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 Exploring the
Diversity of Life

Fifth Canadian Edition

Fenton • Maxwell • Haffie • Milsom • Nickle • Ellis • Riskin

Russell • Hertz • McMillan • Benington
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 Exploring the
Diversity of Life

Fifth Canadian Edition

M. Brock Fenton Bill Milsom

Western University University of British Columbia

Denis Maxwell Todd Nickle

Western University Mount Royal University

Tom Haffie Shona Ellis

Western University University of British Columbia

Shelby Riskin
University of Toronto

Peter J. Russell Paul E. Hertz Beverly McMillan Joel H. Benington

With contributions by Ivona Mladenovic, Simon Fraser University,

Jonathan Ferrier, Dalhousie University, Jeff Baker, University of Saskatchewan,
Jean Becker, University of Waterloo, and Erin Hodson, Wilfrid Laurier University

Australia • Brazil • Canada • Mexico • Singapore • United Kingdom • United States

11140_fm_ptg01.indd 1 21/03/22 4:13 PM

Biology: Exploring the Diversity of Life, © 2023, 2019 Cengage Learning Canada, Inc. ALL RIGHTS RESERVED.
Fifth Canadian Edition
M. Brock Fenton, Denis Maxwell, Tom
Adapted from Biology: The Dynamic Science, Fifth Edition, by Peter J. Russell, Paul E. Hertz,
Haffie, Bill Milsom, Todd Nickle, Shona
Beverly McMillan, and Joel H. Benington. Copyright © Cengage Learning, Inc., 2021.
Ellis, Shelby Riskin, Peter J. Russell, Paul
E. Hertz, Beverly McMillan, and Joel H. No part of this work covered by the copyright herein may be reproduced or distributed in
Benington any form or by any means, except as permitted by Canadian copyright law, without
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Library and Archives Canada Cataloguing in Publication
Production Service: MPS Limited
Title: Biology : exploring the diversity of life / M. Brock Fenton (Western University), Denis
Copy Editor: Frances Robinson Maxwell (Western University), Tom Haffie (Western University), Bill Milsom (University
Compositor: MPS Limited of British Columbia), Todd Nickle (Mount Royal University), Shona Ellis (University of
British Columbia), Shelby Riskin (University of Toronto), Peter J. Russell, Paul E. Hertz,
Art Director: Chris Doughman
Beverly McMillan, Joel H. Benington ; with contributions by Ivona Mladenovic (Simon
Illustrators: Dragonfly Media Group,
Fraser University) [and four others].
MPS Limited
Names: Russell, Peter J., author. | Fenton, M. Brock (Melville Brockett), 1943– author. |
Text Designer: Chris Doughman
Maxwell, Denis, author. | Haffie, Tom, author. | Milsom, Bill, 1947– author. | Nickle,
Cover Designer: Courtney Hellam
Todd, 1967–2021 author. | Ellis, Shona, 1962– author. | Riskin, Shelby, author.
Cover Image: Shona Ellis
Description: Fifth Canadian edition. | Issued also in 3 volumes. | Includes index.

Identifiers: C
 anadiana (print) 20210378492 | Canadiana (ebook) 2021037862X | ISBN
9780176911140 (hardcover) | ISBN 9780176911225 (PDF)

Subjects: LCSH: Biology—Textbooks. | LCGFT: Textbooks.

Classification: LCC QH308.2 .R88 2022 | DDC 570—dc23

ISBN: 978-0-17-691114-0

Ebook ISBN: 978-0-17-691122-5

Cengage
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Printed in the United States of America

Print Number: 01 Print Year: 2022
 For, and because of,
our generations of students.

11140_fm_ptg01.indd 3 21/03/22 4:13 PM

 In Memoriam: Dr. Todd Nickle
The entire Cengage Canada pro-
duction team is saddened, yet
honoured, to dedicate this fifth
Canadian edition of Biology:
Exploring the Diversity of Life
to our friend, colleague, and
coauthor Dr. Todd Nickle, as a
celebration of his life.
Before his death in August
of 2021, Todd completed his
revisions for this edition with
characteristic passion, dedica-
tion, and enthusiasm. He was
a particularly generous man,
Penny Nickle
quick with praise and quicker
with helpful support for col-
leagues and students. A very
early adopter of bleeding-edge educational technologies for learning
and assessment, Todd brought many innovative ideas to the table,
while always keeping student learning at the centre of our discussions.
If there was a new aspect of the project we wanted to explore, like The
Purple Pages videos or the COVID-19 supplement, Todd was always
happy to step up and mock up an example. He once apologized for the
birdsong on the audio track of one of his draft video commentaries
iv
11140_fm_ptg01.indd 4
on research figures. He had been working on the piece at his cabin in
the woods.
Always the good-hearted and playful provocateur, Todd had a knack for
engaging the author team in lively debate on important issues of student
experience. While he did not live to see publication of this edition, his leg-
acy and influence on this work will continue for many iterations to come.
Todd received his PhD from Oklahoma State University in 1998 and
taught biology at Mount Royal University, advocating active learning:
students come to class prepared to work with material rather than just
hear about it. Student preparation involves reading the text and apply-
ing the concepts to online exercises, the results of which inform what
the next lecture will be about. Class time focuses on exploring connec-
tions between concepts and ideas in biology and how they relate to other
disciplines. This inspired Todd to coauthor a handbook for first-year
science students, Science3. A compelling candidate for the 3M National
Fellowship, Todd’s work put him in the first cohort of full professors
at Mount Royal University in 2012. He also garnered the 2015 ACIFA
Innovation in Teaching Award and the 2016 Distinguished Faculty Award
from MRU. Todd’s interest in promoting best teaching practices among
educators beyond his home campus saw him expand and lead the Alberta
Introductory Biology Association (AIBA) to official society status in
Alberta as the Undergraduate Biology Educators of Alberta (UBEA). At
the national level, Todd also leaves many close colleagues in the Open
Consortium of Undergraduate Biology Educators (oCUBE).
21/03/22 4:13 PM
 About the Canadian Authors
M. B. (Brock) Fenton received his PhD from the
University of Toronto in 1969. Since then, he has been a
faculty member in biology at Carleton University, then
at York University, and then at Western University. In
addition to teaching parts of first-year biology, he has
also taught Vertebrate Biology, Animal Biology, and
M. B. Fenton
Conservation Biology, as well as field courses in the
biology and behaviour of bats. Brock has received awards
for his teaching (Carleton University Faculty of Science
Teaching Award; Ontario Confederation of University
Faculty Associations Teaching Award; and a 3M Teaching
Fellowship, Society for Teaching and Learning in Higher
Education) in addition to recognition of his work on public
awareness of science (Gordin Kaplan Award from the
Denis Maxwell received his PhD from the University
of Western Ontario in 1995. His thesis focused on
photosynthetic acclimation in green algae. Following
his doctorate, he undertook postdoctoral training at the
Department of Energy Plant Research Laboratory at
Denis Maxwell
Michigan State University, where he studied the function
of the mitochondrial alternative oxidase. After taking up
a faculty position at the University of New Brunswick in
Tom Haffie is a graduate of the University of Guelph
and the University of Saskatchewan in the study of
microbial genetics. Newly retired, he devoted his 33-year
Mitch Zimmer
career at Western University to teaching large biology
classes in lecture, laboratory, and tutorial settings. He
led the development of the innovative core laboratory
course in the Biology program; he was an early adopter
of computer animation in lectures; and, most recently,
he led a deep blended redevelopment of introductory
biology informed by a students-as-partners approach to
collaborative course design. Tom was a founding force
in the Western Biology Undergraduate Society (BUGS),
Bill Milsom received his PhD from the University
of British Columbia and is a professor in UBC’s
Department of Zoology, where he has taught a variety
of courses, including first-year biology, for almost 40
Bruce Moffat
years. His research interests include the evolutionary
origins of respiratory processes and the adaptive
changes in these processes that allow animals to exploit
diverse environments. He examines respiratory and
cardiovascular adaptations in vertebrate animals in
rest, sleep, exercise, altitude, dormancy, hibernation,
diving, and so on. This research contributes to our
understanding of the mechanistic basis of biodiversity
11140_fm_ptg01.indd 5
Canadian Federation of Biological Societies; Honorary Life
Membership, Science North, Sudbury, Ontario; Canadian
Council of University Biology Chairs Distinguished
Canadian Biologist Award; The McNeil Medal for the
Public Awareness of Science of the Royal Society of Canada;
and the Sir Sandford Fleming Medal for Public Awareness
of Science, the Royal Canadian Institute). He also received
the C. Hart Merriam Award from the American Society
of Mammalogists for excellence in scientific research. Bats
and their biology, behaviour, evolution, and echolocation
are the topics of his research, which has been funded by
the Natural Sciences and Engineering Research Council of
Canada (NSERC). In November 2014, Brock was inducted
as a Fellow of the Royal Society of Canada.
2000, Denis moved in 2003 to the Department of Biology
at Western University. He served as Associate Chair for
Undergraduate Education for the Department of Biology
from 2009 to 2016. Currently, he is Assistant Dean for
the Faculty of Science, with a portfolio that includes
Recruitment and First-Year Studies and outreach. Denis
has taught first-year Biology to over 15 000 students, most
of the time with coauthor Tom Haffie.
the Open Consortium for Undergraduate Biology
Educators (oCUBE), and the Western Conference on
Science Education (WCSE). Tom’s educational practice
was honoured with several awards, including a Western
University Students’ Council Award for Excellence
in Teaching, a Western University Edward G. Pleva
Award for Excellence in Teaching, a Western University
Fellowship in Teaching Innovation, a Western University
Teaching Fellowship for Science, a Province of Ontario
Award for Leadership in Faculty Teaching (LIFT), and
a 3M National Teaching Fellowship for excellence in
teaching.
and the physiological costs of habitat selection. Bill’s
research has been funded by NSERC, and he has
received several academic awards and distinctions,
including the Fry Medal of the Canadian Society of
Zoologists, the August Krogh Distinguished Lectureship
Award of the American Physiological Society, the
Bidder Lecture of the Society for Experimental Biology,
and the Izaak Walton Killam Award for Excellence
in Mentoring. He has served as President of the
Canadian Society of Zoologists and as President of the
International Congress of Comparative Physiology and
Biochemistry.
21/03/22 4:13 PM
 Shona Ellis received her MSc from the University
of British Columbia and is a professor of teaching in
the Botany Department. She developed a keen interest
in forests and oceans growing up on the central coast
of British Columbia in Heiltsuk Territory. As an
undergraduate, Shona pursued her interests in botany
and entomology. Her MSc research incorporated tissue
culture, phytochemistry, and plant anatomy. She realized
a passion for teaching and joined the faculty at UBC in
Andy Cotton
1998. Shona teaches botany courses that include vascular
plants, economic botany, bryology, and plant systematics,
as well as Introductory Biology. She also currently teaches
Shelby Riskin received her BA from Grinnell College
in Iowa, where she was first exposed to the wonderful
world of biological research. She received her PhD from
Brown University in a joint program with the Marine
Biological Laboratory in Woods Hole, Massachusetts. Her
dissertation focused on ecosystem-level biogeochemical
consequences of conversion to agriculture in the Brazilian
Dan Riskin
Amazon, the region of the world where deforestation and
conversion to large-scale farming are happening most
rapidly. Shelby then joined the University of Toronto as
vi A B O U T T H E C A N A D I A N AU T H O R S
11140_fm_ptg01.indd 6
a course through the Haida Gwaii Institute that takes
her and her class into the natural environment of Haida
Territory. Shona teaches in a number of settings; in large
and small lecture classes, in laboratories, and on field
trips. While she feels the best classroom is outdoors,
she integrates online technologies into all her courses;
she is an early adopter of online teaching and learning
resources. Shona has received two Killam Teaching
Awards, a Dean of Science Service Award (UBC), UBC’s
Margaret Fulton Award for student development, and
the Charles Edwin Bessey Teaching Award from the
Botanical Society of America.
Assistant Professor in the teaching stream of the Ecology
and Evolutionary Biology Department. She teaches a
variety of undergraduate courses exploring ecosystem
ecology and conservation biology and is Director of the
U of T National Biology Competition, an international
competition for high school students. Shelby is also
passionate about mentoring undergraduates through
independent research projects and about teaching biology
and ecology to students outside the life sciences—
understanding the diversity of life is for everyone.
21/03/22 4:13 PM
 About the US Authors
Peter J. Russell received his BSc in biology from the
University of Sussex, England, in 1968 and his PhD in
genetics from Cornell University in 1972. He has been
a member of the Biology faculty of Reed College since
Courtesy of Peter J. Russell
1972 and is currently Professor of Biology, Emeritus.
Peter taught a section of the introductory biology course,
a genetics course, and a research literature course on
molecular virology. In 1987, he received the Burlington
Northern Faculty Achievement Award from Reed
College in recognition of his excellence in teaching. Since
1986, he has been the author of a successful genetics
textbook; the current edition is iGenetics: A Molecular
Approach. Peter’s research was in the area of molecular
genetics, with a specific interest in characterizing the role
of host genes in the replication of the RNA genome of a
pathogenic plant virus and the expression of the genes of
Paul E. Hertz was born and raised in New York City.
He received a BA in biology from Stanford University
in 1972, a master’s degree in biology from Harvard
University in 1973, and a PhD in biology from Harvard
University in 1977. While completing field research for
Courtesy of Aaron Kinard
his doctorate, he served on the Biology faculty of the
University of Puerto Rico at Rio Piedras. After spending
two years as an Isaac Walton Killam Postdoctoral Fellow
at Dalhousie University, Paul accepted a teaching position
at Barnard College, where he has taught since 1979. He
was named Ann Whitney Olin Professor of Biology in
2000 and the Claire Tow Professor of Biology in 2016. He
received The Barnard Award for Excellence in Teaching
in 2007. In addition to serving on numerous college
committees, Paul chaired Barnard’s Biology Department
for eight years and served as Acting Provost and Dean of
the Faculty from 2011 to 2012. He was also the founding
Program Director of the Hughes Science Pipeline Project
at Barnard, an undergraduate curriculum and research
program funded continuously by the Howard Hughes
Beverly McMillan holds undergraduate and graduate
degrees from the University of California, Berkeley. She
has worked extensively in educational and commercial
publishing as an author, science writer, project manager,
Courtesy of Beverly McMillan
and multimedia content developer. In addition to her
contributions to college textbooks, Beverly has written or
coauthored multiple popular books on topics in natural
11140_fm_ptg01.indd 7
the virus; yeast was used as the model host. His research
has been funded by agencies such as the National
Institutes of Health, the National Science Foundation, the
American Cancer Society, the Department of Defense,
the Medical Research Foundation of Oregon, and the
Murdoch Foundation. He has published his research
in a variety of journals, including Genetics, Journal of
Bacteriology, Molecular and General Genetics, Nucleic
Acids Research, Plasmid, and Molecular and Cellular
Biology. Peter has a long history of encouraging faculty
research involving undergraduates, including cofounding
the biology division of the Council on Undergraduate
Research in 1985. He was Principal Investigator/Program
Director of a National Science Foundation Award for the
Integration of Research and Education (NSF–AIRE) to
Reed College, 1998 to 2002.
Medical Institute, from 1992 to 2016. The Pipeline Project
included the Intercollegiate Partnership, a program for
local community-college students that facilitated their
transfer to four-year colleges and universities. Since
2016, he has served as Director of the Science Pathways
Scholars Program, which provides support and research
opportunities to first-generation college students and
students of colour at Barnard. He teaches one semester
of the introductory sequence for biology majors and
pre-professional students, lecture and laboratory courses
in vertebrate zoology and ecology, and seminars that
introduce first-year students to scientific research. Paul is
an animal physiological ecologist with a specific research
interest in the thermal biology of lizards. He has conducted
fieldwork in the West Indies since the mid-1970s, most
recently focusing on the lizards of Cuba and Puerto
Rico. His work has been funded by the National Science
Foundation, and he has published his research in such
prestigious journals as The American Naturalist, Ecology,
Nature, Oecologia, and The Proceedings of the Royal Society.
history and human health and biology, as well as field guides
to the flora and fauna of more than 20 US states. She has
also created Web and print content for such clients as the US
National Park Service, the Science Museum of Virginia, The
Mariners’ Museum, the San Francisco Exploratorium, the
University of California system, and the Virginia Institute of
Marine Science/College of William and Mary.
vii
21/03/22 4:13 PM

Courtesy of Joel H. Benington
viii
11140_fm_ptg01.indd 8
Joel H. Benington received his BA from St. John’s
College in Annapolis, Maryland, in 1985 and his PhD in
biology from Stanford University in 1992. He performed
postdoctoral research at the University of Los Angeles
and Stanford University until 1996. Since then, he has
been a member of the Biology faculty of St. Bonaventure
University, where he is currently Professor of Biology
and Director of the Bioinformatics and Health and
Society programs. He has twice served as Chair of the
Department of Biology. During his entire time at St.
Bonaventure University, he has taught one or both
semesters of the general biology sequence for first-
year life science majors. He also teaches upper-level
A B O U T T H E U S AU T H O R S
Neurobiology, Genomics, and Evolution courses and
has led a variety of seminar courses in the university’s
Honors Program. He has published his research in
journals such as Progress in Neurobiology, Brain Research,
The American Journal of Physiology, and The Scientist.
In addition to laboratory research, he has published
hypotheses concerning the role of sleep in brain energy
metabolism, the functional relationship between REM
sleep and non-REM sleep, and connections between
sleep and learning. Joel’s research has been funded by
the National Institutes of Health, and he has served as
Principal Investigator of a National Grid grant to support
K–12 STEM education in Cattaraugus County, New York.
21/03/22 4:13 PM
 Brief Contents
VOLUME 1: BIOLOGY OF THE CELL 1 UNIT 6 ECOLOGY AND BEHAVIOUR
1 Defining Life and Its Origins 5 26 Population Ecology 689
27 Species Interactions and Community Ecology 717
UNIT 1 SYSTEMS AND PROCESSES—THE CELL
28 Ecosystems 753
2 The Cell: An Overview 31 29 Conservation of Biodiversity 779
3 Energy and Enzymes 59
4 Cell Membranes and Signalling 83 THE CHEMICAL AND PHYSICAL FOUNDATIONS OF
5 Cellular Respiration 109 BIOLOGY (THE PURPLE PAGES) F-1
6 Photosynthesis 135
VOLUME 3: SYSTEMS AND PROCESSES 801
UNIT 2 GENES
7 Cell Cycles 161 UNIT 7 SYSTEMS AND PROCESSES—PLANTS
8 Genetic Recombination 185 30 Organization of the Plant Body 805
9 The Chromosomal Basis of Inheritance 211 31 Transport in Plants 833
10 Genetic Linkage, Sex Linkage, and Other Extensions 32 Reproduction and Development in Flowering
to Basic Inheritance Mechanisms 237 Plants 853
33 Plant Nutrition 875
UNIT 3 DNA AND GENE EXPRESSION 34 Plant Signals and Responses to the Environment 895
11 DNA Structure, Replication, and Repair 265
12 Gene Structure, Expression, and Mutation 291 UNIT 8 SYSTEMS AND PROCESSES—ANIMALS
13 Regulation of Gene Expression 321 35 Introduction to Animal Organization and
14 DNA Technologies 347 Physiology 925
15 Genomes 375 36 Animal Nutrition 947
37 Gas Exchange: The Respiratory System 977
VOLUME 2: EVOLUTION, ECOLOGY, AND THE 38 Internal Transport: The Circulatory System 1001
DIVERSITY OF LIFE 403 39 Regulation of the Internal Environment: Water, Solutes,
and Temperature 1027
UNIT 4 EVOLUTION AND CLASSIFICATION 40 Control of Animal Processes: Endocrine Control 1059
16 Evolution: The Development of the Theory 407 41 Animal Reproduction and Development 1085
17 Microevolution: Changes within Populations 429 42 Control of Animal Processes: Neural Control 1123
18 Speciation and Macroevolution 451 43 Muscles, Skeletons, and Body Movements 1179
19 Systematics and Phylogenetics: Revealing the Tree 44 Animal Behaviour and Responses to the
of Life 471 Environment 1201
45 Defences against Disease 1237
UNIT 5 THE DIVERSITY OF LIFE
20 Bacteria and Archaea 497 APPENDIX A: ANSWERS TO SELF-TEST
21 Protists 519
QUESTIONS A-1
GLOSSARY G-1
22 Fungi 547
INDEX I-1
23 Plants 571
24 Animals 603
25 Viruses, Viroids, and Prions: Infectious Biological
Particles 671

ix

11140_fm_ptg01.indd 9 21/03/22 4:13 PM

 Contents 4 Cell Membranes and Signalling 83
4.1 An Overview of the Structure of Membranes 84
Figure 4.2 Experimental Research The Frye–Edidin Experiment
Demonstrating That the Phospholipid Bilayer Is Fluid 85

In Memoriam: Dr. Todd Nickle iv Figure 4.3 Research Method Freeze Fracture 86

About the Canadian Authors v 4.2 The Lipid Fabric of a Membrane 86

About the US Authors vii 4.3 Membrane Proteins 89
Letter to Students xvii 4.4 Passive Membrane Transport 91

New to This Edition xxi

4.5 Active Membrane Transport 95
4.6 Exocytosis and Endocytosis 98
Welcome to Biology: Exploring the Diversity
of Life, 5Ce xxiv 4.7 The Role of Membranes in Cell Signalling 100
SUMMARY ILLUSTRATION FOR CHAPTER 4 104
Active Learning xxvi

Student and Instructor Resources xxx 5 Cellular Respiration 109

Acknowledgments xxxi 5.1 The Chemical Basis of Cellular Respiration 110

5.2 Cellular Respiration: An Overview 112

VOLUME 1: BIOLOGY OF THE CELL 1

5.3 Glycolysis: The Splitting of Glucose 113
5.4 Pyruvate Oxidation and the Citric Acid Cycle 116
1 Defining Life and Its Origins 5
5.5 Oxidative Phosphorylation: Electron Transport
1.1 What Is Life? 6
and Chemiosmosis 119
1.2 The Chemical Origins of Life 8
5.6 The Efficiency and Regulation of Cellular Respiration 122
1.3 The Evolution of Information Flow: RNA, DNA, Protein 11
5.7 Oxygen and Cellular Respiration 125
1.4. The Development of Metabolism and the First Cells 13
SUMMARY ILLUSTRATION FOR CHAPTER 6 130
1.5 The Tree of Life 16
1.6 Eukaryotes and the Rise of Multicellularity 19 6 Photosynthesis 135

1.7 The Fossil Record 24 6.1 The Physical Nature of Light 136

UNIT 1 SYSTEMS AND PROCESSES—THE CELL 6.2 Photosynthesis: An Overview 138

6.3 The Photosynthetic Apparatus 140
2 The Cell: An Overview 31 6.4 The Light Reactions 144
2.1 Basic Features of Cell Structure and Function 32 6.5 The Calvin Cycle 146
2.2 Prokaryotic Cells 36 6.6 Photorespiration and CO2-Concentrating Mechanisms 149
2.3 Eukaryotic Cells 37 6.7 Photosynthesis and Cellular Respiration Compared 154
Figure 2.8 Research Method Cell Fractionation 38 SUMMARY ILLUSTRATION FOR CHAPTER 5 156
2.4 Specialized Structures of Plant Cells 49
UNIT 2 GENES
2.5 The Animal Cell Surface 51
SUMMARY ILLUSTRATION FOR CHAPTER 2 54 7 Cell Cycles 161
7.1 The Cycle of Cell Growth and Division: An Overview 162
3 Energy and Enzymes 59
7.2 The Cell Cycle in Prokaryotic Organisms 163
3.1 Energy and the Laws of Thermodynamics 60
7.3 Mitosis and the Eukaryotic Cell Cycle 164
3.2 Free Energy and Spontaneous Processes 63
7.4 Formation and Action of the Mitotic Spindle 172
3.3 Thermodynamics and Life 66
7.5 Cell Cycle Regulation 174
3.4 Overview of Metabolism 67
Figure 7.19 Experimental Research Movement of Chromosomes
3.5 The Role of Enzymes in Biological Reactions 70 during Anaphase of Mitosis 176
3.6 Factors That Affect Enzyme Activity 74
SUMMARY ILLUSTRATION FOR CHAPTER 7 180
SUMMARY ILLUSTRATION FOR CHAPTER 3 78

11140_fm_ptg01.indd 10 21/03/22 4:13 PM

 8 Genetic Recombination 185 Figure 11.2 Experimental Research The Hershey and Chase
8.1 Mechanism of Genetic Recombination 186 Experiment Demonstrating That DNA Is the Hereditary Molecule 268
8.2 Genetic Recombination in Bacteria 187 11.2 DNA Structure 269
Figure 8.2 Research Method Replica Plating 188 11.3 DNA Replication 272

Figure 8.3 Experimental Research Genetic Recombination in Figure 11.9 Experimental Research The Meselson and Stahl
Bacteria 189 Experiment Demonstrating the Semiconservative Model for DNA
Replication to Be Correct 274
8.3 Genetic Recombination Occurs in Eukaryotes during
Meiosis 195 11.4 Repair of Damage in DNA 283

SUMMARY ILLUSTRATION FOR CHAPTER 8 206 SUMMARY ILLUSTRATION FOR CHAPTER 11 286

9 The Chromosomal Basis of Inheritance 211 12 Gene Structure, Expression, and Mutation 291

9.1 Mendel’s Experiments with Garden Peas 212 12.1 The Connection between DNA, RNA, and Protein 292

Figure 9.2 Research Method Making a Genetic Cross between Two Figure 12.2 Experimental Research The Gene–Enzyme
Pea Plants 213 Relationship 294

Figure 9.4 Experimental Research The Principle of Segregation: 12.2 Transcription: DNA-Directed RNA Synthesis 297
Inheritance of Flower Colour in Garden Peas 216 12.3 Processing of mRNAs in Eukaryotes 299
12.4 Translation: mRNA-Directed Polypeptide Synthesis 303
Figure 9.7 Experimental Research Testing the Predicted Outcomes
12.5 Mutations Can Affect Protein Structure and Function 313
of Genetic Crosses 220
SUMMARY ILLUSTRATION FOR CHAPTER 12 316
Figure 9.8 Experimental Research The Principle of Independent
Assortment 221 13 Regulation of Gene Expression 321
13.1 Regulation of Gene Expression in Prokaryotic Cells 322
9.2 Later Modifications and Additions to Mendel’s Hypotheses 224
13.2 Regulation of Transcription in Eukaryotes 328
Figure 9.12 Experimental Research Experiment Showing
13.3 Posttranscriptional, Translational, and Posttranslational
Incomplete Dominance of a Trait 225
Regulation 335
SUMMARY ILLUSTRATION FOR CHAPTER 9 232 13.4 The Loss of Regulatory Controls in Cancer 339
SUMMARY ILLUSTRATION FOR CHAPTER 13 342
10 Genetic Linkage, Sex Linkage, and Other Extensions
to Basic Inheritance Mechanisms 237
14 DNA Technologies 347
10.1 Genetic Linkage and Recombination 238
14.1 DNA Cloning 348
Figure 10.2 Experimental Research Evidence for Gene Linkage 240
Figure 14.3 Research Method Identifying a Recombinant Plasmid
10.2 Sex-Linked Genes 243 Containing a Gene of Interest 351
Figure 10.8 Experimental Research Evidence for Sex-Linked Figure 14.4 Research Method Synthesis of DNA from mRNA Using
Genes 246 Reverse Transcriptase 352
Figure 14.5 Research Method The Polymerase Chain Reaction
10.3 Chromosomal Mutations That Affect Inheritance 248
(PCR) 353
10.4 Human Genetic Traits, Pedigree Analysis, and
Genetic Counselling 252 Figure 14.6 Research Method Separation of DNA Fragments by
Agarose Gel Electrophoresis 354
10.5 Additional Patterns of Inheritance 256
SUMMARY ILLUSTRATION FOR CHAPTER 10 260 14.2 Applications of DNA Technologies 355
Figure 14.8 Research Method Southern Blot Analysis 357
UNIT 3 DNA AND GENE EXPRESSION
Figure 14.11 Research Method Making a Knockout Mouse 361
11 DNA Structure, Replication, and Repair 265 Figure 14.14 Experimental Research The First Cloning of a
11.1 Establishing DNA as the Hereditary Molecule 266 Mammal 364
Figure 11.1 Experimental Research Griffith’s Experiment with Figure 14.16 Research Method Using the Ti Plasmid of Rhizobium
Virulent and Nonvirulent Strains of Streptococcus pneumoniae 267 radiobacter to Produce Transgenic Plants 366

CONTENTS xi

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 SUMMARY ILLUSTRATION FOR CHAPTER 14 370 18.4 Genetic Mechanisms of Speciation 461
Figure 18.16 Observational Research Chromosomal Similarities and
15 Genomes 375
Differences among Humans and the Great Apes 465
15.1 Genomics: An Overview 376
15.2 Genome Sequencing 377 SUMMARY ILLUSTRATION FOR CHAPTER 18 466

Figure 15.1 Research Method Sanger Sequencing 378 19 Systematics and Phylogenetics: Revealing the
Tree of Life 471
Figure 15.2 Research Method Pyrosequencing 382
19.1 Nomenclature and Classification 472
15.3 Annotation Identifies Genes 385
19.2 Phylogenetic Trees 474
Figure 15.8 Research Method Analysis of Gene Expression Levels 19.3 Sources of Data for Phylogenetic Analyses 477
using RNA-seq 390
19.4 Traditional Classification and Paraphyletic Groups 480
15.4 Comparative Genomics Can Reveal How Genes and 19.5 The Cladistic Revolution 481
Genomes Evolved 391
Figure 19.11 Research Method Using Cladistics to Construct a
SUMMARY ILLUSTRATION FOR CHAPTER 15 400 Phylogenetic Tree 483

19.6 Phylogenetic Trees as Research Tools 486

VOLUME 2: EVOLUTION, ECOLOGY, AND Figure 19.13 Research Method Using Genetic Distances to
THE DIVERSITY OF LIFE 403 Construct a Phylogenetic Tree 487
UNIT 4 EVOLUTION AND CLASSIFICATION 19.7 Molecular Phylogenetic Analyses 489

16 Evolution: The Development of the Theory 407 SUMMARY ILLUSTRATION FOR CHAPTER 19 490

16.1 What Is Evolution through Natural Selection? 408 UNIT 5 THE DIVERSITY OF LIFE
16.2 Evidence for Evolution through Natural Selection 411
16.3 Development of the Theory of Evolution 414 20 Bacteria and Archaea 497
20.1 The Full Extent of the Diversity of Bacteria and Archaea
Figure 16.9 Experimental Research Adaptation of E. coli to a Change
Is Unknown 498
in Temperature 415
20.2 Prokaryotic Structure and Function 498
16.4 Evolutionary Theory since Darwin 421
Figure 20.5 Experimental Research Genetic Recombination in
SUMMARY ILLUSTRATION FOR CHAPTER 16 424 Bacteria 501

17 Microevolution: Changes within Populations 429 20.3 The Domain Bacteria 508
17.1 Variation in Natural Populations 430 20.4 The Domain Archaea 510
17.2 Population Genetics 432 SUMMARY ILLUSTRATION FOR CHAPTER 20 514
17.3 The Agents of Microevolution 434
21 Protists 519
Figure 17.8 Research Method Using the Hardy–Weinberg
21.1 The Vast Majority of Eukaryotes Are Protists 520
Principle 435
21.2 Characteristics of Protists 521
Figure 17.13 Experimental Research Do Humans Experience
21.3 Protists’ Diversity Is Reflected in Their Habitats, Structure,
Stabilizing Selection? 440
Metabolism, and Reproduction 522
17.4 Non-random Mating 441
21.4 Eukaryotic Supergroups and Key Protist Lineages 523
Figure 17.16 Experimental Research Sexual Selection in
Figure 21.8 Observational Research Isolation and Identification
Action 443
of Marine Diplonemids, Potentially the Most Abundant Marine
17.5 Maintaining Genetic and Phenotypic Variation 444 Organism 525
SUMMARY ILLUSTRATION FOR CHAPTER 17 446 21.5 Some Protist Lineages Arose from Primary Endosymbiosis
and Others from Secondary Endosymbiosis 539
18 Speciation and Macroevolution 451
SUMMARY ILLUSTRATION FOR CHAPTER 21 542
18.1 What Is a Species? 452
18.2 Maintaining Reproductive Isolation 455 22 Fungi 547
18.3 The Geography of Speciation 458 22.1 General Characteristics of Fungi 548

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 22.2 Evolution and Diversity of Fungi 550 26.5 Models of Population Growth 697
22.3 Fungal Life Styles 561 26.6 Population Regulation 702
Figure 22.20 Experimental Research Hidden Third Partner in Lichen Figure 26.16 Experimental Research Evaluating Density-Dependent
Symbiosis 562 Interactions between Species 704

SUMMARY ILLUSTRATION FOR CHAPTER 22 566 26.7 Human Population Growth 707
SUMMARY ILLUSTRATION FOR CHAPTER 26 712
23 Plants 571
23.1 Defining Characteristics of Land Plants 572 27 Species Interactions and Community Ecology 717
23.2 The Transition to Life on Land 573 27.1 Population Interactions Shape Communities 718
23.3 Bryophytes: Nonvascular Land Plants 579 27.2 Symbioses: Close Associations 719
23.4 Seedless Vascular Plants 583 27.3 Energy Intake and Exchange 721
23.5 Gymnosperms: The First Seed Plants 588 27.4 Defence 723
23.6 Angiosperms: Flowering Plants 593 27.5 Competition 727
Figure 23.30 Experimental Research Exploring a Possible Early Figure 27.15 Experimental Research Gause’s Experiments on
Angiosperm Adaptation for Efficient Photosynthesis in Dim Interspecific Competition in Paramecium 728
Environments 595 Figure 27.18 Experimental Research Demonstration of Competition
between Two Species of Barnacles 730
SUMMARY ILLUSTRATION FOR CHAPTER 23 598
Figure 27.19 Experimental Research The Complex Effects of a
24 Animals 603 Herbivorous Snail on Algal Species Richness 731
24.1 What Are Animals? 604
27.6 The Nature of Ecological Communities 732
24.2 Animal Origins: Animals Probably Arose from a Colonial
27.7 Community Characteristics 734
Flagellate 605
27.8 Effects of Population Interactions on Community
24.3 Key Features Used to Classify Animals 606
Structure 738
24.4 Basal Phyla 611
27.9 Succession 740
24.5 The Protostomes 619
27.10 Variations in Species Richness among Communities 743
24.6 The Deuterostomes 638
SUMMARY ILLUSTRATION FOR CHAPTER 27 748
SUMMARY ILLUSTRATION FOR CHAPTER 24 666
28 Ecosystems 753
25 Viruses, Viroids, and Prions: Infectious Biological
28.1 Ecosystems and Energy 754
Particles 671
Figure 28.12 Experimental Research A Trophic Cascade in Salt
25.1 What Is a Virus? Characteristics of Viruses 672
Marshes 763
25.2 Viruses Infect Bacterial, Animal, and Plant Cells by Similar
Pathways 675 28.2 Nutrient Cycling in Ecosystems 764
25.3 Treating and Preventing Viral Infections 680 28.3 Anthropogenic Global Change 771
25.4 Virotherapy: Using Viruses to Cure Disease 681 SUMMARY ILLUSTRATION FOR CHAPTER 28 774
25.5 Viruses May Have Evolved from Fragments of Cellular
DNA or RNA 681 29 Conservation of Biodiversity 779

25.6 Viroids and Prions Are Infective Agents Even Simpler in 29.1 The Anthropocene 780
Structure than Viruses 682 29.2 Threats to Biodiversity 781
SUMMARY ILLUSTRATION FOR CHAPTER 25 684 Figure 29.3 Observational Research Near-Complete Extinction of
Small Mammals in Tropical Forest Fragments 783
UNIT 6 ECOLOGY AND BEHAVIOUR
29.3 Ecosystem Services Highlight the Benefits of Ecosystems 787
26 Population Ecology 689
29.4 Conservation Biology: Principles and Theory 789
26.1 Introduction 690
Figure 29.16 Experimental Research Effect of Landscape Corridors
26.2 Population Characteristics 690 on Plant Species Richness in Habitat Fragments 793
26.3 Demography 693
29.5 Assessing Threatened Species and Prioritizing Efforts 793
26.4 Evolution of Life Histories 695

CONTENTS xiii

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 29.6 Protecting Biodiversity Requires a Diversity of Solutions 794 34 Plant Signals and Responses to the Environment 895
SUMMARY ILLUSTRATION FOR CHAPTER 29 798 34.1 Plant Hormones 896
Figure 34.3 Experimental Research The Darwins’ Experiments on
Phototropism 899
THE CHEMICAL AND PHYSICAL FOUNDATIONS OF
BIOLOGY (THE PURPLE PAGES) F-1 Figure 34.4 Experimental Research Two Experiments by Frits
Went Demonstrating the Effect of Indoleacetic Acid (IAA) on an Oat
Coleoptile 900
VOLUME 3: SYSTEMS AND PROCESSES 801
UNIT 7 SYSTEMS AND PROCESSES—PLANTS 34.2 Plant Chemical Defences 907
34.3 Plant Movements 911
30 Organization of the Plant Body 805 34.4 Plant Biological Clocks 915
30.1 Plant Structure and Growth: An Overview 806 34.5 Learned Behaviour in Plants 919
30.2 The Three Plant Tissue Systems 810
SUMMARY ILLUSTRATION FOR CHAPTER 34 920
Figure 30.9 Experimental Research Networking the Secondary
Cell Wall 813 UNIT 8 SYSTEMS AND PROCESSES—ANIMALS
30.3 Primary Shoot Structure 815 35 Introduction to Animal Organization and
30.4 Primary Root Structure 820 Physiology 925
30.5 Secondary Growth 823 35.1 Organization of the Animal Body 926

SUMMARY ILLUSTRATION FOR CHAPTER 30 828 35.2 Animal Tissues 928

35.3 Coordination of Tissues in Organs and Organ Systems 935
31 Transport in Plants 833 35.4 Homeostasis 935
31.1 Principles of Water and Solute Movement in Plants 834 Figure 35.12 Experimental Research Demonstration of the Use of
31.2 Uptake and Transport of Water and Solutes by Roots 837 the Bill for Thermoregulation in Birds 939
31.3 Long-Distance Transport of Water and Minerals
in the Xylem 839 SUMMARY ILLUSTRATION FOR CHAPTER 35 942

31.4 Transport of Organic Substances in the Phloem 844 36 Animal Nutrition 947
Figure 31.12 Experimental Research Translocation Pressure 845 36.1 Nutrients Are Essential Components of Any Diet 948
SUMMARY ILLUSTRATION FOR CHAPTER 31 848 36.2 Feeding to Obtain Nutrients 952
36.3 Digestive Processes 954
32 Reproduction and Development in Flowering 36.4 Structure and Function of the Mammalian Digestive
Plants 853 System 957
32.1 Overview of Flowering Plant Reproduction 854 36.5 Regulation of Digestive Processes 967
32.2 Flower Structure and Formation of Gametes 856
Figure 36.20 Experimental Research Association of Bacterial
32.3 Pollination, Fertilization, and Germination 860 Populations in the Gut Microbiome with Obesity in Humans 969
32.4 Asexual Reproduction in Flowering Plants 866
36.6 Reflections on the Chapter 971
Figure 32.16 Research Method Plant Tissue Culture
SUMMARY ILLUSTRATION FOR CHAPTER 36 972
Protocol 868

32.5 Early Development of Plant Form and Function 868 37 Gas Exchange: The Respiratory System 977

SUMMARY ILLUSTRATION FOR CHAPTER 32 870 37.1 General Principles 979

37.2 Ventilation of the Gas-Exchange Organs 983
33 Plant Nutrition 875
37.3 The Mammalian Respiratory System 987
33.1 Plant Nutritional Requirements 876 37.4 Exchange of Gas with Blood 989
Figure 33.2 Research Method Hydroponic Culture 877 37.5 Transport of Gases in Blood 990
33.2 Soil 880 37.6 Reflections on the Chapter 994
33.3 Root Adaptations for Obtaining and Absorbing Nutrients 883 Figure 37.21 Experimental Research Demonstration of a Molecular
SUMMARY ILLUSTRATION FOR CHAPTER 33 890 Basis for High-Altitude Adaptation in Deer Mice 995

SUMMARY ILLUSTRATION FOR CHAPTER 37 996

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 38 Internal Transport: The Circulatory System 1001 Figure 41.12 Experimental Research Vocal Cues to Ovulation in
38.1 Animal Circulatory Systems: An Introduction 1003 Human Females 1097
38.2 Blood and Its Components 1006 41.5 Development 1104
38.3 The Heart 1009 SUMMARY ILLUSTRATION FOR CHAPTER 41 1118
38.4 Blood Vessels of the Circulatory System 1013
38.5 Maintaining Blood Flow and Pressure 1017 42 Control of Animal Processes: Neural Control 1123
42.1 The Basis of Information Flow in Nervous Systems:
Figure 38.19 Experimental Research Demonstration of a
An Overview 1124
Vasodilatory Signalling Molecule 1019
Figure 42.13 Experimental Research Demonstration of Chemical
38.6 The Lymphatic System 1019 Transmission of Nerve Impulses at Synapses 1136
38.7 Reflections on the Chapter 1021
42.2 Sensory Inputs: Reception 1137
SUMMARY ILLUSTRATION FOR CHAPTER 38 1022
Figure 42.25 Experimental Research How Do Sea Urchins Detect
39 Regulation of the Internal Environment: Water, Solutes, Light? 1147
and Temperature 1027 Figure 42.42 Experimental Research Magnetic Sense in Sea
39.1 Introduction to Osmoregulation and Excretion 1028 Turtles 1158
39.2 Osmoregulation and Excretion in Invertebrates 1032
42.3 The Central Nervous System: Integration 1159
39.3 Osmoregulation and Excretion in Non-mammalian
42.4 The Peripheral Nervous System: Transmission and
Vertebrates 1034
Response 1171
39.4 Osmoregulation and Excretion in Mammals 1036
SUMMARY ILLUSTRATION FOR CHAPTER 42 1174
Figure 39.15 Experimental Research ADH-Stimulated Water
Reabsorption in the Kidney Collecting Duct 1042 43 Muscles, Skeletons, and Body Movements 1179
43.1 Vertebrate Skeletal Muscle: Structure and Function 1180
39.5 Introduction to Thermoregulation 1043
39.6 Ectothermy 1045 Figure 43.5 Experimental Research The Sliding Filament Model of
Muscle Contraction 1183
39.7 Endothermy 1048
39.8 Reflections on the Chapter 1053 43.2 Skeletal Systems 1189
SUMMARY ILLUSTRATION FOR CHAPTER 39 1054 43.3 Vertebrate Movement: The Interactions between Muscles and
Bones 1193
40 Control of Animal Processes: Endocrine Control 1059 SUMMARY ILLUSTRATION FOR CHAPTER 43 1196
40.1 Hormones and Their Secretion 1060
40.2 Mechanisms of Hormone Action 1063 44 Animal Behaviour and Responses to the
Environment 1201
Figure 40.6 Experimental Research Demonstration That
44.1 Introduction 1203
Epinephrine Acts by Binding to a Plasma Membrane
Receptor 1066 Figure 44.3 Experimental Research The Role of Sign Stimuli in
Parent–Offspring Interactions 1204
40.3 The Hypothalamus and Pituitary 1068
44.2 Genes and Behaviour: Nature or Instinct 1204
40.4 Other Major Endocrine Glands of Vertebrates 1071
44.3 Environment and Behaviour: Learning or Nurture 1206
40.5 Endocrine Systems in Invertebrates 1076
44.4 Hormones and Behaviour 1207
Figure 40.16 Experimental Research Demonstration That Growth
and Moulting in Insects Is Hormonally Controlled 1078 44.5 Neurophysiology and Behaviour 1209
44.6 Finding Food/Defences against Predation 1211
40.6 Reflections on the Chapter 1079
44.7 Communication 1212
SUMMARY ILLUSTRATION FOR CHAPTER 40 1080
44.8 Choice of Habitat 1216
41 Animal Reproduction and Development 1085 Figure 44.28 Experimental Research Experimental Analysis of the
41.1 The Drive to Reproduce 1086 Indigo Bunting’s Star Compass 1221

41.2 Asexual and Sexual Reproduction 1086 44.9 Mating 1224

41.3 Mechanisms of Sexual Reproduction 1088 44.10 Social Behaviour 1226
41.4 Sexual Reproduction in Mammals 1095 SUMMARY ILLUSTRATION FOR CHAPTER 44 1232

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 45 Defences against Disease 1237 45.4 Impairment and Peculiarities of the Immune System 1251
45.1 Three Lines of Defence against Invasion 1238 45.5 Immune Systems in Other Organisms 1254
45.2 Innate Immunity: Nonspecific Defence 1239 SUMMARY ILLUSTRATION FOR CHAPTER 45 1256
45.3 Adaptive Immunity: Specific Defences 1242
Appendix A: Answers to Self-Test Questions A-1
Figure 45.10 Research Method Production of Monoclonal
Glossary G-1
Antibodies 1248
Index I-1

xvi CONTENTS

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 Letter to Students
As you take another step toward increasing your understanding
of the biological world, we welcome you to this exploration of
the diversity of life. The main goal of this textbook is to guide
you on a journey of discovery about life’s diversity across inter-
connected levels of organization, ranging from molecules to
genes, from cells to organs, and from species to ecosystems.
Along the way, we will explore many questions about the mech-
anisms underlying diversity as well as the importance of main-
taining diversity for all species, including our own.
Diverse perspectives…
With this edition of Biology: Exploring the Diversity of Life, we have
been deliberate in beginning an ongoing process of opening this
resource to a wider range of sources, voices, and perspectives
devoted to a more fulsome understanding of the interconnected
biological world. In particular, we offer this edition as a first step in
responding to the Truth and Reconciliation Commission of
Canada’s (TRC) Calls to Action and to the United Nations Dec-
laration on the Rights of Indigenous Peoples (UNDRIP). The
truth of the relationship between science and Indigenous Peo-
ples in Canada has often, unfortunately, been one of oppression
and exploitation. The racism that fuelled the colonization of the
land that is now Canada influenced all aspects of social life,
including the delivery of education and the conduct of science.
The residential school system used education as a tool of cul-
tural assimilation, with tragic effects on Indigenous communi-
ties that continue to play out today. Science education was rarely
provided in these schools, thus limiting the capacity for Indige-
nous peoples to effectively interact with scientists representing
government, industry, or academia. A significant underrepre-
sentation of Indigenous peoples in Canadian science- and tech-
nology-related careers continues to this day.
In many Indigenous communities, science and research
have been viewed fearfully as tools of oppression. Some scien-
tists have engaged in unethical practices in Indigenous com-
munities that have resulted in deliberate harm to people and/or
their environment. Indigenous Traditional Knowledge and
technologies have been taken, and sometimes commercialized,
without consent from or benefit to the original Knowledge-
Keeping Elders and their communities.
Part of the work of reconciliation in this edition is a contri-
bution to the ongoing shift in perspectives of science from
those of Eurocentrism (i.e., science as emanating from, and
largely exclusive to, European nations and their current or
former colonies) to a more open, decolonized, “multi-science”
perspective that acknowledges all Indigenous Peoples as having
scientific knowledge of the natural world, albeit through their
own cultural lenses. One such perspective is Indigenous
Science, the accumulated empirical knowledge that has been
11140_fm_ptg01.indd 17
generated by thriving and prosperous Indigenous peoples
through their observations of and relationships with natural
phenomena over millennia.
This book is one result of deepening relationships among
the primary authors and their professional colleagues, the edi-
torial and production teams at Cengage Canada, students of
biology, Indigenous Elders and scholars, as well as members of
our Indigenous Advisory Board. The author team participated
in Ceremony with Indigenous advisors, experiencing first-
hand Indigenous cultural practices to acknowledge the collec-
tive desire for the success of this project as an example of and
inspiration for reconciliatory learning. With open minds and
hearts, we are committed to creating a meeting place where sci-
entists of all backgrounds can tell their stories.
As an author, advisory, and production team including
both settlers and Indigenous People of Turtle Island, we want
our work to contribute to reconciliation and improved relations
among settlers and First Nations, Inuit, and Métis Peoples. As
educators, we want our classes to be widely accessible. We
understand that this means making efforts to increase the level
of safety felt by Indigenous students and others who have been
excluded in our science education system. As authors of educa-
tional resources, we want to promote innovative, wholistic,
effective andragogy that supports teaching and learning by all
students and instructors. As global citizens, we want to pro-
mote collaboration in harnessing the creative power of a wide
variety of perspectives in the face of global challenges.
An emphasis on the diversity of life…
At first glance, the riot of life that animates the biosphere over-
whelms our minds. One way to begin to make sense of this
diversity in a textbook is to divide it into manageable sections
on the basis of either similarities or differences. We examine
how different organisms solve the common problems of finding
nutrients, energy, and mates on the third rock from our Sun.
Evolution provides a powerful conceptual lens for viewing
and understanding the roots and history of the diversity of
living things. What basic evolutionary principles inform the
relationships among life forms regardless of their different
body plans, habitats, or life histories?
We will demonstrate how knowledge of evolution helps us
appreciate the changes we observe in organisms. Whether the
focus is the conversion of free-living prokaryotic organisms
into mitochondria and chloroplasts or the loss of eyes in cave
fish, selection for particular traits over time can explain the
current observations.
Examining how biological systems work is another theme
pervading this text and underlying the idea of diversity. We have
intentionally tried to include examples that will tax your imagina-
xvii
21/03/22 4:13 PM
 tion, from sea slugs that steal chloroplasts for use as solar panels, to
the molecular basis of high-altitude adaptations in deer mice, to
adaptive radiation of viruses. In each situation, we examine how
biologists have explored and assessed the inner workings of organ-
isms, from gene regulation to the challenges of digesting cellulose.
Solving problems is another theme that runs throughout
the book. Whether the topic is gene therapy to treat a disease in
people, preventing harmful algal blooms, or reducing the inci-
dence of human cancer, both the problem and the solution lie
in biology. We will explore large problems facing planet Earth
and the social implications that arise from them.
Emphasizing the big picture…
While many biology textbooks use the first few chapters to review
fundamentals of chemistry and biochemistry as well as informa-
tion on the scientific method, this book immediately engages in
some of the excitement, mystery and controversy that is modern
biology right up front in Chapter 1. Important background infor-
mation appears in the centre of the book as a distinct reference
section entitled The Chemical and Physical Foundations of
Biology. With their purple borders, these pages are distinct and
easy to find, and have become affectionately known as The Purple
Pages. These pages enable information to be readily identifiable
and accessible to you as you move through the textbook rather
than it being tied to a particular chapter. The concepts of atoms,
molecules, and macromolecules are connected through the
theme of “emergent properties.” By considering how the “stuff of
life” interrelates as a function of increasing complexity rather
than just memorizing the attributes of individual items, you can
better grasp why biology works the way it does rather than be
awed by how much information we know about it.
Throughout this book, we have highlighted the work of
Indigenous and non-Indigenous scientists in Canada and else-
where, and we present many examples and biological concepts
in Canadian contexts. We hope that you, as a student studying
in Canada, will find this context accessible and engaging.
Focusing on research to engage the living
world through a scientific lens…
A primary goal of this book is to evoke and sustain your curiosity
about biology, rather than dulling it with a mountain of discon-
nected facts. We can help you develop the mental habits of scien-
tists and a fascination with the living world by conveying our
passion for biological research. We want to excite you not only
with what biologists know about the living world but also with how
they know it and what they still need to learn. In doing so, we can
encourage some of you to accept the challenge and become biolo-
gists, posing and answering important new questions through
your own innovative research. For those of you who pursue other
careers, we hope that you will leave your introductory—and
perhaps only—biology course armed with intellectual skills that
will enable you to evaluate future knowledge with a critical eye.
xviii L E T T E R TO S T U D E N T S
11140_fm_ptg01.indd 18
In this book, we introduce you to a biologist’s “way of
learning.” Research biologists constantly integrate new observa-
tions, hypotheses, questions, experiments, and insights with
existing knowledge and ideas. To help you engage the world as
biologists do, we must not simply introduce you to the current
state of knowledge, we must also foster an appreciation of the
historical context within which those ideas developed and iden-
tify the future directions that biological research is likely to take.
Because advances in science occur against a background of
research, we also give you a feel for how biologists of the past
formulated basic knowledge in the field. By fostering an appre-
ciation of such discoveries, given the information and theories
available to scientists in their own time, we can help you under-
stand the successes and limitations of what we consider cut-
ting-edge today. This historical perspective also encourages
you to view biology as a dynamic intellectual enterprise, not
just a collection of facts and generalities to be memorized.
We have endeavoured to make the science of biology come
alive by describing how biologists formulate hypotheses and
evaluate them using hard-won data, how data sometimes tell
only part of a story, and how the results of studies often end up
posing more questions than they answer. Our exploration of
the Tully Monster in Chapter 23 is a case in point. Since its
fossil discovery and description, this mainly soft-bodied animal
has been tentatively classified with species in five different
groups of animals. Through this example and throughout
Chapter 23, we explore the current recognition that the histor-
ical and traditional grouping of animals into protostomes and
deuterostomes is more artificial than real.
Although you might prefer simply to learn the right answer
to a question, we encourage you to embrace the unknown—
those gaps in knowledge that create opportunities for further
research. An appreciation of what biologists do not yet know
may draw you into the field. And by defining why scientists do
not understand interesting phenomena, we encourage you to
think critically about possible solutions and to follow paths dic-
tated by your own curiosity. We hope that this approach will
encourage you to make biology a part of your daily life by
having informal discussions and debates about new scientific
discoveries with members of your personal and academic
communities.
This textbook is a toolbox…
You may see your professor as primarily a teacher, but as prac-
ticing scientists, professors are always learning from colleagues,
from the scientific literature, from their research programs, and
from their students. You may see yourself as primarily a learner,
but as a member of a social network, you are very likely to find
yourself explaining biological concepts to your friends, family
members, and the general public. Recognizing that we are all
lifelong learners and teachers may help you to see how each part
of your learning, both in and out of class, fits into the continu-
ously developing picture of your life.
21/03/22 4:13 PM
 Although it’s usual to think of this resource as merely a
book, it may be more helpful to imagine it as a toolbox con-
taining various interrelated components carefully designed to
inspire, inform, structure, connect, and confirm your learning.
Modern science courses are often blended and/or flipped.
That is, you will likely be assigned an online learning component
to complement subsequent face-to-face classroom interaction.
Professors will often assign textbook readings in advance of class.
If, for example, you are assigned to read Sections 14.2 to 14.4 of the
text, be aware that educational research has identified specific
strategies for approaching this reading that are more likely to
result in durable learning. The least-effective strategy is to go
directly to the assigned material and simply read it as you would
read a novel or social media post. Learning is more durable if new
concepts can be connected to ideas that are already known or rel-
evant to you. This means that the structure and context for new
learning is important. Assume that you are responsible for reading
certain sections of a chapter. Begin with the brief Why It Matters
section at the opening of the chapter. This will help you to see the
relevance of the material and, hopefully, allow you to make some
personal connection to the concepts. Check the chapter-opening
Roadmap to see how the assigned topics are related to other ideas
within the chapter and also to material in other chapters. Then
look over the Summary Illustration at the end of the chapter to
get a broad visual sense of the scope of the material. What do you
recognize from previous learning? What looks new? What looks
especially interesting? Then, go to the assigned sections of the
chapter with the awareness of how they fit into the bigger picture
of the chapter and to the concepts that are already known or inter-
esting to you. Even then, don’t just start reading at the first word.
Instead, continue the exercise of noting the structure of the mate-
rial by just noticing the main headings and perusing the figures.
Once you begin to read, create your own organization of
the material by making your own notes, highlights, etc.
Learning science requires that you understand new vocabu-
lary; consult the Glossary to confirm the meaning of unfa-
miliar terms (or look up elsewhere terms not included in the
Glossary—get comfortable with all of the vocabulary). Use
the embedded Study Break questions as an opportunity to
pause and help confirm that you have identified and understood
the major ideas. If you don’t know the answer to a Study Break
question, this is a sign that you should go back and read more
carefully and/or seek additional help. Pay particular attention to
the Concept Fix feature as well. These brief sections identify
and clarify many misconceptions that biology students are
prone to experience as they are learning. As your notes develop,
notice any connections that arise to what you have already
learned in previous classes and other courses. Highlighting such
relationships will help you to retain your new knowledge.
As you read, use the complementary photos and illustra-
tions to supplement and confirm your understanding. Notice
that graphs and anatomical drawings are annotated with inter-
pretive explanations that lead you step by step through the
major points they convey.
Science is a progressive enterprise in which answers open
new questions for consideration. As you read a chapter, watch
for the underlying thread of the research process that gave rise
to the information you are learning. Notice the three types of
specially designed research figures providing more detailed
information about how biologists formulate specific hypoth-
eses and test them by gathering and interpreting data. Experi-
mental Research figures describe specific studies in which
researchers used both experimental and control treatments,
either in the laboratory or in the field, to test hypotheses or
Why It Matters will help
you see the relevance of the
material and allow you to
make some personal
connections to the concepts.

Study Break Questions

provide an opportunity to
pause and help confirm that
you have identified and
understood the major ideas.
Assigned Readings
The Summary Illustration
will give you a broad, visual
sense of the scope of the material.
• What do you recognize from
previous learning?
• What looks new?
• What looks especially
interesting?

The Chapter Roadmap will

show you how each of the
chapters’ topics are related to
other ideas within the chapter
and to the material in other
chapters.

L E T T E R TO S T U D E N T S xix

11140_fm_ptg01.indd 19 21/03/22 4:13 PM

 answer research questions by manipulating the system they
studied. Observational Research figures describe specific
studies in which biologists have tested hypotheses by com-
paring systems under varying natural circumstances. Research
Method figures provide examples of important techniques,
such as light and electron microscopy, the polymerase chain
reaction, making a knockout mouse, DNA microarray analysis,
plant cell culture, producing monoclonal antibodies, radio-
metric dating, and cladistic analysis. Each Research Method
figure leads you through the purpose of the technique and pro-
tocol and describes how scientists interpret the data it
generates.
To complement the textbook material relating to research,
we encourage you to take any opportunity to participate in
hands-on laboratory or field investigations. Such first-hand
experiences can help you to see how the theoretical knowledge
in the book relates to your life and the aspects of biology that
interest you most.
As a final check of your learning, you might address selected
Self-Test Questions at the end of the chapter. These chapter-
review questions are organized according to Bloom’s Taxonomy
into three sections: Recall/Understand, Apply/Analyze, and
Create/Evaluate. This structure allows you to review the mate-
rial in a sequence that moves from the basic knowledge of fac-
tual material to more challenging and sophisticated applications
of that knowledge to novel situations. Although answers to the
xx L E T T E R TO S T U D E N T S
11140_fm_ptg01.indd 20
Self-Test Questions are found in an appendix at the back of the
book, looking at answers before generating your own can lead to
hindsight bias and misplaced confidence. Working through the
questions before consulting the answers will give you a more
accurate assessment of your understanding of the material
before you are faced with high-stakes exam questions.
In summary, we have applied our collective experience as
teachers, researchers, and writers to create text and comple-
mentary visuals that provide accessible explanations of funda-
mental concepts from an evolutionary perspective that binds
all the biological sciences. We hope that you are as captivated
by the biological world as we are and are drawn from one
chapter to another. But don’t stop there; use MindTap and other
resources to broaden your search for understanding and, most
importantly, observe and enjoy the diversity of life around you.
M. Brock Fenton
Denis Maxwell
Tom Haffie
Bill Milsom
Todd Nickle
Shona Ellis
Shelby Riskin
London, Toronto, Calgary, and Vancouver
January 2022
21/03/22 4:13 PM
 New to This Edition
This section highlights the changes we made to enhance the
effectiveness of Biology: Exploring the Diversity of Life, Fifth
Canadian Edition. Every chapter has been updated to ensure
currency of information. We made organizational changes to
more closely link related topics and preferred teaching
sequences. Key chapters have been extensively revised to pro-
vide a full treatment of the subject matter that reflects new
developments in these specialized fields.
Organization
We begin the fifth Canadian edition with the question “What is
life?” Chapter 1: Defining Life and Its Origins starts with the rec-
ognition that the origin and definition of life are widely consid-
ered to be one of the great mysteries and challenges in science.
It explores how we define the characteristics of life, why sci-
entists cannot agree on a universal definition of life, and how the
way a discipline defines life is intimately connected to the ques-
tions it explores. This opening chapter situates biology as a
method of inquiry that has at its core curiosity and a sense of awe.
This sense of wonderment is once again brought to the fore
in Unit 5: The Diversity of Life, which features a two-page spread
introducing students to the Tree of Life. The starburst tree pres-
ents students with a perspective on the truly astounding organ-
ismal diversity on the planet, the relative place of humans within
it, and the equally astounding amount of biological diversity that
remains unknown. This unit has been reorganized so that Bac-
teria and Archaea (Chapter 20) now precedes Protists (Chapter
21), followed by Fungi (Chapter 22), Plants (Chapter 23), Animals
(Chapter 24), and Viruses, Viroids, and Prions (Chapter 25).
Conservation of Biodiversity (Chapter 29) was moved from this
unit to the end of Unit 6: Ecology and Behaviour.
Plant Signals and Responses to the Environment (Chapter 34)
and Animal Behaviour and Responses to the Environment
(Chapter 44) have been moved to their respective units cov-
ering the systems and processes of plants and animals. And
finally, the fifth Canadian edition concludes with a new and
timely Chapter 45: Defences against Disease. Reintroduced into
the book by popular demand, this chapter covers innate and
adaptive immunity, vaccines, convalescent plasma and anti-
body therapies, malfunction and failures of the immune
system, and defence systems in other organisms.
Diversity in Ways of Knowing and Learning
The fifth Canadian edition constitutes a modest first step in our
contribution to the ongoing shift in perspectives of science from
Eurocentrism to a more open, decolonized, “multi-science”
perspective that acknowledges all Indigenous Peoples as having
scientific knowledge of the natural world through their own cul-
tural lenses. Throughout the book, we have made connections to
11140_fm_ptg01.indd 21
Indigenous Ways of Knowing and Learning in the Why It Matters
sections. We have included specific examples from First Nations,
Inuit, and Métis communities, using the traditional names of
Indigenous Nations and the traditional languages of Indigenous
groups. We have attempted to cite research from Indigenous
researchers; research where Indigenous communities have part-
nered with the researchers; and research where Indigenous
knowledge has informed, enhanced, or changed our under-
standing of a particular phenomenon. To authentically represent
concepts and stories that we have been given permission to share,
we have committed to ensuring that any associated artwork is
created by an Indigenous artist.
Points of Interest in Each Chapter
At the beginning of each chapter are redesigned Chapter Road-
maps. Instead of providing a visual representation of how the
chapter is organized, we revisited this feature with an eye to pro-
viding a clear visual overview of how the major topic areas in the
chapter relate to one another. In addition to showing how con-
cepts relate to each other within the chapter, the roadmaps also
show the connections between the topics in the chapter and the
other chapters in the book.
Chapter 1: Defining Life and Its Origins
● NEW. While we know so much about life, science struggles
to arrive at a simple definition of what it means to be living.
We discuss this interesting question.
● NEW. More robust discussion of the origins of metabolism
and how it may have evolved in close association with alka-
line hydrothermal vents.
● NEW discussion of how science has moved from a three-
domain view of the tree of life to a two-domain view
● NEW exploration of the latest evidence showing that eukary-
otes are descended from a lineage of the Archaea
Chapter 5: Cellular Respiration
● NEW Why It Matters exploring the role of mitochondria and
cold tolerance in Inuit individuals
Chapter 6: Photosynthesis
● NEW section on the physical nature of light
● NEW Concept Fix—Why are there three membranes in
chloroplasts and only two in mitochondria, both of which
evolved through endosymbiosis?
● NEW section on electron transport
Chapter 7: Cell Cycles
● NEW references to the fastest-growing prokaryotic cell,
Bibrino natiegens
xxi
21/03/22 4:13 PM
 ● Expanded relevance to cancer through reference to Peto’s
Paradox as commentary on underlying genetics of low
cancer incidence in large mammals
● NEW Summary Illustration
Chapter 8: Genetic Recombination
Reconstructed chapter to integrate sources of variability
●
with the description of respective stages of meiosis
● NEW Summary Illustration
Chapter 9: The Chromosomal Basis
of Inheritance
● NEW Why It Matters begins the process of decolonization
by acknowledging that people have applied their under-
standing of inheritance to purposefully increase desirable
characteristics among successive generations of plants and
animals for tens of thousands of years before Mendel
● NEW Summary Illustration
Chapter 10: Genetic Linkage, Sex Linkage,
and Other Extensions to Basic Inheritance
Mechanisms
● NEW reference to Indigenous clan system and ethical,
respectful efforts to address the lack of Indigenous sequences
in DNA databases
● NEW discussion of the three-parent baby controversy
● NEW connection to epigenetics and the possibility of a role
for epigenetics in multigenerational trauma
● NEW Summary Illustration
Chapter 11: DNA Structure, Replication, and
Repair
● Refined and clarified text descriptions of early experiments
● NEW Summary Illustration
Chapter 12: Gene Structure, Expression, and
Mutation
● NEW Summary Illustration
Chapter 13: Regulation of Gene Expression
● Substantial update of the paradigm of the mechanism under-
lying diauxic growth with respect to the lac operon
● NEW material on cancer incidence in Canada
● NEW Summary Illustration
Chapter 14: DNA Technologies
● Updated discussion of CRISPR
Chapter 15: Genomes
● NEW Why It Matters discussing how the Golden State Killer
was caught because a relative submitted their DNA to a
public database
xxii N E W TO T H I S E D I T I O N
11140_fm_ptg01.indd 22
Chapter 16: Evolution: The Development
of the Theory
● Thoroughly revised and reorganized, bringing the theory of
natural selection itself to the fore
Chapter 17: Microevolution: Changes within
Populations
● Clarified discussion around the four key aspects of
mutation
● Selected figures reviewed and revised for clarity
Chapter 18: Speciation and Macroevolution
● NEW Why It Matters describing how Native Papuans and
Ernst Mayr identified a nearly identical number of bird
species using very different methods
Chapter 19: Systematics and Phylogenetics:
Revealing the Tree of Life
● Revised Summary Illustration
Chapter 20: Bacteria and Archaea
● NEW discussion around Pseudomonas aeruginosa as
common co-infecting pathogen in COVID-19 patients
● Discussion of how tuberculosis is a leading cause of death
due to infectious disease throughout history and how Indig-
enous people have been disproportionately affected by this
disease
● Revised Summary Illustration
Chapter 21: Protists
● NEW Why It Matters about how the people of Haíl~zaqv,
Heiltsuk Nation, collect, process, and use seaweeds from
intertidal areas
● NEW Summary Illustration
Chapter 22: Fungi
● NEW material on the four main invasive fungal co-infec-
tions with COVID-19
● NEW discussion of Ch’ãłtthi (willow tinder fungus), used by
the Denesuline (Chipewyan) people as tinder and
packaging
Chapter 23: Plants
● Reorganized for flow and clarity
● Updated and revised selected illustrations for clarity
● NEW discussion of the story behind the name of Indian pipe
(Monotropa uniflora)
NEW discussion of sk’in (“horsetails” in the Haida language)
●
and how it was used
● NEW discussion of the pitch from species of the pine family,
which has many traditional uses by Indigenous Peoples
● Revised Summary Illustration
21/03/22 4:13 PM
 Chapter 24: Animals
● Completely reorganized and streamlined to shift the
approach to understanding and recognizing patterns and
variations
● Emphasis on key groups rather than a description of all
groups
● More emphasis on concepts and less on details
● NEW figures providing an overview of the group of animals
discussed in each section
Chapter 25: Viruses, Viroids, and Prions:
Infectious Biological Particles
● NEW Why It Matters discussing the impact of smallpox on
Indigenous communities
● Thoroughly revised and updated, including material on
SARS-CoV-2
● NEW section on virotherapy
Chapter 26: Population Ecology
● NEW Why It Matters discussing a collaboration between the
University of Guelph and the Chippewa of Nawash Unceded
First Nation identifying five general principles associated
with Nawash harvesting
● Thoroughly revised, streamlined, and updated
● NEW definitions and discussion of semelparity and
iteroparity
Chapter 28: Ecosystems
● NEW Why It Matters about the impact of 19th-century colo-
nial development along Lake Erie
● NEW discussion of Three Sisters crop complementarity
● Thoroughly revised, streamlined, and updated
● Updated discussion of climate change and disruption of the
global carbon cycle
NEW Summary Illustration
●
Chapter 30: Organization of the Plant Body
● NEW Why It Matters featuring a discussion of culturally
modified trees written by researcher and Heiltsuk archeolo-
gist Elroy White (Q̌íx̌ itasu) of central coast British Columbia
● Revised Summary Illustration
Chapter 31: Transport in Plants
● Figures revised and updated for clarity
Chapter 33: Plant Nutrition
● NEW Why It Matters about the role of the salmon cycle and
plant growth, featuring original artwork by Indigenous artist
Chester Lawson of the Haíłzaqv (Heiltsuk) Territory, Central
Coast, British Columbia.
● NEW discussion of Palliser’s Triangle in Alberta/Saskatch-
ewan, a hunting ground for Cree, Sioux, and Blackfoot
11140_fm_ptg01.indd 23
Chapter 34: Plant Signals and Responses
to the Environment
● NEW Why It Matters about the observation of plant behav-
iour to predict the timing of other resources, such as the
soapberry harvest of Nlaka’pamux First Nation of Thompson
River, British Columbia, and the link between elderberry
blooms and halibut fishing among the First Nations of Nuu-
chah-nulth on Vancouver Island
● NEW discussion of heliotropism
Revised Summary Illustration
●
Chapter 36: Animal Nutrition
● NEW Why It Matters about dietary requirements in animals
and across cultures, and how there are no “essential foods,”
only essential nutrients
● NEW material on the human gut biome
Chapter 37: Gas Exchange: The Respiratory
System
● NEW Why It Matters featuring a discussion of the many
roles that breathing plays across different cultures
Chapter 38: Internal Transport: The Circulatory
System
NEW Why It Matters featuring the significance of the notion
●
of the beating heart across different cultures
● NEW section—Heart Attacks and Strokes Arise from Poor
Blood Flow to the Heart Muscle and Brain
● Revised Summary Illustration
Chapter 40: Control of Animal Processes:
Endocrine Control
● Revised Summary Illustration
Chapter 41: Animal Reproduction and
Development
● NEW Why It Matters featuring the reproductive process of
Pacific herring off the coast of British Columbia
● Revised Summary Illustration
Chapter 44: Animal Behaviour and Responses
to the Environment
● NEW Why It Matters featuring a discussion of how Indige-
nous hunting societies around the globe have relied on
knowledge of the behaviours of their food species for
survival
● Thoroughly revised and streamlined
NEW Chapter 45: Defences against Disease
● NEW chapter
● Why It Matters featuring a discussion of plagues, epidemics,
and pandemics, including COVID-19
N E W TO T H I S E D I T I O N xxiii
21/03/22 4:13 PM
 Welcome to Biology:
Exploring the Diversity of Life, 5Ce
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.

◀ Each chapter begins with Why Does ___ Matter to Me?,

an engaging reading that explores a real-life application of
the chapter topic, providing students with context around
the relevance of the material they are learning.

▲ Learn It! The Media Library, with its dynamic animations

and videos, offers a different entry point into the content that ▲ Study It! Exam Prep auto-graded questions have been
helps students process and understand concepts in a different carefully curated to replicate the type and level of questions
way. that students will encounter in a testing scenario. Chapter
Flashcards help students remember key term in each chapter.

xxiv

11140_fm_ptg01.indd 24 21/03/22 4:13 PM

 ◀ Apply It! Think and Engage Like a Scientist by taking
gradable short-answer quizzes:

• Apply Evolutionary Thinking questions ask students to

interpret a relevant topic in relation to the principles of
evolutionary biology.
• Design an Experiment challenges students’
understanding of the chapter and helps them deepen
their understanding of the scientific method as they
consider how to develop and test hypotheses about a
situation that relates to a main chapter topic.
• Interpret the Data questions help students develop
analytical and quantitative skills by asking them to
interpret graphical or tabular results of experimental or
observational research experiments for which the
hypotheses and methods of analysis are presented.

Also available in MindTap for Biology are engaging and

informative videos that accompany The Purple Pages. From
matter to polypeptides, author Todd Nickle of Mount Royal
University (pictured) will walk students through these
foundational concepts, strengthening their understanding
and helping them build a strong base of knowledge and
understanding of biology.

Test students’ mastery of concepts

with Conceptual Learning
Activities that complement the
text and help them learn and
understand key concepts through
focused assignments, an engaging
variety of problem types,
exceptional text/art integration,
and immediate feedback.

xxv

11140_fm_ptg01.indd 25 21/03/22 4:13 PM

 Active Learning
Visually stunning features are provided engage your students in the process of
learning because an engaged student is a successful student.
Chapter Roadmap
Plant Nutrition

33.1
To Chapter 6 for details about
photosynthesis and how some
plants are modified for hot and Plant Nutritional Requirements
dry environments
Plants can make all the macromolecules,
including their subunits. Plants supply
macromolecules to other trophic levels. Nitrogen
acquisition often limits plant growth. Symbiosis

© Chester Lawson
and predation on animals are adaptations to help
From Chapter 30 to make
plants thrive.
connections between root
structure and water/nutrient
absorption

PT Spawning Salmon by Chester Lawson

33
To Chapter 27 make connections
between nutrients and organism
interactions Plant Nutrition
33.2 33.3
Why it matters. . . Coastal British Columbia is known for its rugged mountains, rivers,
Soil Root Adaptations for and lush forests. The ocean is connected through the rivers to what have been termed the “salmon
Plant health depends on soil for both Obtaining and Absorbing Nutrients forests.” Salmon has sustained the peoples of these lands for thousands of years; First Nations fish-
texture and dissolved nutrients. Humus Microorganisms such as bacteria and eries were run sustainably, ensuring the returning runs of coho, pink, sockeye, chinook, and chum.
provides nutrition and also stores water, both of fungi contribute to nutrient cycling. Symbiotic
which are absorbed by roots. Soil acidity and ion
Fishing technologies were designed so that only what was needed was taken. The migration runs
associations are important in making nutrients
content also affect availability of dissolved occur at certain times of the year, when enough salmon would be collected and processed to pro-
from the soil more available to the plant. Root
nutrients to the plant. adaptations encourage symbiotic interactions.
vide food all year round. Drying and smoking were important forms of preservation; more recently,
Xylem carries dissolved materials to the shoots canning and freezing have become important. These salmon are on their migratory path to their
from the roots. place of birth in rivers, streams, and lakes, where they spawn and then die.
What does this have to do with plant nutrition? Salmon are not only integral to the health and
To Chapter 31 to consider how nutrients nutrition of First Nations, they are crucial to the health of the ecosystems of the coast as well as
are taken up and distributed in plants
further inland as the fish venture up into the interior through the networks of waterways. Imagine
To Chapter 22 to consider
how nutrients are taken up thousands of fish making their way up rivers, with bears, otters, eagles, and ravens depending on
and distributed in plants To Chapter 34 to make their arrival. The salmon that make it to their spawning grounds lay their eggs, fertilize them, and
connections between
To Chapter 28 to investigate nutrients and auxin as
then die. Bears in particular drag the fish carcasses up into the forest. They are selective eaters and
how nutrients are cycled well as other plant often consume only particular portions of the fish, leaving the carcasses to provide food and nesting
within an ecosystem hormones sites (e.g., for blowflies), and for decomposers to recycle their nutrient-rich bodies (Figure 33.1). There
is a distinct fragrance that fills the air during these runs, the smell of decomposition, as you find
carcasses along the riverside, the riparian areas, and deep in the forest. The nutrients within the fish,
particularly nitrogen, are made available to other organisms. As we will see in this chapter, nitrogen
is a limiting nutrient in plants, and a number of researchers have followed its path through coastal
ecosystems. Nitrogen does not have a direct route into plants; microorganisms are instrumental in
breaking down the carcasses, and important intermediaries make the nitrogen available for uptake
by plants. Animals that consume the fish release nitrogen in their excrement and urine (urea), which
get broken down by soil enzymes and microbes. Plants absorb nitrogen in the form of nitrate and
ammonium, often facilitated by fungi through mycorrhizal associations or bacterial symbioses.
Within the plants, this nitrogen is incorporated into organic molecules such as proteins and nucleic

875

▲ Chapter Roadmaps The Chapter Roadmaps provide a visual overview of how the major topic areas in the chapter
relate to one another and show the connections between the topics in the chapter and other chapters in the book.

Why It Matters Why It Matters draws students in with an engaging vignette that is linked to the concepts discussed in
the chapter.

Concept Fix Concept Fix sections draw on the extensive

research literature dealing with misconceptions commonly
held by biology students. Strategically placed throughout
the text, these short segments help students identify—and
correct—a wide range of misunderstandings. ▼

▲ Study Break The Study Break features fall at the end of

each major section and encourage students to pause and
review what they have learned before going on to the next
topic within the chapter.

xxvi

11140_fm_ptg01.indd 26 21/03/22 5:52 PM

 ◀ Experimental Research Experimental Research
FIGURE 10.2 Experimental Research figures describe specific studies in which research used both
Evidence for Gene Linkage
experimental and control treatments—either in the
Question: Do the purple-eye and vestigial-wing genes of Drosophila assort independently?
laboratory or in the field—to test hypotheses or answer
Experiment: Morgan crossed true-breeding, wild-type flies having red eyes and normal wings with purple-eyed, vestigial-winged flies. The F 1
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
research questions by manipulating the system they
analyzed the phenotypes of the progeny.
studied.
1. P generation Wild-type female Double mutant male
(red eyes, wild-type 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, wild-type wings) (red eyes, wild-type 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, wild-type 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,

wild type 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, wild type. If the genes assorted independently, the expectation is for a 1:1:1:1 ratio for testcross progeny: approximately 1420
of both parental and recombinant progeny.

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.
Research Method Research Method figures provide
240 UNIT 2 GENES
examples of important techniques, lead students through
the purpose of the technique and protocol, and describe
how scientists interpret the data generated. ▼

FIGURE 14.3 Research Method

FIGURE 18.16 Observational Research Identifying a Recombinant Plasmid Containing a Gene of Interest
Purpose: To identify a recombinant plasmid containing a gene of
Chromosomal Similarities and Differences among Humans and the Great Apes Protocol: interest from a ligation reaction mixture containing a bacterial
Question: Does chromosome structure differ between humans and their closest relatives among the apes? 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
Hypothesis: Large-scale chromosome rearrangements contributed to the development of reproductive isolation between species within the
evolutionary lineage that includes humans and apes.
1. The ligation reaction produces recombinant plasmids (the only
Prediction: Chromosome structure differs markedly between humans and their close relatives among the great apes: chimpanzees, gorillas,
and orangutans. products that might contain the gene of interest), nonrecombinant
plasmids, and joined pieces of genomic DNA (not shown).
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
Recombinant plasmid Nonrecombinant plasmid
present in humans and in the 3 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–
Results: Analysis of human chromosome 2 reveals that it was produced by the fusion of two smaller chromosomes that are still present in the
E. coli (which cannot make
other three species. Although the position of the centromere in human chromosome 2 matches that of the centromere in one of the
β-galactosidase) with a sample of the
chimpanzee chromosomes, in 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
Centromere position is similar a plasmid or transformed
in humans and chimpanzees
Chimpanzee Selection: with gene fragments
Bacteria transformed with plasmids
grow on medium containing ampicillin
because of ampR gene on plasmid.
Matching bands Untransformed bacteria 3. Spread the bacterial cells on a plate of growth medium
or bacteria transformed
Gorilla Screening: with gene fragments
containing ampicillin and X-gal, and incubate the plate
Blue colony contains bacteria cannot grow on medium until colonies appear.
with a nonrecombinant containing ampicillin.
Compared to the chromosomes of humans and chimpanzees, plasmid; that is, the lacZ +
the region that includes the centromere is inverted (its position gene is intact.
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
Conclusion: Differences in chromosome structure between humans and both gorillas and orangutans are more pronounced than they are in this case the gene of interest. ampicillin and X-gal
Restriction lacZ+ gene
between humans and chimpanzees. Structural differences in the chromosomes of these four species may contribute to their reproductive site Plasmid
isolation.
cloning
Source: 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)

▲ Observational Research STUDY

chromosome rearrangement becomes established within a
Observational
BREAK QUESTIONS
Research 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

figures describe specific studies

population, that population will diverge more rapidly from
populations lacking the rearrangement. The genetic diver-
in which
biologists have 1. How can natural selection promote reproductive isolation in
allopatric populations?
recombinant plasmids. Bacteria making up blue colonies contain nonrecombinant plasmids. These plasmids have intact lacZ 1 genes and produce
β-galactosidase, which changes X-gal to a blue product. Bacteria that form the white colonies contain recombinant plasmids. Each recombinant

tested hypotheses by comparing systems under varying

gence eventually causes reproductive isolation. 2. How does polyploidy promote speciation in plants? plasmid has a DNA fragment (in this example, the gene of interest) inserted into the lacZ1 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 quantities of the recombinant plasmid that can be isolated and purified for analysis and/or manipulation of the gene 5 “gene of interest.”
natural circumstances.
465
xxvii
CHAPTER 18 S P E C i AT i o n A n d M A C R o E v o lu T i o n

C H A P T E R 14 D N A T E C H N O LO G I E S 351

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 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 Citric Acid Cycle

Acetyl CoA
C C – CoA CoA
Six-carbon molecule formed
from acetyl CoA + oxaloacetate
Cellular respiration is a collection of reactions that converts the chemical Four-carbon acceptor molecule
C C C C C C Citrate
energy present in carbohydrates, fats, and proteins into an energy currency, (regenerated in each cycle)
ATP, that is readily used by the cell. The phases of cellular respiration C C C C Oxaloacetate
are (i) glycolysis, (ii) pyruvate oxidation and the citric acid cycle, and NAD+
One carbon
(iii) oxidative phosphorylation. The overall process is represented by the lost as CO2
chemical equation for the oxidation of glucose. NADH NADH + CO2

C C C C C
NAD+
Cellular respiration is a redox process. NAD+ Second carbon
lost as CO2
• The overall change in free energy (ΔG) is NADH + CO2
negative. C C C C
• Energy released during the formation
C6H12O6 + 6O2 6CO2 + 6H2O
Reduced electron carrier
of the bonds in CO2 and H2O is greater similar to NADH GDP, P
FADH2 i
than the energy required to break the
FAD
bonds in glucose and O2. GTP Energy-carrying molecule
equivalent to ATP
• The goal is to couple the energy released
Glucose to the synthesis of ATP!
Pyruvate is transported into the mitochondrial matrix and converted to acetyl-CoA,
which enters the citric acid cycle and is oxidized. Notice the products formed.
Glycolysis
Glycolysis occurs in the cytosol and splits
glucose into pyruvate, which yields ATP and
NADH. The fate of pyruvate depends on Low [O2] Oxidative Phosphorylation
the availability of oxygen. Pyruvate Lactate/ethanol
Cytosol
Fermentation Outer
mitochondrial
membrane

High [O2]
H+ H+
While prokaryotic cells lack mitochondria, H+ H+
H+ H+ H+
H+ + H+ H+ H+
many bacteria and archaea possess a H+ H H+ ATP
complete respiratory pathway that is H+ H+ H+ synthase
H+ H+
similar to eukaryotes. Intermembrane
Citric Acid Cycle e–
compartment cyt c
e– Complex
Inner UQ e– IV
NADH , Complex
mitochondrial Complex e–
I UQ III
FADH2 membrane
e– Complex
Oxidative
e– II H+
Phosphorylation
H+
ATP NADH H+ NAD+ FADH2 FAD 2e– + 2 H+ + 1/
2 O2
H2O
Mitochondrial
matrix
ADP + P H+ ATP
i

Electron transport Chemiosmosis

Electrons from the oxidation of NADH and FADH2 produced by glycolysis and the
citric acid cycle are passed along an electron transport chain. Electron transport
and chemiosmotic synthesis of ATP are coupled by a proton gradient.

130 131

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 the appendix at the back of the book. ▼

self - test questions Appendix A: Answers to Self-Test

Recall/Understand 8. Suppose you are a doctor and your patient complains of feeling
hot all the time, even on the coldest winter days. The young
Questions
1. Which of these factors is found in organic molecules that are man perspires constantly and his skin is always flushed. He also
considered good fuels?
a. many C—H bonds
eats a lot but is rather thin. You perform some laboratory tests, Chapter 1 15. Anchoring junctions function to rein- Chapter 4
and find that the patient consumes lots of oxygen in his meta- 1. c 2. b 3. a 4. a 5. d 6. d 7. b 8. c 9. d force cell-to-cell connections made by 1. a 2. c 3. b 4. c
b. many C=C double bonds bolic pathways. What would you suspect this patient suffers 10. b 11. b 12. d adhesion molecules. Tight junctions 5. Some proteins perform transport;
c. an abundance of oxygen from and why? 13. To determine relative dates, scientists seal the spaces between cells. Gap junc- others have enzymatic activities; some
d. a high molecular weight order fossils found in different strata tions create direct channels for com- are a part of signal transduction pro-
2. What is one of the places in a cell where cellular respiration municating between adjacent cells.
Create/Evaluate in sequence from the lowest (oldest) cess; and others are involved in attach-
occurs? to the highest (newest) strata. This ment and/or recognition.
9. In cellular respiration, which of the following does the term un-
a. in plant mitochondria, but not in animal mitochondria
coupled refer to, specifically?
sequencing reveals their relative ages. Chapter 3 6. b 7. c 8. b 9. c 10. c 11. b 12. a 13. d
b. in plant chloroplasts By using a technique called radio- 1. c 2. d 3. b 4. a 5. b 6. c 7. c 8. d 9. c 14. Passive transport occurs down the
c. in the mitochondria of both animals and plants a. The two parts of glycolysis are running independently of metric dating, scientists can estimate 10. d concentration gradient of the solute,
d. in animal mitochondria, but not in plant mitochondria each other. the age of a rock by noting how much 11. As they dissolve, the sugar molecules and active transport occurs against
b. Respiratory electron transport is operating, but chemiosmo- of an unstable “parent” isotope has raise their entropy. However, the crys- the gradient of the transported solute.
3. Which of these processes occurs during glycolysis? sis is inhibited.
a. oxidation of pyruvate decayed to another form. By mea- tals re-form because the water Active transport therefore requires a
c. Respiratory electron transport is operating, but proton suring the relative amounts of the decreases in its order, changing from protein and energy to perform.
b. reduction of glucose pumping is inhibited. parent radioisotope and its break- compact liquid to disordered vapour. 15. They are both a form of passive trans-
c. oxidative phosphorylation d. Oxidative phosphorylation is occurring, but the proton- down products, and comparing this 12. Although sugar breakdown is an exer- port, but facilitated diffusion utilizes
d. substrate-level phosphorylation motive force remains high. ratio with the isotope’s half-life—the gonic process, the activation energy proteins to speed up the transport of
4. Which of these processes feeds glycolysis products into the citric 10. Phosphofructokinase (PFK) is regulated by a number of metabo- time it takes for half a given amount of required for the reaction to proceed is solute across the membrane.
acid cycle? lites. In addition to those mentioned in the text, which one of the radioisotope to decay—researchers high, which makes sugar unreactive.
a. chemiosmosis can estimate the absolute age of the
b. formation of G3P
following would also make sense?
rock.
The addition of heat or an enzyme can Chapter 5
a. Pyruvate could function as an activator of PFK. increase the rate of breakdown. 1. a 2. c 3. d 4. d 5. b 6. c 7. a
c. reduction of NAD+ b. Glucose could function as an inhibitor of PFK. 14. LUCA stands for Last Universal 13. Any substance in an ordered state
d. pyruvate oxidation 8. This patient might have defective
c. ADP could function as an activator of PFK. Common Ancestor. We know it existed (minimum entropy) will contain mol- mitochondria in his cells. This condi-
d. Acetyl-CoA could act as an activator of PFK. because of the core similarities that all ecules with maximum free energy. On tion is common in a number of dis-
Apply/Analyze 11. Which of these statements best describes the “paradox of aerobic
forms of life on Earth today possess. the contrary, any substance in a disor- eases. The reason why it was suspected
The likeliest explanation for these dered state (maximum entropy) will
5. The breakdown of fats releases fatty acids. In what form do the life”? is that, based on his symptoms, prob-
common traits is all organisms today contain molecules with minimum
carbon molecules enter the respiratory pathway? a. Humans are completely protected from the toxic effects of ably little ATP is synthesized, in spite
are descended from a single ancestor free energy. The relationship is
a. as NADH oxygen. of high oxygen consumption, since his
that also possessed all those traits. reversed.
b. as acetyl-CoA b. Hydrogen peroxide is formed when a single electron is cells dissipated a lot of heat (the
15. Bacteria and Archaea have circular 14. In an exergonic reaction, reactants
c. a glucose donated to O2. patient was hot all the time).
chromosomes; eukaryotes have linear contain more free energy than the
d. as pyruvate c. Cytochrome oxidase is a major source of reactive oxygen 9. b 10. c 11. d
chromosomes. The location of DNA products; energy is released and the
species. 12. Direct burning of glucose is an uncon-
6. You are reading this text while breathing in oxygen and breathing in prokaryotes is in the nucleoid reaction is spontaneous. In an ender- trolled process; cellular respiration
out carbon dioxide. Which two processes are the sources of the d. Strict anaerobes often lack the enzyme(s) superoxide dis- region, and in eukaryotes it is in the gonic reaction, reactants contain less
mutase and/or catalase. occurs in a series of steps and is there-
carbon dioxide? nucleus. Chromosomes segregate by free energy than the products; energy fore a form of controlled combustion.
a. glycolysis and pyruvate oxidation 12. Compare direct burning of glucose and cellular respiration with binary fission in prokaryotes, and by is required and the reaction is not 13. Reduction is the acceptance of elec-
b. glycolysis and oxidative phosphorylation reference to their progression. mitosis/meiosis in eukaryotes. Introns spontaneous. trons during a redox reaction. Oxida-
c. pyruvate oxidation and the citric acid cycle are rarely present in bacteria, but are 15. At any time in a cell, there must be
13. Distinguish between reduction and oxidation during redox tion is the loss of electrons during a
d. the citric acid cycle and oxidative phosphorylation common in archaea and eukaryotes. exergonic reactions happening to pro-
reactions. redox reaction.
vide enough energy for endergonic
7. Under conditions of low oxygen, what key role is played by fer- 14. Explain what happens with hydrogen and its bonding electrons 14. Hydrogen and its electrons move from
mentation in overall metabolism? during cellular respiration. Chapter 2 reactions. In addition, the energy sugar to oxygen, forming water.
1. d 2. c 3. c 4. a 5. a 6. b 7. b 8. a 9. d released by exergonic reactions must 15. The process of oxidative phosphory-
a. It regenerates the NAD+ required for glycolysis.
15. Cyanide is a strong toxin that reacts with the final protein in the 10. a 11. d 12. d 13. c be higher than the energy needed for lation produces the large number of
b. It synthesizes additional NADH for the citric acid cycle.
electron transport chain (ETC). Explain why it can kill a human 14. ribosomes, rough ER, transport ves- endergonic reactions because some ATP molecules needed for the ender-
c. It allows for pyruvate to be oxidized in mitochondria.
within a few minutes. icle, Golgi complex, secretory vesicle, energy is always transferred to heat gonic reactions in the cell that we are
d. By activating oxidative phosphorylation, it allows for the
plasma membrane (second law of thermodynamics). so dependent on. One of the major
synthesis of extra ATP.

A-1

xxviii

132 UNIT 1 SYSTEMS AND PROCESSES—THE CELL

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 ◀ The Chemical and Physical Foundations of
Biology (The Purple Pages) While many textbooks use
the first few chapters to introduce and/or review, we believe
that the first chapters should convey the excitement and
interest of biology itself. We therefore placed important
background information about biology and chemistry in a
reference section entitled The Chemical and Physical
Foundations of Biology in the centre of the book. With their
purple borders, these pages are distinct and easy to find
and have become affectionately known as The Purple Pages.
References to material covered in The Purple Pages are set in
purple in the Chapter Roadmaps and in the Index.

The Green Pages—Unit 5: The Diversity of Life opens with the Tree of Life, emphasizing the richness and tremendous
variability among living organisms explored in the chapters of The Green Pages. With their green borders, these pages identify
chapters that introduce and explore the diversity of life. ▼

Staphylococcus
UNIT 5 THE DIVERSITY OF LIFE
(Tenericutes)

Bacteria
Zixibacteria
Cloacimonetes
Atribacteria
Aqui cae
Actinobacteria Armatimonadetes
Chloro exi
Nomurabacteria Kaiserbacteria
Adlerbacteria
Campbellbacteria
Tree of Life
Fibrobacteres
Gemmatimonadetes
Calescamantes
Caldiserica Firmicutes
Researchers at the University of California, Berkeley, recently generated a Tree of Life
WOR-3 Cyanobacteria
TA06
Dictyoglomi
Thermotogae Cyanobacteria (opposite) comprising 3083 species, representing the major lineages of life on Earth.
Poribacteria Deinococcus-Therm.
Latescibacteria
BRC1
Synergistetes
Fusobacteria
Giovannonibacteria
Wolfebacteria
The tree was constructed by comparing the DNA sequences of genes that code for spe-
Melainabacteria
Marinimicrobia
RBX1
Jorgensenbacteria
cific ribosomal proteins. Organisms with fewer sequence differences were grouped more
Bacteroidetes Ignavibacteria
Chlorobi Caldithrix WOR1
closely together than species where the sequences were more dissimilar.
Azambacteria
Parcubacteria This tree is called a starburst tree. The distant past is at the centre and the outer edge
PVC
superphylum
Planctomycetes
Yanofskybacteria
Moranbacteria
represents today. This type of representation is called an un-rooted tree because, in the
Elusimicrobia
Chlamydiae,
Lentisphaerae, Magasanikbacteria centre, you will notice there is no clear starting point. The research that produced the
Verrucomicrobia Uhrbacteria
Falkowbacteria Candidate tree was not designed to provide clarity about the origin of life or how early lineages
Omnitrophica
SM2F11
Phyla Radiation developed—questions related to the distant past are met with tremendous uncertainty.
Aminicentantes Rokubacteria NC10 Peregrinibacteria
Acidobacteria
Tectomicrobia, Modulibacteria Gracilibacteria BD1-5, GN02
Absconditabacteria SR1
Instead, the research team was focused on looking at the relationships among the diver-
Nitrospinae
Nitrospirae
Dadabacteria
Saccharibacteria
Berkelbacteria sity of life that exists today. And this is the focus of the chapters that make up Unit 5.
Deltaprotebacteria
(Thermodesulfobacteria)
Chrysiogenetes
Of the 3083 DNA sequences used to build the tree, over 1000 represent different
Deferribacteres
Hydrogenedentes NKB19
Woesebacteria
microbial species that scientists have never actually studied in a laboratory! Their DNA
Spirochaetes Shapirobacteria
TM6
Wirthbacteria Amesbacteria
Collierbacteria
was discovered in various environmental samples (e.g., soil, water) and is represented
Epsilonproteobacteria Pacebacteria
Beckwithbacteria on the tree by red dots. Several research groups are developing methods to try and iso-
Roizmanbacteria
Dojkabacteria WS6
CPR1
Gottesmanbacteria
Levybacteria late and grow these microbes in the laboratory, a critical step to linking an actual
CPR3 Daviesbacteria Microgenomates
Alphaproteobacteria
Katanobacteria Curtissbacteria
WWE3
organism to the DNA sequences isolated from the environment. This includes members
Zetaproteo. of a specific group of archaeans called Asgard, which is represented on this tree by Loki.
Acidithiobacillia As discussed in Chapter 1, evidence indicates that the Asgard are the closest prokary-
Betaproteobacteria
Major lineages with isolated representative - italics otic relatives of eukaryotes.
Major lineage lacking isolated representative -
0.4
You will notice that some branches of the tree are longer than others. Branch length
Gammaproteobacteria
is related to the amount of evolutionary change. Along longer branches, more sequence
change has occurred than along shorter branches. For example, the branch leading to
E. coli
eukaryotes is particularly long, meaning there are many genetic differences between the
Salmonella organisms in that group compared to other organisms on the tree. To figure out exactly
how much genetic change has occurred you need to use the scale bar. The 0.4 on the
scale means that, for that length on any branch of the tree, there have been 0.4 changes
Micrarchaeota
Diapherotrites Eukaryotes at each nucleotide position in the RNA genes used to produce the tree.
Nanohaloarchaeota
Aenigmarchaeota Loki.
Parvarchaeota Thor.

Korarch.
0.4 DPANN Crenarch.
Pacearchaeota Bathyarc.
YNPFFA
Nanoarchaeota Aigarch.
Woesearchaeota Opisthokonta
Altiarchaeales Halobacteria
Z7ME43

Archaea
Methanopyri
Methanococci TACK Excavata
Hadesarchaea
Thermococci Thaumarchaeota Archaeplastida Animals
Methanobacteria and
Thermoplasmata
Archaeoglobi
Chromalveolata fungi
Methanomicrobia Amoebozoa
Plants, red and
green algae

xxix
495

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 Student and Instructor Resources
Student Resources
MindTap
Modern students require modern solutions. MindTap is a flex-
ible all-in-one teaching and learning platform that includes the
full ebook, a customizable learning path, and various course-
specific activities that drive student engagement and critical
thinking.
Download the Cengage Mobile App
Get access to a full, interactive ebook, readable online or off;
study tools that empower anytime, anywhere learning; and 24/7
course access. The Cengage Mobile app keeps students focused
and ready to study whenever it is convenient for them.
Instructor Resources
Test Bank
The test bank is available on a cloud-based platform. Testing
Powered by Cognero® is a secure online testing system that
allows instructors to author, edit, and manage test-bank content
from anywhere internet access is available. No special installa-
tions or downloads are needed, and the desktop-inspired inter-
face, with its drop-down menus and familiar, intuitive tools,
allows instructors to create and manage tests with ease. Multiple
test versions can be created in an instant, and content can be
imported or exported into other systems. Tests can be delivered
from a learning management system, the classroom, or wher-
ever an instructor chooses. Testing Powered by Cognero for
Biology: Exploring the Diversity of Life, Fifth Canadian Edition,
can be accessed through http://login.cengage.com.
PowerPoint
Microsoft PowerPoint® lectures have an average of 80 slides per
chapter, many featuring key figures, tables, and photographs
from Biology: Exploring the Diversity of Life, Fifth Canadian Edi-
tion. Instructor notes have been incorporated throughout,
making it simple for instructors to customize the deck for their
courses. The PowerPoint slides also feature “build slides”—
selected illustrations with labels from the book that have been
reworked to allow optimal display in PowerPoint.
xxx
11140_fm_ptg01.indd 30
TurningPoint®
Another valuable resource for instructors is TurningPoint® class-
room response software customized for Biology: Exploring the
Diversity of Life, Fifth Canadian Edition. Now instructors can
author, deliver, show, access, and grade, all in PowerPoint, with
no toggling back and forth between screens! JoinIn on Turning-
Point is the only classroom response software tool that gives
instructors true PowerPoint integration. With JoinIn, instruc-
tors are no longer tied to their computers. Instructors can walk
about the classroom as they lecture, showing slides and col-
lecting and displaying responses with ease. There is simply no
easier or more effective way to turn a lecture hall into a per-
sonal, fully interactive experience for students. If an instructor
can use PowerPoint, they can use JoinIn on TurningPoint! It
contains poll slides and pre- and post-test slides for each chapter
in the text.
Image Library
This resource consists of digital copies of figures, tables, and
photographs used in the book. Instructors may use these JPEGs
to customize the PowerPoint slides or create their own Power-
Point presentations. The Image Library Key describes the images
and lists the codes under which the JPEGs are saved.
Instructor Guide
The Instructor Guide is organized according to the textbook
chapters and addresses key educational concerns, such as typical
stumbling blocks student face and how to address them.
MindTap
MindTap is the digital learning solution that powers students from
memorization to mastery. It gives instructors complete control of
their course—to provide engaging content, challenge every indi-
vidual, and build student confidence. Instructors can customize
interactive syllabi to emphasize priority topics as well as add their
own material or notes to the ebook as desired. This outcome-
driven application gives instructors the tools needed to empower
students and boost both understanding and performance.
21/03/22 4:13 PM
 Acknowledgments
We would like to begin by expressing our gratitude for the con-
tribution of Jonathan Ferrier. Jonathan has been an important
member of this team from the very beginning, and his passion,
insight, and guidance have informed and grounded our ongoing
journey to a more open, “multi-science” perspective that
acknowledges all Indigenous Peoples as having scientific knowl-
edge of the natural world that enriches more Eurocentric views.
We are very grateful for the quiet wisdom of Jean Becker, the
patient guidance and expertise of Jeff Baker, and the warmth and
insight of Erin Hodson. The three of you have been a source of
inspiration, guidance, and encouragement.
With great appreciation we acknowledge the contributions
of Chester Lawson and Elroy White of the Haíłzaqv (Heiltsuk)
Territory, Central Coast, British Columbia. Chester (Q'vúmkviá)
is Yím'ás (Haíɫzaqv hereditary chief). He is a community artist
and retired teacher. Chester has been creating art for over
70 years in several traditional and western art forms. Elroy
(Q'íxitasu) is Yím'ás, a potlatch historian, and an archaeologist
(Central Coast Archeology). We are grateful for the cultural
knowledge he generously shared about seaweed harvest and
processing and his work on assessing culturally modified trees
(CMTs). As an archeologist, Elroy approaches his research in a
way that combines academia and oral history; he calls this
approach Mnuxvit (to unite/become one).
We are deeply appreciative of the willingness of Elders Dan
Smoke and Mary Lou Smoke, Cliff Summers, John Brown, and
others to facilitate the Pipe Ceremony expressing our collective
desire for a positive and productive impact of this project through
strengthened relationships and respectful collaborations.
We are also grateful to the members of the fifth Canadian
edition Editorial Advisory Board and Student Advisory Board,
who provided us with valuable feedback and alternative per-
spectives (special acknowledgments to these individuals are
listed below).
We also thank Richard Walker at the University of Calgary
and Ken Davey at York University, who began this journey with
us but were unable to continue. We are very grateful to Heather
Addy of the University of Calgary for her significant contribu-
tions to the first three editions.
The authors are all indebted to Ivona Mladenovic of Simon
Fraser University for her excellent work on the Self-Test Questions
and to Johnston Miller, whose extensive background research
anchored our Concept Fix feature in the education literature.
We are profoundly grateful to Toni Chahley, Content
Development Manager, who was an insightful and reliably
cheerful constant on this project. We appreciate the warm pro-
fessionalism of Carmen Yu, Portfolio Manager, and Imoinda
Romain, Senior Content Production Manager. We thank
Christine Myaskovsky, Senior Intellectual Property Analyst,
11140_fm_ptg01.indd 31
and Betsy Hathaway, Senior Intellectual Property Project Man-
ager, for their hard work with the numerous visuals in the book
and Frances Robinson for her careful and thoughtful copy-
editing. Finally, we thank Scott Brayne, Marketing Manager,
for making us look good.
Paul Fam was a warm friend and a truly fearless leader
through the previous four editions of this text. The direction
and emphasis of this fifth Canadian edition was sparked by his
energy and enthusiasm and carries his vision forward.
It is never easy to be the family of an academic scientist.
We are especially grateful to our families for their sustained
support over the course of our careers, particularly during
those times when our attentions were fully captivated by bac-
teria, algae, fungi, parasites, snakes, geese, or bats. Saying “yes”
to a textbook project means saying “no” to a variety of other
pursuits. We appreciate the patience and understanding of
those closest to us that enabled the temporary reallocation of
considerable time from other endeavours and relationships.
Many of our colleagues have contributed to our develop-
ment as teachers and scholars by acting as mentors, collabora-
tors, and, on occasion, “worthy opponents.” Like all teachers,
we owe particular gratitude to our students. They have gath-
ered with us around the discipline of biology, sharing their
potent blend of enthusiasm and curiosity, and leaving us ener-
gized and optimistic for the future.
Indigenous Advisory Board
Jonathan Ferrier, Assistant Professor, Department of Biology,
Dalhousie University
Jeff Baker, Assistant Professor and Chair in Indigenous
Education, University of Saskatchewan
Jean Becker, Associate Vice-President, Indigenous Relations,
University of Waterloo
Erin Hodson, Educational Developer, Wilfrid Laurier University
Editorial and Student Advisory Boards
We were very fortunate to have the assistance of some extraordi-
nary students and instructors of biology across Canada who
provided us with feedback that helped shape this textbook into
what you see before you. As such, we would like to say a very
special thank you to the following people:
Editorial Advisory Board
Dora Cavallo-Medved, University of Windsor
Emma Despland, Concordia University
Heidi Engelhardt, University of Waterloo
Mark Fitzpatrick, University of Toronto
William Huddleston, University of Calgary
Sobia Iqbal, Wilfrid Laurier University
Amy Jeon, Kwantlen Polytechnic University
xxxi
21/03/22 4:13 PM
 Ivona Mladenovic, Simon Fraser University Western University
Ken Otter, University of Northern British Columbia Catherine Andary
Melissa Perrault, University of Guelph Deepshikha
Alexandra Pettit, University of Ottawa Tessa Fortnum
Roy Rea, University of Northern British Columbia Muhammad Hussain Jan
Ross Shaw, MacEwan University Helen Jiang
Edward Liu
Student Advisory Boards
Maryam Oloriegbe
University of Calgary Sofia Varon
Brooklyn Mattila Jonathan Wei
Korbin Nunez Michelle Yu
Gurleen Randhawa
Yvaine Stelmack
Rayyan Zuberi

xxxii ACKNOWLEDGMENTS

11140_fm_ptg01.indd 32 21/03/22 4:13 PM

 BIOLOGY OF THE CELL

VOLUME 1
Lebendkulturen.de/Shutterstock.com
Volvox is a genus of green algae. As a photosynthetic eukaryote, Volvox is a simple multicellular
organism that exists as spheres of two distinct cell types. Small non-reproductive cells surround
an interior that contains six to eight much larger reproductive cells.

What is life? We can make a list of the characteristics of living things, but why is it difficult to
simply define life? In the introductory chapter of this opening volume, we tackle this question
before exploring hypotheses around how life evolved some 4 billion years ago. Today, Earth is
teaming with life; some estimates peg the total number of species at over 1 billion (with most yet
to be described!). Yet, what this 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, every-
thing that is alive on Earth employs a variation on that remarkable innovation: the cell. No matter
whether that cell is communicating with other cells in the brain of a fruit fly, or capturing sun-
light 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.

11140_ch01_ptg01.indd 1 20/03/22 12:27 PM

 The invention of the microscope allowed scientists to
finally understand how living organisms were built from cells,
which to early scientists were astonishingly small. Further work
clearly delineated two major divisions in cell types: those
without a nucleus (prokaryotic) and those with a nucleus
(eukaryotic). And within both these groups there are clear sub-
divisions: for example, plants, fungi, and animal cells, in
eukaryotes.
The chapters of Unit 1 talk a lot about energy because
without it living cells would die. Like the non-living world, all
forms of life abide by the foundational laws of thermodynamics
and need to bring in energy and matter from the environment
to maintain their highly ordered state. In part, energy is
required to build complex things (e.g., proteins) out of simpler
things (e.g., amino acids). In addition to energy, the evolution
of life is tied to the development of a remarkable group of pro-
teins called enzymes, which when they get tied to a biochemical
reaction can increase its rate by 1010 times!
Another remarkable feature of cells that we dedicate a
chapter to in Unit 1 is membranes. These self-forming lipid
bilayers act as the gatekeepers of the cell: they allow certain
things in but keep other things out. How they do this is by
acting in concert with membrane-specific proteins that shuttle
molecules from one side to another. Membrane proteins also
play a remarkable role in transducing signals from outside the
cells and compartments to the inside. As we will see, the trans-
duction of signals associated with hormones, for example, can
profoundly affect cell function.
What you may not realize is that virtually all the energy
used by living systems comes ultimately from sunlight being
harvested and converted into a useable chemical form through
photosynthesis. This process evolved perhaps as early as 3.5
billion years ago and used photons of light energy to extract
electrons from water, releasing oxygen as a by-product. The rise
in oxygen in an atmosphere that previously had none led to an
explosion of life as the mechanism of cellular respiration
evolved that could use that oxygen and enabled cells to produce
huge amounts of energy.
The chapters of Units 2 and 3 are dedicated to molecular
biology and genetics, the central player being the gene. All cells
possess genes that are coded by the molecule DNA that,
2 V O LU M E 1 B I O LO G Y O F T H E C E L L
11140_ch01_ptg01.indd 2
through the process of transcription, get copied into RNA. All
cells contain ribosomes where some kinds of RNAs get
translated into proteins, the fundamental structural, func-
tional, and regulatory molecule of the cell.
Genes are stretches of DNA sequence in an organism that
collectively comprise a kind of library of information about
how a cell functions. Recent advances in technology have made
it relatively easy to determine the entire DNA sequence of an
organism, including individual humans. As a result, modern
biology is awash in the As, Ts, Gs, and Cs of DNA sequences
revealed by thousands of sequencing projects. New insights
into evolutionary history as well as gene structure and func-
tion are arising from bioinformatic analysis of such extensive
data sets.
The elegant double-strandedness of DNA, whereby two
long strands of nucleotides are held together by hydrogen bonds
formed between complementary base pairs, affords a straight-
forward mechanism for replication that was recognized early
on by Watson and Crick. Although conceptually simple, the
mechanism for unwinding the DNA double helix and polymer-
izing new complementary bases is rather complicated and
managed by a suite of interacting enzymes. Again, we see that
all DNA on the planet is replicated using variations on one
underlying strategy.
DNA genes provide the cell with needed RNA by tran-
scription. One remarkable feature of all protein-coding genes is
that, with minor exceptions, the information they carry is
specified by a universal code. That is, a gene from one organism
can be “understood” by any other organism, even if only dis-
tantly related: a gene from a spider can be expressed by a goat;
a gene from a jellyfish can be expressed in a flower. The field of
genetic engineering is devoted to developing the tools and
applications of this technology for moving genes from one
organism to another.
In a story that is about to come full circle, synthetic biolo-
gists have extensively customized naturally occurring cells and
have made important advances toward their ultimate goal of
creating novel life forms artificially in the lab. As students of
biology in the early 21st century, you can well expect to witness
a momentous event in Earth’s history, the creation of one life
form by another.
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11140_ch01_ptg01.indd 3 20/03/22 12:27 PM
 Chapter Roadmap

Defining Life and Its Origins

What is life, and what do we know about how it may have begun about 4 billion years ago?

1.1 1.2
What Is Life? The Chemical Origins of Life
Biology is the study of life, but a The conditions on early
simple definition of life is not easy to Earth allowed for the synthesis
come up with. of key molecules essential for life.

The Purple Pages provides

an overview of the key
molecules of life.

1.3
The Evolution of Information
Flow: DNA, RNA, Protein
1.4
RNA was likely the first of these three molecules to
evolve. It could carry information and The Development of Metabolism
act as a catalyst.
and the First Cells
Life may have evolved in alkaline hydrothermal vents,
The chapters of Unit 3 introduce an environment with a constant source of geothermal
major concepts related to DNA energy and mineral catalysts.
and gene expression.

Chapter 4 provides more

1.5 details on energy
and metabolism.

The Tree of Life

Recent research indicates that there are two
primary domains of life: Bacteria and Archaea.
Eukaryotes evolved from a lineage of Archaea. 1.6
In Chapter 19, we discuss the tree
of life and the discipline of Eukaryotes and the
phylogenetics in more detail.
Rise of Multicellularity
Eukaryotic cells have structural and
1.7 functional complexity not found in prokaryotic
cells. In addition to the nucleus, this
includes mitochondria and chloroplasts
The Fossil Record that are descended from bacteria.
Much of our understanding of the
history of life comes from the fossil record. In Chapter 2, we
However, the fossil record has a lot provide an overview
of gaps and does not provide a of both prokaryotic and
record of the earliest forms of life. eukaryotic cells.

11140_ch01_ptg01.indd 4 20/03/22 12:27 PM

 Triff/Shutterstock.com
Perseverance landed on Mars on February 18, 2021, to search for signs of life.

Defining Life and Its Origins

Why it matters… Finding evidence of life somewhere else would be the most profound dis-
covery in human history. On February 18, 2021, the NASA rover Perseverance landed on Mars and,
along with characterizing the Martian climate and geology, it is looking for signs of life. The landing
1
site of Perseverance is the 45-kilometre–wide Jezero Crater that a few billion years ago was the site of a
large lake. As the rover slowly roams over the Jezero lakebed, it employs an instrument with the
acronym SHERLOC that uses laser light to detect within the sediments the presence of organic mol-
ecules that, on Earth, we know are produced through biological activity. Detailed analysis of samples
that Perseverance collects will have to wait until they are sent back to Earth as part of a later mission.
A hope of many astrobiologists is that Perseverance comes across a surface feature that can’t
be attributed to anything other than ancient microbial life. On Earth, a good example of that is
stromatolites, which are sediment formations generated by the action of photosynthetic bacteria.
But therein lies one of the problems with searching for life elsewhere: we only know about life on
Earth, and maybe life on other planets is very different. And then, of course, what is life anyway?
It is actually really hard to come up with even a simple definition. While we continue to search
for life elsewhere, our understanding of life and how it may have developed is limited to
understanding how life may have developed on Earth.
The events we are going to discuss in this chapter may seem improbable: the synthesis of
organic molecules; the development of the first cells; the evolution of DNA, RNA, and proteins; the
building of metabolic pathways; and all the way to the evolution of multicellular eukaryotes.
Improbable? Perhaps. But we must keep in mind that these events took place over hundreds of
millions of years. As scientist and author George Wald of Harvard University put it, given so much
time, “the impossible becomes possible, the possible probable, and the probable virtually certain.”

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 1.1 What Is Life?
Picture a frog sitting in a pond, slowly shifting its head to follow
the movements of an insect flying nearby (Figure 1.1). Most of us
instinctively know that the frog is alive and the water is not. Or
is water alive? The atoms found in living and non-living entities
are the same. The oxygen and hydrogen atoms, for example,
aren’t any different within cells than they are in a pool of water
or a rock. As well, living cells may be extraordinary things but
they abide by the same fundamental laws of chemistry and
physics as the abiotic (non-living) world. Before we embark on a
discussion of how life may have developed billions of years ago
and the characteristics that early forms of life must have pos-
sessed it would be helpful if we had a clear notion of what dis-
tinguishes something that is living from something that is not
living.
1.1a Why It’s Hard to Define Life
Biology is the study of life, and the purpose of Biology: Exploring
the Diversity of Life is to provide you with an introduction to
many aspects of living things: from the unique features of bio-
logical molecules, reaching outward to cells, organisms, and the
interconnectedness that defines ecosystems.
It may surprise you that, while we can describe in consid-
erable detail the features that one organism has compared to
another, and certainly we can think about the things that living
things can do that non-living things can’t do, we struggle to
come up with a clear definition of what life actually is. Doesn’t
that seem a little odd? If you think about the other core disci-
plines often taught in first year, chemistry and physics, most of
the concepts in those disciplines are pretty easy to define, even
if they may not necessarily be easy to learn.
A major hurdle in coming up with a definition for life is the
pursuit of something that is universal. Ideally, the definition of
life should be acceptable to scientists and non-scientists alike,
and be useful and applicable across disciplines. As you will see,
the way a discipline defines life is intimately connected to the
questions that it explores. Let’s consider at three disciplines
Alexander Sviridov/Shutterstock.com
FIGURE 1.1 Northern leopard frog resting in a pond
6 CHAPTER 1 DEFInInG LIFE AnD ITs ORIGIns
11140_ch01_ptg01.indd 6
where defining life seems to be of fundamental importance.
First, let’s think about evolutionary biology, the study of how
life on Earth changes over time through processes such as nat-
ural selection and speciation. Second, there is the growing sci-
ence of astrobiology, which considers questions related to the
possibility of life beyond Earth and the factors necessary to
support life on other planets. Any definition of life that we can
come up with would need to apply to entities we may discover
(or meet) on other worlds. Third, there is the field of artificial
life, which uses computer simulations, robotics, and synthetic
biochemistry to simulate many of the attributes of life.
As shown in Figure 1.2, part of the problem with defining
life is that each of these groups of researchers approach life
from a slightly different perspective, and what one group thinks
is an essential aspect of life, another group may not.
One of the first people to take a crack at defining life was
Aristotle, who mentioned that it is something that “grows, is self-
sustaining and reproduces.” This seems like a pretty solid start;
the ability to make more of itself seems a central feature of all
forms of life. Another central feature seems to be the cell. Peering
into the primitive light microscopes in the 17th century, it soon
became obvious that all living things were formed of cells. As we
will discuss in detail in the next chapter, this became part of cell
theory: The cell was defined as the smallest unit of life and new
cells were derived only when old cells divided. So, for a long time,
if we couldn’t strictly define life, at least everyone could agree on
one universally accepted characteristic: the cell. We will see later
in this section how the discovery of viruses has muddled this
cell-centric view of life.
An entity is alive for An entity is alive for
evolutionary biologists only astrobiologists only if it
if it has properties A, B, and C. has properties B, D, and F.
An entity is alive for artificial-life
researchers only if it has
properties A, D, and E.
FIGURE 1.2 One of the issues with defining life is that, depending
upon the branch of biology where it is important, the properties of
life don’t completely match. For example, does life have to be composed
of cells, evolve, contain DNA, undergo metabolism, reproduce? Some of
these properties are important to some researchers and not important to
others. We have yet to arrive at a definition of life that is universally accepted.
20/03/22 12:27 PM
 In 1992, the NASA exobiology advisory board put forward
a working definition of life as being “a self-sustaining system
capable of Darwinian evolution.” As we will see in Chapter 16,
“Darwinian evolution” means the system can undergo the pro-
cess of natural selection. While this definition was not formally
adopted by NASA, it did help to define what types of chemistry
would constitute life on another planet. The problem with the
NASA definition is that there are many exceptions to it. There
are many clearly living entities that are unable to reproduce.
Think of a mule, which is the sterile hybrid offspring of a horse
and a donkey.
There are also entities that science deems “non-living” that
possess qualities of life such as reproduction, evolution, or
something else. For example, fire can certainly make more of
itself (reproduce) and it can grow and consume matter. For
decades, it has been known that mineral crystals can grow and
maintain a highly ordered structure not unlike an organism. It
was the physicist Erwin Schrödinger in 1945 who saw the
maintenance of order as one key defining property of life. As
we will see in Chapter 3, maintaining order (or low entropy) is
a central reason for why life undergoes energy-requiring
metabolism. Yet, within the past few years, researchers have
shown that energy from light can be used to drive lifelike, self-
organizing behaviours (think schooling fish or flocking birds)
in particles. Published in the prestigious journal Science, the
authors made reference to these as “living crystals.” As well,
many computer viruses can reproduce and change over time
(e.g., evolve), which, as we will see, are two key attribues.
1.1b Viruses Complicate Our Idea of Life
Whether or not viruses should be considered life has been an
argument that goes back to when they were first visualized
early in the 20th century, after the advent of the electron
microscope. Viruses, which we will discuss in detail in
Chapter 25, don’t have their own metabolism; they don’t respire
or carry out photosynthesis. But, like traditional forms of cel-
lular life, they do contain nucleic acid and replicate, and they
can evolve rapidly by natural selection. But they can’t complete
their lifecycle without invading living cells, whether that be a
bacterium or a human (Figure 1.3). Taking up residence within
living cells is required for a virus because it does not have its
own protein-making factories—ribosomes and key enzymes
required for it to replicate. Because of this last fact, many put
viruses clearly in the “not life” camp. But if viruses are not con-
sidered life because they are parasitic, what about Rickettsia?
This bacterium is considered to be alive despite its inability to
live outside a host cell.
Things became more complicated on the virus front in
1992 with the discovery of a virus initially thought to be a bac-
terium. Named mimivirus for “mimicking microbe,” no one
had ever observed a virus that was so large you could visualize
it using conventional light microscopy. Its genome, at 1.2 mil-
lion base pairs, far exceeds the size of any other known virus
11140_ch01_ptg01.indd 7
Image courtesy of John Wertz, Yale University
FIGURE 1.3 Bacteriophage infecting a bacterium. Notice the
bacteriophages on the cell surface as well as inside the bacterium. A
bacteriophage is a type of virus, and most scientists do not consider viruses
to be alive.
and is larger than many bacteria. Its genome has been predicted
to encode over 900 proteins, many of them associated with
functions never before seen in a virus. This includes proteins
central to protein synthesis and metabolism, two hallmarks of
life. However, like other viruses, mimivirus is still dependent
upon host cell ribosome for protein synthesis. Clearly, if mimi-
virus is not allowed to be a member of the life “club,” it stands
right outside the door.
1.1c Seven Characteristics Shared
by All Cellular Life Forms
From the above discussion it is pretty clear that we don’t have a
universally accepted definition of life. In fact, a growing number
of researchers reject the need for a common definition. Many
want a “theory of living systems” that can fully describe life (its
origins and characteristics) without forcing a strict definition.
Others argue that, since we are constantly discovering new spe-
cies and have yet to detect evidence of life on another planet,
trying to strictly define it is silly because we simply don’t under-
stand enough about it.
For now, we need to make a compromise. While we can
use the word entity to include a wide range of self-replicating
and evolving systems, we are going to reserve the word organism
for forms of life that are made of cells. It is cellular life that is
going to hold our attention through the next 44 chapters of this
textbook.
Without a strict definition of life, organisms on Earth all
possess a key list of attributes. As detailed in Figure 1.4, all life
displays order, harnesses and utilizes energy, reproduces,
responds to stimuli, exhibits homeostasis, grows and develops,
and evolves.
STUDY BREAK QUESTIONS
1. Why is it difficult to arrive at a definition for life?
2. Why is a virus often considered not to be alive?
CHAPTER 1 DEFInInG LIFE AnD ITs ORIGIns 7
20/03/22 12:27 PM
 a. Display order b. Harness and utilize energy c. Reproduce

Steve Byland/Shutterstock.com

SCIMAT/Science Source
harmeet/StockXchng
d. Respond to stimuli e. Exhibit homeostasis f. Grow and develop g. Evolve

Dr. Jeremy Burgess/Science Source

Stanislav Duben/Shutterstock.com
Karin Duthie/Alamy Stock Photo
Uwe Krejci/Stone/Getty Images
FIGURE 1.4 Although it is hard to strictly define life, all cellular forms of life on Earth display these seven characteristics. (a) Display order.
All forms of life, including this flower, are arranged in a highly ordered manner, with the cell being the fundamental unit that exhibits all properties of life.
(b) Harness and utilize energy. Like this hummingbird, all forms of life acquire energy from the environment and use it to maintain their highly ordered state.
(c) Reproduce. All organisms have the ability to make more of their own kind. Here, some of the bacteria have just divided into two daughter cells.
(d) Respond to stimuli. Organisms can make adjustments to their structure, function, and behaviour in response to changes to the external environment.
A plant can adjust the size of the pores (stomata) on the surface of its leaves to regulate gas exchange. (e) Exhibit homeostasis. Organisms are able to regulate
their internal environment such that conditions remain relatively constant. Sweating is one way in which the human body attempts to remove heat and
thereby maintain a constant temperature. (f) Grow and develop. All organisms increase their size and/or number of cells. Many organisms also change over
time. (g) Evolve. Populations of living organisms change over the course of generations to become better adapted to their environment. The snowy owl
illustrates this perfectly.

1.2 The Chemical Origins of Life life arose out of a mixture of molecules that existed on early
Earth. In this section, we present hypotheses for how biologi-
One of the tenets of cell theory states that cells arise only from cally important molecules could have been synthesized on early
the growth and division of preexisting cells. This tenet, which Earth in the absence of life.
we will discuss in detail in the next chapter, has probably been
true for a few billion years, yet there must have been a time 1.2a Earth Is 4 600 000 000 Years Old
when this was not the case. There must have been a time when
Earth was formed about 4.6 billion years ago, at the same time
no cells existed, when there was no life. It is thought that, over
as the rest of the planets in the solar system. The age of our
the course of millions of years, cells with the characteristics of
planet has been arrived at using the technique of radiometric
Oxygen produced dating, which looks at specific isotopic ratios in rocks and
January 1 by photosynthetic knowledge of their rate of decay.
Earth bacteria December 31, The decay of uranium to lead, in
forms Earliest 11:43 p.m.
particular, has been used to age
life Modern
humans Earth. To give us some sense of
First
eukaryotes appear just how long 4.6 billion years is,
Earliest
land as well as the relative timing of
plants some major events in the his-
January February March
April First tory of life on Earth, Figure 1.5
Ma animals
y condenses the entire history of
Jun
e Earth into a unit of time that we
July er
em b are more familiar with: 1 year.
Aug Dec
ust
m b e r With 4.6 billion years con-
Septemb Nove
er October densed into a single year, each
day represents an interval of
FIGURE 1.5 The history of Earth condensed into one year 12.6 million years!

8 CHAPTER 1 DEFInInG LIFE AnD ITs ORIGIns

11140_ch01_ptg01.indd 8 20/03/22 12:27 PM

 Using our condensed version of the history of Earth, we In comparison to the proposed reducing atmosphere of
set the date of the formation of Earth as January 1 at 12:00 a.m. early Earth, today’s atmosphere is classified as an oxidizing
We will discuss this in detail later but, based on chemical evi- atmosphere. The presence of high levels of oxygen prevents
dence, life may have started as early as 4 billion years ago. This complex, electron-rich molecules from being formed because
translates to mid-March in our one-year calendar. The first oxygen is a particularly strong oxidizing molecule and would
clear fossil evidence of prokaryotic cells occurs in late March, itself accept the electrons from organic molecules and be
or about 3.5 billion years ago. Fossil evidence of eukaryotes reduced to water. Apart from allowing for the buildup of
has been dated to about 2 billion years ago, which is not until electron-rich molecules, the lack of oxygen in the young atmo-
early July using our 1-year analogy. Perhaps surprisingly, ani- sphere also meant that there was no ozone (O3) layer, which
mals do not make an appearance until mid-October (about only developed after oxygen levels in the atmosphere began to
525 million years ago) and land plants until the following increase. Both Oparin and Haldane hypothesized that, without
month. The extinction of dinosaurs, which was completed by the ozone layer, energetic ultraviolet light was able to reach the
about 65 million years ago, does not occur until late December. lower atmosphere and, along with abundant lightning, pro-
What about humans? We may think humans, Homo sapiens, vided the energy needed to drive the formation of biologically
have been around a long time but, relative to other forms of important molecules.
life, the roughly 150 000 years that modern humans have Experimental evidence in support of the Oparin–Haldane
existed is a very short period of time—a blip on our time scale. hypothesis came in 1953, when Stanley Miller, a graduate
Using our year analogy, modern humans have existed only student in the lab of Harold Urey at the University of Chicago,
since December 31; more precisely, since December 31 at created a laboratory simulation of a reducing atmosphere.
11:43 p.m.! Miller mixed the gasses hydrogen, methane, and ammonia in a
closed apparatus and exposed the gases to an energy source in
the form of continuously sparking electrodes (Figure 1.6). Water
1.2b Biologically Important Molecules Can vapour was added to the “atmosphere” in one part of the appa-
Be Synthesized outside Living Cells ratus and subsequently condensed back into water by cooling
in another part. After running the experiment for one week,
All organisms are composed of four classes of essential macro- Miller found a large assortment of organic compounds,
molecules: nucleic acids, proteins, lipids, and polysaccharides including urea; amino acids; and lactic, formic, and acetic acids
(see The Purple Pages for an overview of these molecules). These in the condensed water. In fact, as much as 15% of the carbon
molecules are constantly being synthesized within cells by var- that was originally in the methane at the start of the experi-
ious biochemical pathways and metabolic processes. If at least ment ended up in molecules that are common in living
some of these molecules are an absolute requirement for life, organisms.
then simple forms of them must have come before the first cells;
they must have been produced in the absence of life, what is
referred to as abiotic synthesis.
When the earliest forms of life developed about
∼4.5 billion years ago
4 billion years ago, Earth was vastly different from
Electrodes
what it is today. The atmosphere probably con- Spark
tained an abundance of water vapour from the discharge Simulated early
evaporation of water at the hot surface, as well as Inorganic Earth’s atmosphere
large quantities of hydrogen (H2), carbon dioxide molecules CH4
simulating
(CO2), ammonia (NH3), and methane (CH4). You early Earth’s
NH3
Gases
might be surprised to know there was an almost atmosphere H 2O
H2
complete absence of oxygen (O2) in the early atmo- Water out
sphere. In the 1920s, two scientists, Aleksandr
Condenser
Oparin and John Haldane, independently proposed
Water in Boiling water to produce
that organic molecules, essential to the formation of
Water water vapour, and using a
life, could have formed in the atmosphere of early condenser to form water
droplets
Earth. A critical aspect of what is known as the droplets, simulates the
Water containing water cycle on early Earth.
Oparin–Haldane hypothesis is that the early atmo- organic compounds
sphere was a reducing atmosphere because of the Liquid water in trap
presence of large concentrations of molecules such Boiling water
as hydrogen, methane, and ammonia. These mole-
FIGURE 1.6 The Miller–Urey experiment. Using this apparatus, Stanley Miller, a
cules possess chemical bonds that allow them to graduate student, demonstrated that organic molecules can be synthesized under
enter into reactions with one another that would conditions simulating primordial Earth.
yield larger and more complex organic molecules. Source: Based on S. L. Miller, 1955

CHAPTER 1 DEFInInG LIFE AnD ITs ORIGIns 9

11140_ch01_ptg01.indd 9 20/03/22 12:27 PM

 Other chemicals have been tested in the
Miller–Urey apparatus, including hydrogen cya-
nide (HCN) and formaldehyde (CH 2O), which
are also considered to have been among the sub-
stances formed in the primitive atmosphere.
When cyanide and formaldehyde were added to Microscopic
layers of clay
the simulated atmosphere in the apparatus, all
the building blocks of complex biological mole-
cules were produced: amino acids; fatty acids; the
purine and pyrimidine components of nucleic
acids; sugars such as glyceraldehyde, ribose, glu-
cose, and fructose; and phospholipids, which
form the lipid bilayers of biological membranes.
In the decades since the first Miller–Urey

experiment, considerable debate has developed in
the scientific community over whether Earth’s
atmosphere held enough methane and ammonia
FIGURE 1.7 Clay surfaces catalyze polymerization. The charged, microscopic,
for it to be considered a reducing atmosphere. layered structure of clay allows for the formation of relatively short polymers of proteins
Some geologists have suggested that, based on and nucleic acids.
analysis of volcanic activity, primitive Earth con-
tained other gases that made it probably some-
what less reactive. And as we will see that, later in this
chapter, many scientists have looked to the oceans as the
most likely place where abiotic synthesis took place and life
most likely started. Regardless of these lingering debates,
the significance of the Miller–Urey experiment cannot be
overstated. It was the first experiment to demonstrate the
abiotic formation of molecules critical to life, such as amino
acids, nucleotides, and simple sugars, and it showed that
they could be produced relatively easily. This remarkable
finding laid the groundwork for further research into the
origins of life.
1.2c Life Requires Polymers
Primordial Earth contained very little oxygen and, because of
this, complex organic molecules could have existed for much
longer than would be possible in today’s oxidizing world. Even if
they did accumulate on early Earth, molecules such as amino
acids and nucleotides are monomers, which are simpler and
easier to synthesize than the key chemical components of life,
such as nucleic acids and proteins, which are polymers—macro-
molecules formed from the bonding together of individual
monomers. Nucleic acids are polymers of nucleotides, proteins
are polymers of amino acids, and polysaccharides (starch, cel-
lulose) are polymers of simple sugars. Polymers are synthesized
by dehydration synthesis, which is discussed in The Purple
Pages.
Today, the synthesis of proteins and nucleic acids requires
protein-based catalysts called enzymes and results in macro-
molecules that often consist of hundreds to many thousands
of monomers linked together. So, how do you make the poly-
mers that are required for life without sophisticated enzymes?
Andrew Swift
Nucleotides undergoing
polymerization
A working hypothesis to address this question needs to be
built from the notion that the very earliest forms of life must
have been very simple, far simpler than a modern bacterium,
for example. Scientists hypothesize that a polymer that con-
sists of even 10 to 50 monomers may have been of sufficient
length to impart a specific function (like a protein) or store
sufficient information (like a nucleic acid) to make their for-
mation advantageous to an organism. It is, however, doubtful
that polymerization could have occurred in the aqueous envi-
ronment of early Earth, as it would be very rare for mono-
mers to interact precisely enough with one another to
polymerize. It is more likely that solid surfaces, especially
clays, could have provided the type of environment necessary
for polymerization to occur (Figure 1.7). Clays consist of very
thin layers of minerals separated by layers of water only a few
nanometres thick. The layered structure of clay is also
charged, allowing for molecular adhesion forces to bring
monomers together in precise orientations that could more
readily lead to polymer formation. Clays can also store the
potential energy that may have been used for energy-requiring
polymerization reactions. This clay hypothesis is supported
by laboratory experiments that demonstrate that the forma-
tion of short nucleic acid chains and polypeptides can be syn-
thesized on a clay surface.
STUDY BREAK QUESTIONS
1. For understanding the origins of life, what was the significance
of the Miller–Urey experiment?
2. What is the difference between a reducing atmosphere and an
oxidizing atmosphere?

10 CHAPTER 1 DEFInInG LIFE AnD ITs ORIGIns

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 1.3 The Evolution of DNA RNA Protein

Information Flow:
RNA, DNA, Protein
In the previous section, we discussed how
processes present on early Earth could have
generated macromolecules crucial to the
development of life. However, if we are to
develop a comprehensive model for the
origin of life, we need to explain the evolu-
tion of three key properties of a cell that go
well beyond the synthesis of macromole-
cules. First, a mechanism to store informa-
tion that can be replicated and passed on to
daughter cells must have developed. Second, Information is The information The information in
energy-transforming chemical reactions stored in DNA. in DNA is copied RNA guides the
must have evolved that would have enabled into RNA. production of proteins.
primitive cells to capture energy from their FIGURE 1.8 The central dogma. Information in DNA is used to synthesize proteins through an
surroundings and use it to do work. Third, RNA intermediate. How did such a system evolve when the product, proteins, is required in
these processes would need to take place modern-day cells to catalyze each step?
within defined compartments (e.g., cells)
that are distinct and separate from the
environment. our understanding of how such a system may have evolved
In this section, we discuss the first of these: the develop- came in the early 1980s when Thomas Cech and Sidney Altman,
ment of a genetics system that would allow the passing of infor- working independently, discovered a group of RNA molecules
mation to new cells following cell division. that could themselves act as catalysts. This group of RNA cata-
lysts, called ribozymes, are found in all types of cells. They are
not as common as enzymes, but they carry out critical reac-
1.3a RNA Can Carry Information tions related to the control of gene expression. One type of ribo-
zyme, for example, catalyzes the removal of introns from newly
and Catalyze Reactions synthesized RNA molecules; other ribozymes can cleave spe-
As we will discuss in detail in Chapter 11, DNA (deoxyribose cific messenger RNA (mRNA) molecules, causing their degra-
nucleic acid) is the molecule that provides every cell with the dation. The precise catalytic property of a ribozyme is
instructions necessary to function. The information contained determined by its shape, which is based on the hydrogen
in a sequence of nucleotides in DNA, called a gene, is copied bonding between specific nucleotides of the RNA molecule.
into a unique molecule of RNA. Some of these RNA molecules The link between nucleotide sequence, shape, and thus func-
provide information for the synthesis of specific proteins. Even tion is analogous to an enzyme. The specific reaction catalyzed
the simplest bacterium today contains thousands of genes, RNA by an enzyme is dependent on its precise three-dimensional
molecules, and proteins. The flow of information from DNA to shape, which is the result of its unique amino acid sequence. All
RNA to protein is common to all forms of life and is referred to ribozymes recognize their target by specific base pairing
as the central dogma (Figure 1.8). Each step of the information between the ribozyme nucleotide sequence and the nucleotide
flow requires the involvement of enzymes that catalyze the sequence of the target molecule (Figure 1.9).
transcription of DNA into
RNA, and the translation of Ribozyme
the RNA into protein.
A fundamental question
about the flow of information
from DNA to RNA to protein
is How did such a system
evolve when the final prod- Messenger RNA Ribozyme-mediated cut Cut (cleaved) messenger
ucts, proteins, are required to introduced into RNA message RNA molecules
catalyze each step (e.g., tran- FIGURE 1.9 Ribozyme. An example of a ribozyme binding to an RNA molecule and catalyzing its breakage. Within
scription, translation) of the a modern-day cell, such reactions help control gene expression by altering the abundance of functional messenger
process? A breakthrough in RNA (mRNA) molecules.

CHAPTER 1 DEFInInG LIFE AnD ITs ORIGIns 11

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yolk of an egg and the beaten white. Form into small balls. Drop into
boiling water. Boil two or three minutes.

Quenelles of Chicken, Game, Hare or Rabbit

4 ozs. meat
2 ozs. bread-crumbs
2 ozs. butter
1 whole egg and 1 extra yolk
Chop and pound the meat. Soak the bread-crumbs in a little milk
or broth. Mix all thoroughly together. Season. Pass through a sieve.
Form into balls. Drop into boiling water or broth and simmer for three
minutes.
The best meat should always be reserved for making quenelles.

Quenelles of Marrow
4 ozs. marrow
4 ozs. fine bread-crumbs
1 egg
¹⁄₂ tea-spoonful finely-chopped parsley
Mix all the ingredients thoroughly. Season. Roll in the hand in
small balls. Boil in a little broth for fifteen minutes.

Rice
1 cup of Carolina rice
2 quarts boiling water
1 table-spoon salt
Wash a cup of rice thoroughly. Drain it. Throw it into a large sauce-
pan of salted boiling water and let it boil as fast as possible for
twenty minutes. Do not stir. Drain. Put into cold water for ten
minutes. Drain again. When required warm it by steaming, or set it in
the oven, leaving the door open.

Savoury Rice
(To serve with Clear Soup)
Prepare the rice as above. Add to it one cup of rich stock which
has been highly seasoned. Steam to warm. Add a table-spoon of
butter just before serving.
Or,
Add a table-spoon of chopped onion which has been fried a rich
yellow in a table-spoon of butter, to the cooked rice. Moisten with a
cup of stock and steam for ten minutes.

Rice Balls
(For Cream of Rice or Clear Soups)
¹⁄₄ lb. Carolina rice
1 oz. butter
1 oz. grated Parmesan
2 yolks of eggs
1 whole egg
Boil the rice until quite soft. Drain it. Put it in a sauce-pan with the
butter, cheese and yolks. Stir continually for five minutes. Season.
Take off the fire. Turn out of the sauce-pan to cool. When cold, make
into small balls. Beat the whole egg. Roll the balls first in a little flour,
then in the egg. Fry in very hot lard till a rich yellow.
 Sauces
There is, of course, no end to sauces, and in a book of this size it
is impossible to do justice to their variety. Enough are, however, I
hope, given in the pages that follow for ordinary needs.
It is of the highest importance in making sauces that the materials
used should be of the best. Fresh butter and the finest olive oil
should be used.
When adding the yolks of eggs to sauces it is best to do so in a
bain marie (i.e. to stand the sauce-pan in which the sauce is being
made, inside a larger one full of boiling water), as they must never
be allowed to boil, and a quick fire easily burns them.
For thickening sauces, etc., see remarks on soup on p. 12.
 Hot Sauces for Fish
PAGE
Black Butter 112
Dutch Sauce 112
Genoese Sauce 113
Italian Sauce 113
Maître d’Hotel Sauce 114
Melted Butter 114
Anchovy Sauce 114
Cucumber Sauce 114
Egg Sauce 114
Shrimp Sauce 114
Oyster Sauce 115
Sauce Hollandaise 115

Black Butter
(For Skate, grilled Mackerel)
1 gill vinegar
4 ozs. butter
Several small parsley leaves
Small piece of bay leaf
Boil the vinegar with the bay leaf until it is considerably reduced.
Heat the butter in a pan until it becomes brown. Add the parsley
leaves. Let them fry for a moment. Skim the butter.
 Remove the bay leaf from the vinegar. Add a little salt and pepper.
Pour the butter and parsley leaves into it. Mix and serve.

Dutch Sauce
Butter, size of an egg
1 tea-spoon flour
¹⁄₂ pint milk or cream
Juice of half a lemon
2 yolks of eggs
Melt the butter in a sauce-pan. Stir in the flour and mix till perfectly
smooth. Add the milk or cream. Boil for two or three minutes. Add
lemon juice, and just before serving, stir in the two yolks. After which
do not allow the sauce to boil.

Genoese Sauce
(For Fillet of Sole)
1 oz. butter
2 table-spoons olive oil
2 yolks of eggs
1 table-spoon vinegar
Put the oil and butter into a sauce-pan on the fire and stir till the
butter is melted. Beat the yolks slightly. Add the vinegar to them.
Season. Directly the butter is melted add the yolks and vinegar,
stirring continually over a bain marie until the sauce thickens. Half a
tea-spoonful of mustard may be added.

Italian Sauce
(For Mackerel, etc.)
2 table-spoons olive oil
1 oz. butter
 6 chopped mushrooms
1 shallot, finely chopped
1 tea-spoon chopped parsley
1 clove
1 wine-glass white wine
10 drops Liebig’s extract of meat
Put the butter and oil into a sauce-pan. Add the mushrooms,
shallot, parsley and the clove. Cook for a few minutes. Add the wine
and Liebig. Simmer gently for forty minutes. Season. Pass through a
sieve.

Maître d’Hotel
4 ozs. butter
¹⁄₂ pint milk
1 tea-spoon flour
1 dessert-spoon finely chopped parsley
Juice of a lemon
Mix the flour and butter together till smooth. Melt in a sauce-pan.
Add the boiling milk. Let all boil for three or four minutes, stirring
constantly. Add the parsley and lemon juice.

Melted Butter
1 tea-spoon flour
4 ozs. butter
1 gill boiling milk or water
Mix the flour and butter thoroughly in a basin. When perfectly
smooth put in a sauce-pan. Add to it the boiling milk or water. Let it
boil for two or three minutes. Stir continually from left to right.
Season.
To this sauce the raw yolk of an egg or a finely chopped hard
boiled egg, shrimps, a little essence of anchovy, or a table spoon of
grated cucumber may be added; when it becomes egg, shrimp,
anchovy or cucumber sauce. To the cucumber sauce add a tea-
spoonful of lemon juice.

Oyster Sauce
2 doz. oysters
3 ozs. butter
1 tea-spoon flour
¹⁄₂ pint cream
1 coffee-spoon lemon juice
Prepare the oysters and stew them in their own juice and the
butter until plump and tender. Mix the flour with the cream, until
perfectly smooth. Bring to the boil and let it boil two or three minutes.
Add it to the oysters, etc. Stir quickly together. Season with salt, a
little cayenne and the lemon juice.

Sauce Hollandaise
4 table-spoons vinegar
1 blade mace
1 tea-spoon flour
Yolks of 4 eggs
3 ozs. butter
Season the vinegar, add to it the flour and mix perfectly smooth.
Add the mace. Bring to the boil and boil for two or three minutes.
Take off the fire, and take out the mace. Add the butter cut in small
pieces, and the well-beaten yolks. Stir continually, in one direction,
over a bain marie. Serve directly the butter is melted.
 Hot Sauces for Roasts, Steaks,
Cutlets, etc.
PAGE
Brown Sauce 118
Cucumber Sauce 118
Dutch Horse-radish Sauce 119
Maître d’Hotel I. 120
” ” II. 120
Mushroom Sauce 121
Onion ” 121
Sauce Béarnaise 122
Sauce for Chops and Steaks 122
Sauce Piquante au Citron 123
Sauce Robert 123
Sauce Vinaigrette 124
Tomato Sauce 124

Brown Sauce or Cullis

3 lbs. lean veal
1 lb. raw lean ham
1 oz. butter
6 mushrooms chopped
1 carrot chopped
1 onion chopped
Rind of a lemon
 Small bouquet of herbs
1 tea-spoon allspice
1 quart brown stock
¹⁄₄ lb. brown roux
Slice the veal and ham. Add the vegetables, spice, lemon rind and
herbs, and brown slightly in a sauce-pan with the butter. Add the
stock and brown roux (see p. 13). Boil ten minutes. Stir continually.
Put through a tammy.

Cucumber Sauce
1 cucumber
2 table-spoons brown stock
1 oz. butter
1 table-spoon chopped parsley
Juice of half a lemon
¹⁄₂ pint brown sauce
Peel and split the cucumber lengthwise in four pieces. Take out
the seeds. Cut in small pieces. Put into salted water and boil gently
for seven minutes. Take off and drain. Melt the butter and add to it
the stock, cucumber and parsley. Cook gently for half-an-hour. Add
the brown sauce and lemon juice.

Dutch Horse-radish Sauce

(For Roast Beef or Steak)
1 tea-cup horse-radish
¹⁄₂ pint water
3 ozs. butter
3 table-spoons flour
1 gill cream
4 yolks of eggs
3 table-spoons elder vinegar
 Scrape the horse-radish very finely, and boil it for ten minutes in
water. Drain off the water. Cook the horse-radish with the butter and
flour for four minutes. Add the water in which the horse-radish was
boiled, stirring continually. Heat. Take off the fire. Add the hot cream
and then the beaten yolks. Beat well together. Add pepper, salt and
the vinegar.

Maître d’Hotel—I
2 ozs. butter
1 table-spoon chopped parsley
Juice of half a lemon
Melt the butter. Skim it. Add the parsley (and, if liked, a little finely
chopped shallot), salt, pepper and lemon juice.

Maître d’Hotel—II
4 shallots
1 tea-spoon chopped parsley
1 tea-spoon chopped fennel
1 dozen mushrooms
2 ozs. butter
¹⁄₂ pint brown sauce or béchamel
Chop the shallots. Put them with the parsley, fennel and
mushrooms in a sauce-pan in which the butter has been melted.
Cook gently for five minutes. Add the brown sauce or béchamel (see
pp. 118 and 126). Boil ten minutes. Season and add a squeeze of
lemon juice.

Mushroom Sauce
2 dozen small mushrooms
1 oz. butter
1 table-spoon flour
 1 pint good gravy
¹⁄₂ a lemon
Cook the mushrooms in the butter until brown and tender. Add the
flour. Stir well in and brown. Pour the gravy over the mushrooms.
Boil three minutes. Season and add a little lemon juice.

Onion Sauce
(For Roast Mutton)
4 onions
¹⁄₂ pint melted butter (see p. 114)
Slice and chop the onions finely. Boil until tender. Drain and add to
the hot melted butter. Season. If preferred, the onion can be first
passed through a fine sieve and then added to the melted butter.

Sauce Béarnaise
5 yolks of eggs
2 ozs. butter
1 table-spoon chopped tarragon
1 dessert-spoon vinegar
Put the yolks in a sauce-pan, in a bain marie, and stir into them
one ounce of butter. As soon as the eggs begin to thicken, take off
the fire. Add another ounce of butter, the tarragon and vinegar. This
sauce should be of the consistency of a mayonnaise. Serve with
roast meats.

Sauce for Chops or Steak

2 table-spoons red wine
2 table-spoons ketchup
1 tea-spoon butter
1 tea-spoon vinegar
 Stir altogether in a sauce-pan. Season and serve very hot.

Sauce Piquante au Citron

(For Calf’s Head)
2 table-spoons chopped onions
1 oz. butter
1 table-spoon flour
1 gill white stock
1 gill white wine
1 lemon
Fry the onion in the butter, with the flour, until a rich yellow. Add to
it the stock, which should be boiling, and the wine. Stir together. Add
the juice of the lemon and a little of the grated rind. Simmer for
quarter of an hour. Strain through a fine sieve.

Sauce Robert
(For Pork)
3 onions
1 gill rich brown gravy
1 tea-spoon made mustard
1 tea-spoon vinegar
2 ozs. butter
1 table-spoon flour
Chop the onions. Fry them in the butter. Add the flour. Mix quite
smooth. Add the gravy, salt and pepper. Simmer for half-an-hour.
Skim. Add the mustard and vinegar. Serve with pork.

Sauce Vinaigrette
4 table-spoons vinegar
1 bay leaf
1 table-spoon brown sauce
 1 table-spoon chopped shallots
2 table-spoons chopped gherkins
1 table-spoon capers
1 table-spoon chopped parsley
1 oz. butter
Boil the vinegar for quarter of an hour with the bay leaf. Add the
sauce (see p. 118). Simmer five minutes. Remove the bay leaf. Add
the shallots (which should have been previously cooked in the butter
and allowed to drain upon a sieve), capers, gherkins and parsley.

Tomato Sauce
6 tomatoes
¹⁄₂ an onion chopped
1 clove
1 slice of ham
1 gill rich brown gravy
1 table-spoon brown roux
Remove the seeds from the tomatoes. Stew them with the onion,
ham and clove in an enamel sauce-pan until well cooked. Rub
through a tammy. Return to the sauce-pan. Add the gravy and brown
roux (see p. 12). Simmer for quarter of an hour.
Hot Sauces for Fowls, Ducks, Rabbits,
etc.
PAGE
Apple Sauce 126
Béchamel Sauce 126
Bread Sauce 127
Celery Sauce 127
Gooseberry Sauce 128
Lemon Sauce 128
Parsley Sauce 129
Sauce à la Reine 129
White Sauce 130

Apple Sauce
Set the required quantity of sour apples, pared, cored and sliced,
in a small pan inside a large sauce-pan containing boiling water. Let
the water boil quickly until the apples are done. Mash them and add
sugar to taste.
Or,
Pare, quarter and remove the core of several sour apples. Put
them a sauce-pan with a little water. Boil up quickly. Do not stir until
cooked. Then add sugar and mash.

Béchamel
 1 lb. veal
2 slices ham
2 pints water
¹⁄₄ lb. mushrooms
1 onion
Bouquet of herbs
5 table-spoons white roux
1 pint of cream
Slice the veal, ham, mushrooms and onion and stew them gently
for an hour and a half in the water. Thicken with the roux (see p. 12).
Add the cream. Boil for two or three minutes, stirring continually.
Strain.

Bread Sauce

¹⁄₂ pint milk

1 tea-cup bread-crumbs
1 onion
2 pepper-corns
1 tea-spoon butter
Slice the onion and boil it in the milk with the pepper-corns until
very tender. Strain off the milk and add it to the bread-crumbs which
should be made from stale bread and be very finely grated. Allow the
sauce to stand covered for a few minutes. Add the butter. Stir in
thoroughly. Season and serve very hot.

Celery Sauce
1 large head of celery
¹⁄₂ pint milk or cream
1 table-spoon white roux
Use the best of the celery only. Cut it in small pieces. Cook it in
water until very tender. Put through a sieve. Add it to the cream or
milk. Thicken with a small table-spoon white roux (see p. 12).
Season.

Gooseberry Sauce
(For Duckling or Goose)
1 gill spinach juice
¹⁄₂ pint stock
¹⁄₂ pint gooseberries
1 table-spoon sugar
1 tea-spoon butter
Cook the gooseberries till tender. Rub them through a sieve. Put
them in a sauce-pan on the fire. Add the sugar (more if preferred)
and butter. When thoroughly mixed, add the stock with which the
spinach juice (see p. 104) has been mixed. Make very hot.

Lemon Sauce
(For Rabbit or Fowl)
1 lemon
1 liver of fowl or rabbit
¹⁄₂ pint melted butter
1 table-spoon chopped parsley
Cook the liver, pound it and put it through a sieve. Peel the lemon,
cut the inside, from which the pips must be removed, into very small
dice-shaped pieces. Add the lemon and liver to the melted butter.
Heat gently, but do not boil. Add the parsley.

Parsley Sauce
Small bunch of parsley
¹⁄₂ pint melted butter
 Boil the parsley for five minutes. Drain. Chop finely. Add to the
melted butter.
Or,
To one gill of water in which a fowl has been boiled, add one gill of
cream, one dessert-spoon white roux (see p. 12), seasoning and the
boiled and chopped parsley.

Sauce à la Reine

¹⁄₂ pint veal stock

¹⁄₄ lb. mushrooms
Small bouquet of herbs
¹⁄₂ an onion
¹⁄₂ pint cream
Breast of a fowl
Juice of half a lemon
1 tea-spoon flour
Let the veal stock simmer for half-an-hour with the mushrooms,
onion, and herbs. Then strain. Thicken with the flour. Boil two or
three minutes. Add the boiling cream. Set back on the fire and add
the finely pounded breast, lemon juice and seasoning. Do not allow
the sauce to boil after the chicken has been added.

White Sauce
1 gill veal or chicken stock
1 gill cream
Juice of half a lemon
Juice of half a Seville orange
Mix all together. Heat gently, stirring continually. Season.
 Hot Sauces for Game, etc.
PAGE
Cream Sauce 132
Game Sauce 132
German Sauce 133
Madeira Sauce 133
Orange Sauce 134
Sauce Poivrade 134
Sour Cream Sauce 135

Cream Sauce
The gravy from two roasted birds
1 gill cream
Stir the cream into gravy of the birds with which it is to be served.
Season. Add a few drops of lemon.

Game Sauce
2 onions
A bouquet of thyme, bay leaf and parsley
Several pieces of game
1 slice of ham
1 oz. of butter
4 table-spoons of Madeira
¹⁄₂ pint brown sauce (see p. 118)
 Cut the onions, ham and game into small pieces. Add to them the
bouquet. Fry them gently in the butter. Add the Madeira. Simmer
twenty minutes. Add the sauce and simmer ten minutes. Pass
through a sieve.

German Sauce

¹⁄₂ pint rich brown stock

1 tea-spoon glaze
Pheasant bones
12 mushrooms
1 glass white wine
Break the pheasant bones. Add them to the stock. Simmer half-
an-hour. Add the mushrooms. Simmer till tender. Put through a
sieve. Add glaze, seasoning and glass of wine.

Madeira Sauce

¹⁄₂ onion
¹⁄₂ carrot
1 bay leaf
2 cloves
1 slice ham
1 gill brown stock or gravy
¹⁄₂ pint brown sauce (see p. 118)
1 glass Madeira
Cayenne
Juice of half a lemon
Slice the onion and carrot. Put them, with the bay leaf, clove and
the ham, cut in small pieces, in a sauce-pan. Cover with the brown
stock. Boil up quickly. Simmer half-an-hour. Season. Add Madeira,
brown sauce and lemon juice. Rub through a fine sieve. Colour with
caramel colouring (see p. 13) if not dark enough, and stir in the
butter.
 Orange Sauce
2 Seville oranges
¹⁄₂ lemon
1 glass red wine
1 gill brown gravy
1 lump of sugar
Grate the yellow part of the skin of one orange very finely. Add it to
the brown gravy. Simmer a few minutes. Add the wine, the juice of
two oranges and half a lemon, a little cayenne and the sugar. Serve
with game or wild duck.

Sauce Poivrade
1 oz. butter
2 onions
1 carrot
2 cloves
1 bay leaf
1 tea-spoon flour
1 glass red wine
1 glass water
1 table-spoon vinegar
Melt the butter, add the onions and carrot sliced, the cloves, bay-
leaf and flour. Cook until a good brown, then add the wine, water and
vinegar. Boil half-an-hour. Strain. Season with salt and whole pepper.
Serve with game.

Sour Cream Sauce

2 ozs. butter
2 yolks of eggs
1 table-spoon flour
1 gill sour cream