two The physical nature of light has been the focus of scientific
roles for light are nicely illustrated by the green alga Chlamydo- inquiry for hundreds of years, and in many ways it remains a
monas (Figure 1.1). Chlamydomonas is a single-celled, photosyn- mystery. Unlike the atoms that make up matter, light has no
thetic eukaryote that is commonly found in ponds and lakes. mass. And although the results of some experiments suggest
Chlamydomonas contains a single large chloroplast that harvests
light energy and uses it to make energy-rich molecules through
Flagella Earth
Vacuole
Mitochondrion
Eyespot
Nucleus
NASA
Chloroplast
FIGURE 1.2 The Sun. Like most stars, the Sun generates electromagnetic
FIGURE 1.1 Chlamydomonas. Each cell contains a single chloroplast radiation as a result of the nuclear fusion of hydrogen nuclei into helium.
used for photosynthesis, as well as an eyespot for sensing light in the Note the superimposed image of Earth used to illustrate the relative sizes.
environment.
6 CHAPTER 1 Light and Life NEL
a. Range of the electromagnetic spectrum
The shortest, Range of most Range of heat The longest,
most energetic radiation reaching the escaping from the lowest-energy
wavelengths surface of Earth surface of Earth wavelengths
Gamma X-rays Ultraviolet Near-infrared Infrared Microwaves Radio

rays radiation radiation radiation waves
Visible light
400 450 500 550 600 650 700

Wavelength of visible light (nm)
b. Examples of wavelengths
400 nm 700 nm
wavelength wavelength
FIGURE 1.3 The electromagnetic spectrum. (a) The electromagnetic spectrum ranges from gamma rays to radio waves; visible light and the
wavelengths used for photosynthesis occupy only a narrow band of the spectrum. (b) Examples of wavelengths show the difference between the longest
and shortest wavelengths of visible light.
that light behaves as a wave as it travels through space, other in vision; and indigo, which is used to dye jeans their distinc-
experiments indicate that light behaves more like discrete par- tive blue colour.
ticles of energy called photons. In the end, we are left with a An important question we can ask is: What is it about pig-
compromise description: light is best understood as a wave of ments that enable them to capture light? At first glance, the
photons. An important aspect of light to remember is that there molecules shown in Figure 1.5 seem to be very different from
is an inverse relationship between the energy of a photon and each other structurally. However, they all have a common fea-
the wavelength of light. Looking at Figure 1.3, this means that ture critical to light absorption: a region where carbon atoms
shorter-wavelength blue light consists of photons of higher are covalently bonded to each other with alternating single and
energy than red light, which has a longer wavelength and thus double bonds. This bonding arrangement is called a conjugated
photons of lower energy. system, and it results in the delocalization of electrons. These
electrons are not closely associated with a particular atom or
1.1b Light Interacts with Matter involved in bonding to another atom, and instead are available
to absorb the energy of a photon of light.
Although light has no mass, it is still able to interact with matter While the presence of a conjugated system is common to
and cause change. This change is what allows the energy of light all pigments, differences in the arrangement of the conjugated
to be used by living things. When a photon of light hits an
object, the photon has three possible fates: it can be reflected off
the object, transmitted through the object, or absorbed by the Photon
object. To be used as a source of energy or information by an
organism, the light must be absorbed. Light is absorbed when
the energy of the photon is transferred to an electron within a
Electron at
molecule. This transfer of energy excites the electron, moving it ground state Low-energy level
from its ground state to a higher energy level that is referred to
as an excited state (Figure 1.4). Electron at
High-energy level
Molecules differ considerably in their ability to absorb excited state
photons of light. There is a major class of molecules that is very

efficient at absorbing photons of specific wavelengths; those
molecules are called pigments (Figure 1.5). As you would expect, FIGURE 1.4 The absorption of a photon by a molecule results in the
there is a huge diversity of pigments, including chlorophyll a, energy being transferred to an electron. This causes the energy to move
which is involved in photosynthesis; retinal, which is involved to a higher-energy, excited state.
NEL CHAPTER 1 Light and Life

7
CH2CH3 CH3
H3C O Light
N N Chlorophyll a
CO2CH3
Mg
N N H
H
H2C=CH
CH2CH2CO2CH2CH=C(CH2CH2CH2CH)3CH3
CH3 H CH3
CH3 CH3
O H
N Reflected
light
N
H O
H O
11-Cis-retinal Indigo
Protein
COO– COO– HO HO
H3C S O O
H
H3C HO
H
H
H HO HO OH OH
O N N N N O O
H3C O
H H H
HO
Phycoerythrobilin Carmine
FIGURE 1.6 Why the T-shirt is red. Pigment molecules bound

H3C
to the fabric of the shirt absorb blue, green, and yellow photons
CH3 CH3
H3C CH3 of light. Red photons are not absorbed and are instead
H3C CH3 transmitted through the shirt or are reflected.
CH3 CH3
CH3
Beta-carotene
FIGURE 1.5 Structure of some common pigments. Chlorophyll a, photosynthesis;

11-cis-retinal, vision; indigo, dye; phycoerythrobilin, red photosynthetic pigment found 1.2 Light as a Source of Energy
in red algae; carmine, scale pigment found in some insects; beta-carotene, an orange
accessory photosynthetic pigment. A common feature of all these pigments that is
The ultimate source of the energy used by almost all
critical for light absorption is the presence of a conjugated system of double/single organisms that make up the biosphere is light from the
carbon bonds (shown in red for beta-carotene). Sun. The energy of electromagnetic radiation is made
accessible through the ability of plants and related
organisms to convert the energy of photons into chem-
system as well as differences in the overall chemical structure ical energy. Through photosynthesis, plants absorb photons of
explain why each type of pigment absorbs light of only certain light and use that energy to convert carbon dioxide and water
wavelengths. This is because, for a photon to be absorbed, the into sugars and other molecules.
energy of the photon must match the amount of energy needed Following light absorption by the pigment chlorophyll, the
to move a delocalized electron from its ground state to a spe- high potential energy of excited-state electrons is used in pho-
cific excited state. If the energies don’t match, then the photon tosynthetic electron transport to synthesize the energy-rich
of light is not absorbed and instead is transmitted through the compounds NADPH (nicotinamide adenine dinucleotide
molecule or reflected off the molecule. phosphate) and ATP (adenosine triphosphate). These molecules
The ability of pigments to absorb specific wave- are in turn consumed in the biochemical reactions of the
lengths of light is what determines their colour. A pigment’s Calvin cycle of photosynthesis to convert carbon dioxide into
colour is the result of photons of light that it does not absorb. carbohydrates (Figure 1.7). Although the energy of one photon
Instead of being absorbed, these photons are reflected off the is very small, the photosynthetic apparatus within the chloro-
pigment or transmitted through the pigment and reach your plast of a single plant leaf absorbs millions of photons each
eyes (Figure 1.6). second. And a single cell within a typical plant leaf contains
hundreds of chloroplasts!
While photosynthesis converts carbon dioxide into carbo-
STUDY BREAK QUESTIONS hydrates, it is the process of cellular respiration, which is found
1. What has to occur for a photon to be absorbed? in all organisms, that breaks down carbohydrates and other
2. What structural feature is common to all pigments? energy-rich molecules, trapping the released energy as ATP
(Figure 1.7). The value of ATP is that it is the universal energy
8 CHAPTER 1 Light and Life NEL
Energy is stored as currency, and can be readily used for the energy-
chemical energy. requiring metabolic and biosynthetic processes that
Sugar
are required to maintain all life.
Oxygen Oxygen
Photosynthesis is the dominant process of the
biosphere that directly uses the light of the Sun as a
Electromagnetic source of energy, but you may be surprised to know
energy in sunlight that it isn’t the only one. Another light-driven pro-
cess used to acquire energy is found in a group of
microbes called Halobacterium. These organisms
are not eukaryotes or even bacteria, but rather they
Photosynthesis captures Cellular respiration
electromagnetic energy releases chemical energy belong to the third domain of life, the Archaea. We
from sunlight. from sugar molecules. will introduce you to the three domains of life in
Chapter 21, and will discuss the Archaea in greater
detail in Chapter 22. Most often found in extreme
habitats, Halobacterium contains a pigment–pro-
tein complex called bacteriorhodopsin, which is
found on the plasma membrane and functions as a
Carbon dioxide Released chemical light-driven proton pump. When the pigment com-
energy is made
and
available for other
ponent of bacteriorhodopsin captures a photon of
Water metabolic processes. light, it triggers changes in the protein component,
resulting in the specific transport of protons out of
FIGURE 1.7 Photosynthesis converts light energy into chemical energy. the cell. The resulting difference in H+ concentra-
Photosynthesis uses the energy in sunlight to build sugar molecules from carbon dioxide
tion across the plasma membrane represents a
and water, releasing oxygen as a by-product. The process of cellular respiration breaks
down the products of photosynthesis and releases usable energy.
source of energy that is used by the enzyme ATP
synthase to generate ATP from ADP and inorganic
phosphate (Pi) (Figure 1.8). Like all organisms, the
a. Halobacterium salinarum b. Hutt Lagoon, Western Australia

FIGURE 1.8 Halobacterium is a
genus of Archaea that have a
light-driven proton pump.
(a) Electron micrograph of a
colony of Halobacterium
J Marshall - Tribaleye Images / Alamy Stock Photo

salinarum. (b) Species of
Halobacterium thrive in high-salt
environments, such as Hutt
Lagoon in Australia. The pink
colour of the water is due to the
presence of bacteriorhodopsin
within individual cells. (c) A model
NASA
of bacteriorhodopsin shows the

pigment retinal bound to a
c. A model of bacteriorhodopsin d. Bacteriorhodopsin-driven ATP formation
protein. (d) Bacteriorhodopsin
Light functions as a light-driven proton
pump, the proton gradient being
Inside cell
COO– H+
used to synthesize ATP.
Bacterio- ATP
rhodopsin synthase
ADP + Pi
H+
Outside cell ATP

Phospholipid
membrane Bacterial cell
Retinal
NH+3
NEL CHAPTER 1 Light and Life

9
Another random document with
no related content on Scribd:
—Je ne vous la décrirai donc pas... Mais voici Boucbélère qui revient; la
séance n’a pas été longue; et voici James.»
M. Randal semblait indigné, tristement indigné. Il s’appliquait à garder
un calme que démentait le trouble de sa voix.
«Boucbélère, dit-il d’abord, veuillez vous retirer.»
Puis, quand le délinquant fut parti, l’oreille basse:
«Ma chère Ida et vous, monsieur Delannes, je vous fais juges, tous deux,
d’un cas infâme. Présenter au peuple les images les plus désolantes de la
détresse humaine, cela ne se défend que par l’excellence du but que l’on
veut atteindre. Un pareil spectacle force à réfléchir, à rentrer en soi-même;
il apporte une leçon douloureuse et, par conséquent, un bienfait. On oublie
si vite sa santé! Être normal, cela paraît tout naturel; on n’y songe pas... Je
donne, ici, l’occasion d’y songer et j’incite à en rendre grâces, un jour, à qui
de droit. C’est une prière qui monte, c’est une prière de plus. L’homme sain
remercie Dieu de sa santé, au lieu de le supplier seulement au cours d’une
maladie. Je pense que, pour sa rareté même, cette prière inattendue sera
agréée, comme un don gratuit... Et que vient de me proposer Boucbélère,
pour la seconde fois? un cul-de-jatte qui joue avec son infirmité, qui fait le
singe, qui fait le clown! qui dessine la caricature de sa déchéance et
provoque la gaieté par une parade sacrilège! A la façon de Ned Walkins, il
casse des douzaines d’assiettes, sans arrêt, avec un sourire surpris et cette
expression sottement ravie qui, chez Walkins, était une trouvaille... A-t-on
jamais vu un forçat jongler avec ses chaînes?... A coup sûr, ce cas est
infâme, et vous ne me contredirez pas!»
Il se tut, il s’éloigna d’un pas rapide, sentant qu’il ne se tenait plus en
main.
Mme Randal ne paraissait nullement émue.
«Vous le retrouverez souvent dans cet état. J’avoue que j’ai peine à le
comprendre, car, en somme... N’importe!... Au revoir, Monsieur.»
Il ne restait à Delannes que de prendre congé.
XI
Pendant le jour, Mathieu errait souvent aux abords du camp, et le soir,
après la fermeture des grilles, s’attardait en de longues causeries, jusqu’à
l’heure où un tintement de cloche annonçait pour tous la fin de la veillée.
«Je trouve là, disait-il à son ami Hourgues, des gens qui m’intéressent,
avec qui je m’entends bien: Sam Harland me parle de ses chevaux; je les
connais presque tous et plus d’un m’a déjà fait mordre la poussière. On se
moque de moi qui prétendais être bon cavalier; on me donne des conseils
pratiques; je les suis.
—Avery Leslie me plaît beaucoup: il me décrit ses premiers essais sur la
corde, ses projets, ses tentatives, ses erreurs et ses réussites. Le ton sincère
qu’il met à m’expliquer tout cela finit par me convaincre. Je partage bientôt
ses peines et ses plaisirs... Il m’arrive de chercher avec lui quelque
perfectionnement nouveau à la construction de son balancier, quelque façon
inédite de mettre en valeur son périlleux passage aérien. J’y réussis parfois.
D’autres me racontent de belles histoires, simples comme des images
d’Épinal, mais un peu longues... d’autres me disent leur vie; tous, ils
s’efforcent de se faire comprendre, ce qui attire la sympathie. Assurément,
il y a Boucbélère qu’il faut subir de temps en temps, mais on finit par
excuser sa bassesse: ses discours ont tant de naïveté comique! tant
d’abandon! Cela désarme.
—Oh! s’écria Hourgues, le Boucbélère: un bouffon lugubre! Et que
pensez-vous des patrons de la troupe, du couple Randal?
—Le vieux m’ahurit: il est tellement particulier, étranger... comment
dire?... unique en son genre! Pas bête, certes, assez noble, et, tout de même,
effarant! Quant à sa femme, elle paraît intelligente, mais, en quelque sorte,
pas à sa place. Je la connais peu. Qu’en dites-vous?
—J’ai rarement causé avec elle... Une expression bizarre, n’est-ce pas?
Elle a beaucoup déplu à Alice, tout de suite, parce qu’elle s’entend mal avec
les enfants. Vous savez que ma femme a des opinions très particulières,
certains préjugés: elle se méfiera volontiers de quelqu’un que les enfants ni
les bêtes n’aiment.
—Alice a raison.
—D’ailleurs, Mme Randal est une curieuse figure. Elle exerce sur sa
troupe une influence très forte, dont elle se doute à peine, dirait-on, ou dont
elle a peur... On respecte Randal, on l’admire; elle, on ne la perd jamais de
vue, on obéit à son moindre signe, on a l’air de la considérer comme un
fétiche... le porte-bonheur... le porte-guigne du Randal Circus... Comment
savoir?...
—Sa façon si brusque de s’exprimer me gêne, dit Mathieu, un mélange
de réserve et de passion assez inquiétant: on ignore où l’on va...
—Parlez d’Ida Randal aux hommes de la troupe et vous jugerez de
l’importance de son rôle.
—Que faites-vous, ce soir, Hourgues?
—Des écritures indispensables, puisqu’il nous faut cette machine
agricole dont je vous parlais hier... et vous?
—Je vais me promener un peu, regarder la lune... Elle s’arrondit
délicieusement.
—Rendez donc visite à vos amis du camp. C’est je ne sais quelle fête
d’anniversaire, en Amérique. Ici, l’on veillera jusqu’à minuit, pour
commémorer.
—Excellente idée. Vous ne m’accompagnez pas?
—Non: cette lettre, quelques papiers à classer, et je me couche.
—Tant pis; dormez bien, mon ami.
—Belle promenade, Mathieu!»
Ils se quittent.
Le paysage vaut, en effet, d’être contemplé longuement. Immobiles, sans
un frisson de feuilles, les arbres se dressent, tout argentés, devant leurs
ombres bleues, et le gazon prend d’étranges teintes mauves. Enfin, sur la
mer, c’est une vaste scintillation de féerie, une piste éblouissante, poudrée
de diamants pour quelque divine chevauchée.
Le camp, moins silencieux que d’habitude, ne dort pas encore. Des feux
brillent de-ci, de-là, on entend parfois sonner des rires... Un peu de musique
passe, poussive ou grêle, qui n’offre rien d’émouvant mais qui n’inquiète
pas trop l’oreille.—Sans doute, Sam Harland joue-t-il de l’accordéon, sa
pipe à la bouche, l’œil malin, l’air bonhomme et satisfait, puis ce sera John
Plug, palefrenier de son état, acrobate à ses heures et connu par sa virtuosité
sur un instrument soufflé en figure obèse de citrouille, dont il se sert à
merveille au cours d’un numéro de clowneries fantasques. De ce fruit
démesuré qu’il lui faut saisir à pleins bras, il tire une toute petite mélodie
dessinée en fil de fer, qui monte et se tortille, anormale et falote, presque
plaisante.
On chante aussi: chansons populaires, sentimentales, souvenirs du pays
natal, évocations d’images lointaines... près du foyer, là-bas, une mère
tricote, elle attend; penchée à sa fenêtre, une fiancée rêve; sujets de cartes
postales. Aucun hymne: la fête gardera, ce soir, un ton laïque, un ton très
moral aussi, car personne, bien entendu, ne boit de vin, à l’intérieur du
camp, et toute joie grossière est interdite par un règlement signé James
Randal, dûment affiché, qui, en paragraphes précis, loue ou réprouve,
conseille ou blâme les formes diverses du plaisir. On s’y conforme; on ne
s’amuse pas moins.
Mathieu reste debout devant une barrière de bois, non loin du hangar
illuminé, ruche de chants et de rires. On l’aperçoit, on crie aussitôt à l’ami
«français» d’entrer au plus vite; il est reçu avec des paroles bruyantes de
bon accueil où le «welcome!» domine.
XII
Plus tard, Mathieu se rappela souvent cette nuit et son croissant de lune
et cette longue veillée.
Une trentaine de convives sont installés autour de quelques tréteaux,
devant de hautes cruches pleines de limonade. Chacun a son gobelet;
certains l’accrochent à leur ceinture et, souffrant de rester immobiles,
marchent de long en large, la pipe à la bouche, puis reviennent boire;
certains jouent aux dominos, aux dames, d’autres au bilboquet, le plus
sérieusement du monde, en comptant les coups, sauf un maladroit qui
s’excuse de ses ratés par des contorsions burlesques.—Peu de femmes: miss
Jones, la dactylographe du chef, trois écuyères mariées, la caissière,
personne mûre dont les lunettes n’attristent pas le visage souriant et joufflu;
celle-ci tricote des bas et cause avec tout le monde; qui donc l’a surnommée
«Joy-for-ever», à cause de sa constante et facile gaîté? on ne l’appelle pas
autrement. Sous la visière de sa casquette, une maigre, très maigre dame
interprète, qui sait mal toutes les langues parlées, discourt de mille choses,
sur quel ton d’assurance! enfin Rachel Boucbélère, minuscule, vêtue de
noir, fripée, l’air mécontent et boudeur, fait sans trêve des patiences sur le
coin d’un tréteau, manie nerveusement ses cartes crasseuses, puis son
collier d’ambre, quand «ça ne vient pas», et prend, en désespoir de cause,
une expression sournoise du plus haut comique pour tricher inaperçue.
Boucbélère la surveille de loin, gras, sale, des bagues aux doigts.
Mathieu s’assit entre Sam Harland et Avery Leslie.
«Vous auriez dû arriver plus tôt, dit Harland, notre camarade Boucbélère
vient de chanter une chanson que je n’ai pas très bien comprise, mais qui...
—Ah! c’est qu’il y avait de l’argot de Paris, s’écria Boucbélère d’une
voix alliacée, si vous voulez...
—Merci, je dois la connaître, interrompit Mathieu, craignant qu’il ne
recommençât.
—Un de ces soirs, fit Avery Leslie, moi aussi, je vous chanterai une
chanson. Je l’ai entendue, d’abord, en me promenant sur les quais du Havre,
la nuit, devant les bateaux, et je n’ai pas été long à l’apprendre... Je ne sais
pas qui la chantait. C’est une chanson pour monter le long de la corde
oblique, avec le balancier ou le parasol. Elle exprime le danger, la joie,
l’espoir d’arriver et la prudence qu’il faut garder jusqu’au bout, et
l’impatience qui me travaille à mi-chemin... Je la chanterai en moi-même,
pour moi-même; elle sera mon guide... Non, je ne vous la chanterai pas ici,
car vous n’entendriez rien du tout; c’est une chanson pour le cœur.
—Et comment avez-vous senti que cette chanson vous était destinée?
—Je vais vous le dire, monsieur Mathieu, mais il ne faudra pas vous
moquer... Tous ces cordages, n’est-ce pas, tendus devant la mer, éclairés par
la lune et les feux, et qui s’entre-croisaient, cela me faisait tourner la tête; je
souffrais de ce vertige dont j’ai peur quand je travaille... Mais la chanson
montait si droit, malgré les ficelles et les lumières, qu’elle me rendait toute
ma confiance, tout mon équilibre; le malaise disparut et j’appris la chanson.
—Mon cher Leslie, répondit Mathieu, chacun de nous a besoin d’une
chanson pareille pour les passages difficiles de sa vie, mais certains ne la
trouvent jamais; il faut, je crois, la mériter d’abord, à votre façon.
—Tu vois, Avery, dit Sam Harland, que M. Mathieu n’avait pas envie de
se moquer de toi.»
Auprès des autres causeries, plus bruyantes, celle-ci, à voix presque
basse, se perpétuait entre Mathieu, le danseur de corde et l’écuyer.
«Déjà, dit Mathieu, quand vous montez le long de la corde, vous avez
soin de fixer votre regard à son extrémité. Vous ne faites pas autre chose,
quand vous chantez en vous-même: vous fixez votre pensée...
—Oh! oui!...
—Moi aussi, monsieur Mathieu, dit Sam Harland, je fixe ma pensée. Le
métier d’écuyer, ce n’est pas une route unie. Il faut prendre garde à la
bouteille de gin sur la droite, à la bouteille de whisky sur la gauche, qui
vous font signe, toutes deux, de descendre et de goûter, et puis il y a des
fossés et des caniveaux que l’on ne voit pas d’abord, où le cheval
s’embronche, et surtout, il y a la fatigue de rester en selle si longtemps,
quand on pourrait être mieux assis dans un bar, avec des camarades et des
compagnes, ce qui ne servirait qu’à mener ces hommes et ces femmes dans
la même prison... Alors, moi, pour ne pas trop pécher, je fixe ma pensée,
comme vous dites, je fixe ma pensée sur une belle image, et, tout de suite,
je n’ai plus envie de boire ni de toucher au vice.»
Il parlait simplement, tranquillement, semblant avoir peur de faire des
phrases ou de paraître trop sérieux. Afin de s’excuser un peu, il
accompagna ses dernières paroles d’un sourire...
Mais un incident sut distraire tout le monde. La porte du fond s’ouvrit,
chacun se leva. On se mit à chanter de nouveau, un chœur cette fois, que
l’on eût dit entonné par ordre ou pour faire honneur.
Quelqu’un entrait.
Le chant montait, unanime, véritable hymne de salutation. Les amateurs
de bilboquet haussèrent leurs boules à bout de bras et John Plug, étreignant
passionnément sa citrouille, la délivra d’un cri de petit pourceau...
«Ratée! pour la septième fois!» gémit Rachel Boucbélère en brouillant
ses cartes...
Alors, on vit s’avancer, coiffée d’un voile gris qui serrait ses cheveux,
vêtue d’un tailleur gris de coupe nette, une badine à la main, souriante,
élégante, élancée, le regard posé devant elle comme sur des sujets de sa
dépendance, la reine de la troupe, son idole peut-être: Ida Randal.
XIII
Cette entrée fit sensation. Mathieu songeait à des scènes de cinéma où
l’héroïne, impatiemment attendue, paraît enfin; et pourtant, quoi de plus
naturel? Ida Randal se joignait aux réjouissances de sa troupe réunie, un
soir de fête.
«Plug! s’écria-t-elle en riant clair, n’oubliez surtout pas ce que vous avez
inventé, à l’instant: ce cri nouveau, sorti de votre citrouille! Je vous promets
un beau succès si vous le retrouvez au cirque, dans un sketch!
—On inventerait bien autre chose pour l’amuser, dit Avery Leslie à mi-
voix.
—Ah! pour sûr! affirma Sam Harland, en passant sa pipe dans le coin
gauche de sa bouche.
—Et tous, mes amis, je vous remercie de cet accueil... Bonsoir,
Boucbélère, Leslie, Harland; bonsoir, Joy-for-ever.»
Ravie, les yeux au ciel, la caissière soupira:
«Dear lady!
—Bonsoir, Rachel! Ah! monsieur Delannes, c’est gentil de nous rendre
visite.
—Madame, je passe une soirée excellente...»
En somme, Mathieu se sentait content de la revoir. Leur première

rencontre, leur seule conversation, devant la tente de Randal, lui laissait un
souvenir trouble, et il disait vrai en affirmant à Hourgues qu’il ne
connaissait pas cette femme dont certains propos l’avaient gêné, l’avaient
surpris... Elle l’intriguait: qui était-elle?
Poussant sa chaise, il fit à Ida Randal une place auprès de lui.
«Votre mari viendra-t-il, Madame?
—Non,» dit-elle...
Et, tout de suite après, mais plus bas:
«Il suffit de moi pour tuer l’entrain d’une réunion comme celle-ci.»
Sans avoir disparu, la joie de cette fête n’était cependant plus la même:
on s’entendait mieux, le bavardage sonore s’assourdissait, et il semblait
aussi que chacun, tout en parlant, chantant ou riant, ne perdait pas de vue
celle qui venait de s’asseoir et qui causait avec Mathieu, tantôt en anglais,
tantôt en français, mais toujours d’une façon rapide, impersonnelle et
dégagée, qui passait inaperçue.
«Il faudra revenir souvent, monsieur Delannes. On vous aime bien dans
la troupe.
—J’en suis heureux, Madame, et je compte me faire, au Cirque Randal,
des amis.
—Vous en avez déjà. On apprécie votre bonne camaraderie, votre
simplicité.
—A fréquenter tout ce petit monde dans son décor, je m’instruis et
m’amuse mieux qu’en traînant mes guêtres à Paris. Être simple et bon
camarade, cela ne souffre, ici, pas de difficulté.
—Je le conçois; encore faut-il y mettre du sien, ce que vous faites avec
aisance.»
Ils ne se regardaient pas; ils parlaient, en quelque sorte, devant eux. Ils
ne ressentaient nul besoin de se communiquer leurs pensées autrement que
par des phrases dites sur un ton banal.
Boucbélère se levait. Il chanta de nouveau, et ce fut une lamentable
romance parfumée de roses, palpitante d’hirondelles. Des gloussements
émus, des gestes pathétiques accentuaient les beaux passages amoureux.
«Oh! s’écria Rachel, quand mon Octave dit qu’il aime, moi, je l’adore!
—Elle montre de la vaillance, murmura Mme Randal.
—Comment pouvez-vous chanter ces choses, Boucbélère? demanda
Leslie sur un ton de parfaite candeur.
—Plus tard, petit garçon, tu les chanteras aussi pour plaire aux femmes!
—Je ne pense pas, grogna Sam Harland.
—Mais... vous croyez à tout cela que vous racontez?»
Une explosion de gaîté bruyante fut la seule réponse du chanteur.
«Si vous ne le croyez pas, Boucbélère, alors, c’est vilain! déclara Leslie
qui semblait souffrir.
—Une leçon? à moi! oh! mon petit, va danser sur ta corde!...
—Fichez donc la paix à cet enfant,» interrompit Mme Randal d’une voix
nette.
Boucbélère, ayant pris le ciel à témoin de la pureté de ses intentions, se
rassit, le visage marqué d’une grimace excessive d’ironie. Rachel, très
nerveuse, mais qui n’osait intervenir, le flatta d’un long regard, comme elle
eût déclaré: «Je suis de cœur avec toi, mon bel Octave!»
Et la fête continua, coupée d’intermèdes.
«Avez-vous repensé au discours de mon mari? demanda Mme Randal.

—Souvent, Madame, répondit Mathieu, mais je n’ai guère eu l’occasion
de m’entretenir avec lui; une fois seulement, avant-hier, où il m’a défini et
développé, avec beaucoup de bienveillance, la règle morale de sa troupe.
Cela m’a paru, tout ensemble, très judicieux et très élevé.
—Oui... une police de protestant.
—Si vous voulez, mais qui explique son influence acceptée par chacun.
—Et dont certains ne se félicitent pas!
—La vôtre aussi est intéressante à étudier, Madame.
—La mienne?
—... Si manifeste: elle se retrouve partout et toujours.
—Je l’ignorais.
—Non, Madame, vous la sentez fort bien: votre entrée, il y a deux
heures, dans cette salle où nous sommes, la montrait clairement et prouvait
même que vous en aviez conscience! Il suffisait de suivre votre regard
dominateur. Tous vos sujets tournaient les yeux vers vous, vers vous seule,
et vous leur en saviez à peine gré...
—Sans doute écrivez-vous des romans psychologiques, cher Monsieur...
des romans français!
—Je n’y ai jamais songé, je vous assure, mais il m’arrive de prendre des
notes, de remarquer ceci ou cela, de me souvenir aussi, quand il faut.»
Ce fut à cet instant qu’il considéra le visage d’Ida Randal et s’aperçut
que le beau visage était pâle.
Malgré lui, avec la maladresse que l’on met souvent à réparer, il ajouta:
«Pardon, Madame!»
Sans broncher, elle répondit:
«Je vous pardonne.»
XIV
Certaines paroles d’Ida Randal avaient dérouté Delannes. Il restait
silencieux, prêtant l’oreille, vaguement, aux bruits de la fête finissante,
regardant autour de lui les gestes exaltés ou comiques, mais déjà lassés,
illustrant une joie à son déclin que bientôt le sommeil étouffera.
Ida s’était levée, elle se promenait de table en table, disait bonsoir à
chacun, causait un peu, posait quelque question, donnait un encouragement,
et, de nouveau, Mathieu fut frappé d’une expression commune à tous ces
hommes réunis... Elle ne se retrouvait pas chez les femmes: miss Jones, la
dactylographe, causait de ses affaires, la dame interprète précisait avec
autorité la prononciation d’un vocable français, les trois écuyères
échangeaient des potins à voix basse, seule Joy-for-ever gardait cette
béatitude vivante dont témoignaient sa bouche ronde, ses joues roses et,
sous le verre des lunettes, ses yeux bleus d’enfant.
«Dormez bien, ma chère. C’est un plaisir de vous voir ici; j’aime vous
entendre rire. Je vous fais aussi mon compliment sur la façon remarquable
dont votre caisse est tenue.»
C’en était trop pour Joy-for-ever, trop d’émotion:
«An angel! s’écria-t-elle, an angel from heaven!»
Leslie avait entendu... Il se pencha vers Mathieu et murmura:
«Oh! oui! un ange, un ange du ciel!»
Mme Randal continuait sa promenade et Mathieu la regardait. Cette
beauté, indéniable assurément, n’évoquait rien d’angélique ni de céleste.
Mince, fine, Ida paraissait grande, bien qu’elle fût de taille moyenne. Ses
mouvements avaient quelque chose d’élastique, d’aisé, de facile,
d’entraînant aussi, que l’on retrouve chez les bêtes de chasse ou de course,
et son visage aigu aux yeux jaunes rapprochés donnait une impression de
dureté cruelle, à cause du petit nez courbe et fin, de la mâchoire obstinée et
surtout d’une large bouche frémissante qui, semblait-il, devait sourire
difficilement, méchamment peut-être. De légers cheveux noirs moussaient
avec abondance sous le voile gris, et la robe de même teinte, très simple, au
dessin net, accentuait l’allure de ce corps jeune, plein de santé, de vigueur.
Mathieu avait déjà remarqué les mains intelligentes, la cambrure du pied, la
cheville... Oui, mais que voyait-on là qui fût d’un ange ou vînt du ciel?
«Un ange du ciel, répétait Leslie à mi-voix, un ange descendu droit du
ciel!... n’est-ce pas, monsieur Delannes?»
Comment répondre à pareil propos?
Heureusement, Ida, qui allait franchir la porte, se retourna sur le seuil
même et fit signe à Mathieu. Il s’excusa auprès de ses voisins par quelques
paroles amicales et se hâta de la rejoindre.
«Vos bois sont merveilleux, à cette heure, dit-elle, et la nuit semble très
douce. J’ai envie de suivre jusqu’au bout le petit sentier, vous savez bien,
celui qui passe sous les chênes et coupe le ruisseau. Accompagnez-moi.»
Sans dire mot, Delannes acquiesça par un salut, et ils sortirent.
XV
Douce, tiède, surprenante par sa tranquille pureté, après une telle
atmosphère de tabagie, mais très obscure, la nuit ne portait à son front
qu’un mince croissant mouillé. Ce trait courbe d’argent se découpait seul, à
mi-hauteur du ciel noir, la brume offusquant les étoiles, au-dessus du rideau
des arbres d’un noir plus mat.
Le bois lui-même était opaque et tiède; on y voyait à peine; peu
importait aux deux promeneurs qui semblaient bien connaître le chemin.
Saisis par cette ombre embuée, ils se turent, d’abord, écoutant le bruit de
leurs pas. On n’entendait d’ailleurs que ce bruit mou et, parfois, au sein des
feuilles, un frisson furtif: réveil d’oiseau? battement d’ailes? passage
d’écureuil? Puis Mme Randal se mit à parler, sur un ton très simple, très
posé; elle reprit au point où elle voulait reprendre:
«Un jour, dit-elle, il vous parlera de moi, sans préambule, à sa manière
que je qualifiais de protestante: il me citera, comme il citerait un
personnage quelconque de la Bible ou de l’histoire, pour servir d’exemple à
ce qu’il raconte... Il vous expliquera que, moi aussi, j’ai été ramenée au
bien, qu’il m’a trouvée sur une scène de music-hall, au Canada, où je
dansais des danses singulières, de mon invention, qui lui plurent, dont il
escomptait, je pense, le succès sur un de ses programmes... qu’il voulut me
parler, après la représentation, et qu’aussitôt il comprit qu’il m’aimait, qu’il
ne pouvait me laisser là, que je devais le suivre... Trois mois plus tard, je
m’appelais Mme Randal... Et c’est toute mon histoire: une rencontre
fortuite, à Toronto, une conversation dans un bar avec un directeur de
cirque, un engagement signé sur le bord d’une table sale... un engagement
pour la vie! Parfois, quand il me regarde, je sens que je suis sa proie, celle
qu’il a sauvée du marécage. Il ne ment pas: il m’a sauvée du marécage...
mais pourquoi le dire? et s’il ne vous l’a pas dit, hier, il vous le dira
demain... pourquoi le dire à tout le monde, puisqu’il m’aime?»
Sans violence encore, sans éclats, sa voix s’était cependant réchauffée.
Ida, marchant à côté de Mathieu, ne le voyait pas. Eût-elle osé parler ainsi à
une autre heure, en d’autres lieux? Ses mots, sitôt prononcés, se perdaient
dans la nuit; elle n’en pouvait noter l’effet, elle n’en devinait pas l’action;
elle laissait tomber son aveu comme en un puits sourd.
Sur un ton presque timide, un peu hésitant, Mathieu demanda:
«Du moins, êtes-vous heureuse, Madame?
—Je n’en sais rien, répondit-elle. Je ne suis pas libre!
—Comment l’entendez-vous?»
Elle répéta:
«Je ne suis pas libre! Vous ne sentez donc pas ce que cela veut dire? Oh!
j’ai toute liberté d’agir à ma guise, d’aller à droite, à gauche, où il me plaît,
mais puis-je penser et sentir à ma guise?... Ma tête n’est pas libre! En ce
cas, il vaut mieux être enchaîné pour de bon, comme les forçats.
—Chacun de nous est retenu par quelque lien, Madame...»
Il rougit d’avoir proféré une banalité si plate.
«Oui, oui, mais la contrainte a des moments trop insupportables! James
est un maître d’une bonté terrible: il force ceux qui dépendent de lui à se
rendre compte de tout... il veut que l’on vive ainsi, pas autrement. Tout, à
ses yeux, se dessine en blanc et noir, clairement, tout devient évident. Il faut
avoir conscience de tout pour vivre bien. Ah! que de fois ai-je entendu cette
phrase! Vraiment, elle donne envie de vivre mal! Elle enlève à l’existence
tout son imprévu, tout son hasard, tout ce qui intéresse et qui amuse, tout ce
qui a du goût: la surprise qui fait sourire. Vous concevez bien que, parfois,
l’on veuille ignorer un peu le menu de son repas? Ici, chaque jour apporte
un devoir annoncé, une peine inscrite, comme à la table d’hôte en province,
où le vendredi apporte le plat de morue et de pommes de terre... Cela me
fait perdre l’appétit, même du plaisir!... Et maintenant, dites que je suis
folle, si vous voulez!»
Mathieu ne dit rien d’approchant. La dernière plainte de Mme Randal le
touchait: il s’en fallait de peu qu’il ne sympathisât.
«Non, Madame... Sachez seulement que vous avez, à Villedon, un
compatriote. Je parle votre langue et la gêne que vous ressentez n’a pour
moi rien de mystérieux. La liberté de l’esprit et du cœur me semble un bien
suprême; je conçois que l’on tâche d’y atteindre. A l’occasion, nous
reviendrons sur ce pénible sujet... Oui, reine d’une tribu d’étrangers,
astreinte à suivre les usages de la cour, vous ne cessez d’être en exil. En
somme, vous restez trop française.»
Elle ajouta d’une voix plus gaie:
«Et je vous ai bien dit, n’est-ce pas, que je me trouvais au Canada par le
hasard d’un engagement? Je suis née française, de parents français, à
Château-Thierry (Aisne). Plus tard, j’ai beaucoup, j’ai trop voyagé. Parfois,
je me sens un peu américaine.»
Elle conclut en riant:
«N’importe! le fond demeure, le fond... théodoricien!»
Mais ce rire sonnait faux.
On sortait du bois, l’ombre était moins épaisse, sur la prairie flottait

comme un reste de clarté confuse, des étoiles étincelaient au ciel dégagé de
brume. Alors Mme Randal revit la figure réelle de cet homme qui, par
occasion, avait reçu sa confidence, tandis qu’elle s’appuyait à son bras, et
de nouveau Mathieu aperçut le souple contour d’une femme auprès de lui...
Ils n’étaient plus seulement deux voix, sous les arbres obscurs. Ils ne
pouvaient parler ainsi davantage, ils se séparèrent, ils reprirent leurs
distances.
Puis Mathieu dit encore:
«Nous avons fait le tour du bois et sommes à quelques pas de chez vous,
Madame; permettez que je vous accompagne jusqu’au camp.
—Vous plaisantez! répliqua-t-elle. Je ne suis pas de ces personnes que
l’on accompagne ou que l’on met en voiture: non, non! je rentre par mes
propres moyens... Bonsoir, cher Monsieur; grâce à vous, j’ai fait une
excellente promenade.»
Ils se serrèrent la main par une prise vigoureuse et franche. Un instant
d’arrêt... peut-être pour se rendre bien compte du point où l’on se trouve...
«Amis?... tout de même? demanda-t-elle.
—Amis?... certes!»
Il était sincère.
XVI
Mathieu réfléchissait, assis devant la fenêtre ouverte de son bureau, mais
le spectacle d’une mer nuancée sur laquelle passaient de grands nuages ne
le touchait en rien: il s’occupait de lui-même.
Que ferait-il à Villedon, puisque son propos d’y rester était bien affermi?
Quelle y serait sa vie?—Le cirque ne figurait qu’une distraction de quelques
semaines et pourtant, seule, pensait-il, cette assemblée de gens étrangers par
leur race, leur culture, leur morale et leurs travaux, l’empêchait de
s’ennuyer. Demain, il s’ennuierait, à coup sûr, se sentant de nouveau maître
de ses champs et de ses bois, maître aussi de ses loisirs; demain, il se
trouverait en exil, chez lui.—Rentrer à Paris, il n’y songeait guère. Les gens
qu’il y fréquentait, ceux qu’il s’était habitué à voir, lui faisaient l’effet de
caricatures. Il ne pourrait plus supporter les papotages, les protestations et
les plaintes au sujet d’une robe, d’un souper mal servi, d’une femme de
chambre infidèle ou d’un vaudeville vraiment trop lugubre. Hélas! l’on ne
change pas son entourage comme l’on change de veston. Il y a la rue où
l’on se retrouve, le théâtre où l’on vous aperçoit, le restaurant où Nicole
s’installe par hasard à une table toute proche... Et l’on ne peut cependant
s’enfermer chez soi, se boucler, vivre comme en prison. La prison où l’on
se croit libre est assez rigoureuse déjà!
Mathieu souffre de cette incertitude; des souvenirs lui rendent son mal
plus cuisant. Eh quoi! une enfance orpheline, une jeunesse enfermée, une
adolescence étroite, sans joie, où quelques visites à un vieil oncle singulier
accentuaient encore sa détresse; quelques années de plaisir à Paris... qu’en
avait-il retenu? des grimaces, de petits calculs d’intérêt, de fausses larmes,
de faux serments qui ne prétendaient même pas à convaincre ni à toucher,
étant de passage, comme tout le reste. Lui serait-il donc défendu de goûter
au sel de la vie, à ce que la vie offre de grand et de sincère, à la belle amitié
avec un être qui vous comprend et vous ennoblit, au bel amour qui vous
élève toujours plus haut, qui dégage des nuées, qui rend limpide le ciel que
l’on porte en soi, et dont l’âme s’illumine?
Pourquoi ne pouvait-il toucher à ces fruits spirituels, à ces fleurs
secrètes? Pourquoi ne trouvait-il à portée de sa main que du rebut fait de
grappes gâtées et de corolles fausses?
Mathieu se posait la question, mais ne savait y répondre. Quant à ses
projets de voyage, il les avait écartés pour de bon: courir le monde
deviendrait vite un amusement de touriste; le voyage mieux entendu qu’il
rêvait naguère exigeait une préparation longue qu’il n’avait plus le courage
d’entreprendre; il était envahi de paresse... de quelle paresse étrange,
nouvelle, dont le goût lui semblait inconnu? Cela montait insidieusement,
comme ferait une peur sourde, cela l’écartait de toute action immédiate,
l’engageant à la remettre au lendemain, et surtout cela lui faisait un malaise,
une langueur inquiète, la stupeur que les bêtes ressentent prostrées sous
l’orage menaçant. Mais encore une fois, où trouver une raison à tout cela,
un allégement, un remède? et que faire en attendant?... Continuer
d’attendre?
XVII
Huit jours plus tard, Mathieu, monté sur Flea, le cheval étourneau de
Sam Harland, galopait joyeusement à travers les prés. L’air était encore vif
à cette heure matinale. On ne pouvait que se plaire à pareil exercice, sur une
herbe si fraîche et sous un ciel si pur. Leslie venait de passer qui menait des
bêtes à l’eau avec de grands gestes centauréens et des cris enthousiastes. Un
pantalon de toile bleue pour tout vêtement représentait encore une
concession absurde, à son avis, puisque l’on ne se sent soi-même que nu.
Harland avait fait de beaux essais de saut de barrière, et Plug tâchait de
tomber sans dommage, et le plus ridiculement possible, du dos de l’âne
qu’il enfourchait. D’autres écuyers s’entraînaient au lasso devant un
mannequin servant de but. Mathieu se contentait d’un galop modeste qui le
ravissait; de plus, il remarquait avec satisfaction que Flea, dont l’humeur
était ombrageuse, à l’ordinaire, et qui l’avait désarçonné plusieurs fois, lui
obéissait, maintenant, le mieux du monde. Le front dans le vent, il buvait
l’air, puis il fermait les yeux, un instant, pour goûter sa joie, et les rouvrait
pour reconnaître, alentour, l’herbe, le ciel, les bois et, là-bas, scintillante,
miroitante, déjà criblée de soleil, la mer.
Bientôt il aperçut Mme Randal coiffée d’un béret noir, culottée de noir, à
califourchon sur Mouse, sa jument grise. Elle portait une rose rouge à son
corsage: amazone habillée en adolescent, elle avait vraiment belle allure. Ils
se croisèrent, ils se saluèrent du geste et de la voix. Tout à coup, Mathieu se
ressouvint d’un vers lu jadis: «Contre le sein brûlé d’une antique
amazone...» Il se représenta Mme Randal tenant au poing, en place de cette
cravache inutile, un javelot, et le brandissant, mais l’image s’effaça vite
pour se proposer d’autre façon: une valkyrie qui foulerait des nuées... et
l’héroïque appel sonna à ses oreilles.
Mathieu s’arrêta net. Mme Randal faisait le tour de la prairie, au petit
galop, puis elle la traversa d’une allure plus vive, sauta plusieurs fois le
ruisseau, revint et frôla presque le cheval immobile. Mathieu en ressentit un
léger agacement, car elle n’avait plus tourné la tête; elle semblait tout
occupée de sa course et de cela seulement... Il admirait sa grâce, sa vigueur,
plus manifestes que jamais: cette danseuse se révélait écuyère étonnante, et
son costume peu féminin n’offrait pourtant, si crânement, si simplement
porté, rien de théâtral, malgré la touche de romantisme, et surtout rien
d’équivoque.
Flea piaffait, agacé lui aussi. Mme Randal acheva son tour. Que
n’invitait-elle Mathieu à la rejoindre?... Elle s’éloignait déjà. Il en eut un
surcroît de mauvaise humeur et, pour se justifier, inventa de mauvaises
raisons: ils galoperaient si bien de conserve! à rester seule, ainsi, Mme
Randal lui semblait faire de la parade, un numéro, un sketch d’équitation!
Pourquoi? pour le charmer? pour l’éblouir? Il n’avait nulle envie de
reprendre une promenade solitaire, de sentir la brise sur son front, sur ses
yeux... Mme Randal repassa encore... Subitement, Mathieu ne put se tenir
de toucher du talon le flanc jaune de Flea et de rendre la main.
Flea n’en demandait pas tant pour faire un beau partir en coupant la
prairie, même il dépassa Mouse et, comme l’on se trouvait sur la pente qui
menait à la mer, par prudence, Mathieu ne voulut pas l’arrêter trop court.
Bientôt il s’aperçut que Mme Randal en profitait: elle avait changé de
direction et remontait vers le village. Il la suivit, poussant Flea, l’excitant de
son mieux. Quand l’amazone vit ce cavalier à ses trousses, elle aussi entra
dans le jeu, et Mouse étant vaillante, Flea plus petit, moins robuste, moins
bien monté, fut gagné de vitesse. Course folle... Soudain, Mme Randal
tourna dans le bois et, le ruisseau franchi, disparut, entraînant Delannes
après elle. Quelques instants plus tard, il la revit, bricolant savamment entre
les arbres et les buissons. Mathieu se fatiguait, la tête perdue, les mains
nerveuses, grisé, non plus de vent et de vitesse, mais de chaude colère à
sentir que cette femme se moquait de lui. Il l’atteignit enfin. Elle avait sauté
à terre, sans aide, et caressait le museau de Mouse qui encensait doucement.
«Bonne course, n’est-ce pas?» dit-elle.
Et tout de suite elle ajouta sur un ton de reproche:
«Mais il ne faut pas trop demander à des chevaux délicats...»
Mathieu aurait voulu parler d’autre chose.
«Et «fort comme un cheval» est une expression absurde, indigne d’un
cavalier.»
Allait-il entendre un cours d’équitation sentimentale? Il avait mis pied à
terre aussi et se tenait près d’elle, encore essoufflé, toujours furieux. Il se
reprit un peu, pour la complimenter sur son art d’amazone; il dut le faire
habilement, car elle sourit, mais il se trouvait ridicule et en avait honte... Et
puis, surtout, il eût voulu savoir ce que pensait Mme Randal.

(Download PDF) Biology Exploring The Diversity of Life 4Th Canadian Edition Edition Peter Russell Full Chapter PDF

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

(Download PDF) Biology Exploring The Diversity of Life 4Th Canadian Edition Edition Peter Russell Full Chapter PDF

Uploaded by

Copyright:

Available Formats

Biology: Exploring the Diversity of Life

4th Canadian Edition Edition Peter

Biology: Exploring the Diversity of Life, Fifth

Biology : the unity and diversity of life 15th Edition

(eBook PDF) Campbell Biology, Third Canadian Edition

The Diversity of Fishes: Biology, Evolution and

Biology 12th Edition Peter H. Raven

Marine Biology 10th Edition Peter Castro

Marine Biology 12th Edition Peter Castro

Principles of Biology 4th Edition Robert J. Brooker

EXPLORING THE DIVERSITY OF LIFE

EXPLORING THE DIVERSITY OF LIFE

9780176718886_CVR_hr.indd 1 12/21/17 12:50 PM

EXPLORING THE DIVERSITY OF LIFE

With contributions by Ivona Mladenovic, Simon Fraser University

by Peter J. Russell, Paul E. Hertz,

VP, Product Solutions, K–20: Copy Editor: Cover Image:

tion biology, as well as field courses in the biology and

in lecture, laboratory, and tutorial settings. He led the

UNIT TWO GENES 149 THE CHEMICAL AND PHYSICAL FOUNDATIONS

2 The Cell: An Overview 25 6 Photosynthesis 125

15.1 Genomics: An Overview 362 SUMMARY ILLUSTRATION FOR CHAPTER 18 448

15.3 Annotation Identifies Genes 371 19.2 Phylogenetic Trees 456

24 Protists 559 27.12 Early Tetrapods and Modern Amphibians 696

28.12 Taking Action 740 31.6 Scale, Ecosystems, Species 833

UNIT SIX ECOLOGY AND BEHAVIOUR 747 32 Animal Behaviour 845

30.1 Introduction 778 32.13 Mates as Resources 868

Figure 30.16 Experimental Research Gause’s Experiments on

History of Earth F-42 37.2 Plant Chemical Defences 989

33 Organization of the Plant Body 885

34 Transport in Plants 913 SUMMARY ILLUSTRATION FOR CHAPTER 38 1024

36 Plant Nutrition 955 40.5 Transport of Gases in Blood 1069

36.2 Soil 960 SUMMARY ILLUSTRATION FOR CHAPTER 40 1074

36.3 Root Adaptations for Obtaining and Absorbing Nutrients 964

44 Animal Reproduction 1161 Appendix A: Answers to Self-Test Questions A-1

tional online exercise to further develop student quantitative

versions to help students learn the material.

A streamlined textbook … Emphasizing connections …

NEW material on DNA packaging

NEW figure clarifying replicated versus unreplicated

Chapter 11: DNA Structure, Replication, and

More precise description of fundamentals of

Chapter 5: Cellular Respiration Chapter 13: Regulation of Gene Expression

Chapter 14: DNA Technologies Chapter 20: Humans and Evolution

Archaea (Lokiarchaeota) that has a number of genes in

Chapter 17: Microevolution: Changes within Chapter 24: Protists

●● NEW discussion of the impact of wolf predation on popula-

●● Enhanced discussion of urban ecosystems

●● NEW material on the evolution of human language

and the axon hillock

b Engage, Adapt, and Master! Stay

. Adapt! Reinforce your knowledge of concepts by

m Engage! The learning path for each chapter begins

Test your mastery of concepts with

Assess your knowledge of chapter

there is no DNA synthesis. During

Regents of the University of California 2005/Dr. Uli Weier/Science Source

cell division is called chromosome Mitosis

ATP synthase (Figure 5.16).

inner mitochondrial membrane, from the matrix to the inter- Madeleinecentration

ETC (seewithin Figure 5.14). Within complexes I and IV, specific

energy from ATP to transport ions across membranes (the G in Gagainst

1 stands for gap,