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Understanding
Biology
Fourth Edition

Kenneth A. Mason
University of Iowa

Tod Duncan

Jonathan B. Losos
William H. Danforth Distinguished University Professor and Director,
Living Earth Collaborative, Washington University
UNDERSTANDING BIOLOGY

Published by McGraw Hill LLC, 1325 Avenue of the Americas, New York, NY 10019. Copyright ©2024 by
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ISBN 978-1-266-10043-7
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Brief Contents
About the Authors iv Part V The Diversity of Life 474
Changes to This Edition vi 22 Systematics and Phylogeny 474
23 Prokaryotes and Viruses 494
Acknowledgments viii
24 Protists 520
A Learning Path to Understanding Biology ix 25 Fungi 547
Contents xv 26 Plants 568
27 Animal Diversity 593
Part I The Molecular Basis of Life 1 28 Vertebrates 625

1 The Science of Biology 1

Part VI Plant Form and Function 655
2 The Nature of Molecules and the Properties of Water 21
3 The Chemical Building Blocks of Life 40 29 Plant Form 655
30 Flowering Plant Reproduction 680
Part II Biology of the Cell 65 31 The Living Plant 705

4 Cell Structure 65
Part VII Animal Form and Function 735
5 Membranes 94
6 Energy and Metabolism 115 32 The Animal Body and How It Moves 735
7 How Cells Harvest Energy 131 33 The Nervous System 760
8 Photosynthesis 156 34 Fueling the Body’s Metabolism 789
9 Cell Communication 179 35 Maintaining Homeostasis 822
10 How Cells Divide 198 36 Reproduction and Development 861

Part III Genetics and Molecular Biology 221 Part VIII Ecology and Behavior 893
11 Sexual Reproduction and Meiosis 221 37 Behavioral Biology 893
12 Patterns of Inheritance 236 38 Ecology of Individuals and Populations 919
13 The Chromosomal Basis of Inheritance 256 39 Community Ecology and Ecosystem Dynamics 944
14 DNA: The Genetic Material 276 40 The Living World 979
15 Genes and How They Work 300
16 Control of Gene Expression 328 Appendix: Answer Key A-1
17 Biotechnology 354
Index I-1
18 Genomics 379

Part IV Evolution 404

19 Genes Within Populations 404
20 The Evidence for Evolution 429
21 The Origin of Species 452

iii
About the Authors
Kenneth Mason has held academic positions, as a teacher and researcher, at three different major
universities. He began on the faculty of the University of Kansas, where he designed and established the
genetics lab and taught and published on the genetics of pigmentation in amphibians. At Purdue University,
he successfully developed and grew large introductory biology courses and collaborated with other faculty
in an innovative biology, chemistry, and physics course supported by the National Science Foundation. At the
University of Iowa, where his wife served as president of the university, he taught introductory biology and
Kenneth Mason human genetics. His honor society memberships include Phi Sigma, Alpha Lambda Delta, and, by vote of Purdue
pharmacy students, Phi Eta Sigma Freshman Honors Society. After eight years at the University of Iowa, Kenneth
and his wife both retired; they maintain close ties to the institution as President Emerita and Lecturer Emeritus.

Tod Duncan was formerly Associate Clinical Associate Professor at the University of Colorado Denver, where he
taught first-semester general biology and coordinated first- and second-semester general biology laboratories. He has
also taught general microbiology, virology, the biology of cancer, medical microbiology, and cell biology. A bachelor’s
degree in cell biology with an emphasis on plant molecular and cellular biology from the University of East Anglia in
Lesley Howard England led to doctoral studies in cell-cycle control and then postdoctoral research on the molecular and biochemical
mechanisms of DNA alkylation damage in vitro and in Drosophila melanogaster. Currently, he is interested in factors
affecting retention and success of incoming first-year students from diverse backgrounds. He lives in Boulder,
Colorado, with his Great Dane, Eddie.

Jonathan Losos is the William H. Danforth Distinguished University Professor in the Department of Biology at
Washington University and Director of the Living Earth Collaborative, a partnership between the university, the
Saint Louis Zoo, and the Missouri Botanical Garden. Losos’s research has focused on studying patterns of adaptive
radiation and evolutionary diversification in lizards. He is a member of the National Academy of Sciences, a fellow
of the American Academy of Arts and Science, and the recipient of several awards, including the Theodosius
Dobzhanksy and David Starr Jordan Prizes, the Edward Osborne Wilson Naturalist Award, and the Daniel Giraud
Jonathan Losos
Elliot Medal, as well as receiving fellowships from the John Guggenheim and David and Lucile Packard Foundations.
Losos has published more than 250 scientific articles and has written two books, Lizards in an Evolutionary Tree:
Ecology and Adaptive Radiation of Anoles (University of California Press, 2009) and Improbable Destinies: Fate,
Chance, and the Future of Evolution (Penguin-Random House, 2017). He is currently in the process of writing his next
book, on scientific research on the ecology and evolution of domestic cats.

iv
 A Conceptual Approach
In the decade leading up to 2019, there was a gradual increase in the importance of a variety of forms of
digital instruction. The use of online education was growing, and we lived through fads like the massive open
online course. While much commentary was devoted to a “revolution” in instruction, in fact, it was a relatively
slow evolutionary process. This was completely upended by the COVID pandemic. Institutions that had never
considered online instruction were forced to move partly or entirely online. This was followed by a roller coaster
ride of on again, off again, online and in-person instruction, and hybrid models using both modes of instruction.
McGraw Hill was ideally positioned to respond to this crisis with the best online tools available from any
publisher. This has allowed us to respond to the changing needs of faculty with a variety of tools, and of course,
high-quality textbooks. One unexpected result of all of this turmoil has been the increasing importance of the
actual textbook. As students were required to do more work on their own, the quality and accessibility of the
textbook proved to be critical. While the pandemic may have revealed the importance of a high-quality textbook,
the need will continue afterward. With the emergence of “active learning” in the majors biology classroom,
the responsibility of learning basic information has moved back to the student, giving instructors the time and
opportunity to increase critical thinking and higher-level learning in their courses.
Understanding Biology fits well into this ever-changing environment in two main ways. First, the book is the
result of a thorough analysis using a variety of sources to determine what is actually being taught in most majors
biology courses across the country. This allowed us to produce a book that goes against the trend of encyclopedic
textbooks yet still has all of the material, in the appropriate depth, required for a majors biology course. This helps to
reduce cost, and makes the text less intimidating for students.
The second way that Understanding Biology is ideal for the new landscape is that it was designed from the
beginning using our knowledge of how people learn. There is a strong emphasis on concepts over disconnected
facts, and an organization that provides the student with a clear path to success in learning difficult material. Each
chapter begins with a Learning Path that introduces the major concepts for the chapter. Then within each section,
larger concepts are broken down into more specific supporting concepts. Each of these secondary concepts
comes with a learning objective that tells students what they should be able to do after completing the section.
Each section has a brief review with a question to help students think about the concepts. This organization, and
more important, the content, allows us to focus on promoting student understanding. The end-of-chapter material
includes a review aligned with the organization of the chapter, and assessment based on a modified version of
Bloom’s Taxonomy of Understand, Apply, Synthesize questions.
The art program supports this approach with clear and accessible figures, and stepped-out figures where
appropriate. This includes Scientific Thinking figures that walk a student through how a problem can be solved
experimentally. We have also moved our “Connecting the Concepts” feature to become a tool for active learning
online, which seems to be a better environment to explore the relationships of chapter material to core concepts.
With this edition we have also added a Concept Overview feature. This opens the chapter with a simple figure
containing the main higher-level concepts of the chapter organized as the top levels of a flow diagram. The diagram
is then filled out at the end of the chapter with supporting conceptual statements. The Concept Overviews are
expansions of the Learning Path in a graphic presentation. They provide students with a conceptual overview of
the chapter, and then larger concepts are broken down into more specific supporting concepts. This is particularly
helpful in the second half of the book, where we have combined some topics that might not appear to be
connected, but in fact are related.
We’re excited about the fourth edition of this innovative text, which provides a clear learning path for
a new generation of students who have to deal with unprecedented challenges. The authors have extensive
experience teaching undergraduate biology, which has provided a guide in producing a text that is up to date and
beautifully illustrated and that incorporates a conceptual approach to learning. We have worked hard to provide
explicit learning outcomes clearly tied to each section of the book, which are used as a basis for developing the
assessments. We are also excited about the continually evolving digital environment that provides unique and
engaging learning tools for modern students. We continue to work to closely integrate the text with its media
support materials to provide instructors with an excellent complement to their teaching.

about the authors v

Changes to This Edition
A new pedagogical feature called “Concept Overview” was
added to all chapters in this fourth edition of Understanding
Biology. The chapter opening page contains a diagram laying out
the main concepts in the chapter. In the online eBook in Connect,
Progressive Concept Overviews are inserted in relevant places
in the chapter and provide a visual presentation of concept
statements that support the main concepts. These Progressive
Concept Overviews are available in the Instructor Resources
for students who are using a printed text. All of the Progressive
Concept Overviews are combined in an end-of-chapter diagram
that presents a conceptual hierarchy of the chapter, highlighting
the important concepts.
Also, McGraw Hill is dedicated to creating products that foster
a culture of belonging and are accessible to all the diverse global
customers we serve. Within this edition, content has been reviewed
to implement inclusive content guidelines around topics including
generalizations and stereotypes, gender, abilities/disabilities, race/
ethnicity, sexual orientation, diversity of names, and age. Similarly,
the illustrations throughout were evaluated and revised to ensure
meaningful text and images are distinguishable and perceivable by
users with limited color vision and moderately low vision.
Part I The Molecular Basis of Life
Chapter 1 A number of edits were made for clarity. The section
on the process of science was edited for clarity and
content, including the social context of science.
Chapter 2 Some minor edits were made for clarity.
Chapter 3 Edits were made to clarify the structure of nucleic
acids, and the process of protein folding. One figure
was converted into two figures to better fit the text
and to increase student understanding.
Part II Biology of the Cell
Chapter 4 The section on prokaryotic cell structure was
rewritten to incorporate new information on
organization, substructure, and compartments.
This provides a different view of the complexity
of prokaryotic cells. A new figure was added
comparing bacterial flagella and the archaeal
archaellum.
Chapter 5 A variety of edits were made for clarity and student
comprehension.
Chapter 6 The material on the Second Law of Thermodynamics
was rewritten for clarity and accessibility. This includes
one new figure to clarify the exergonic and endergonic
reactions. The nature of enzymes and how they
function was also rewritten for clarity.
Chapter 7 Light edits were made for clarity and student
understanding.
Chapter 8 The section on the light-independent reactions
was rewritten for clarity and accuracy. This
included edits to one figure for accuracy and
understanding. This also makes this chapter more
vi
congruent with chapter 7 in terms of names of
compounds that have multiple names.
Chapter 9 The chapter was edited for clarity, including
content on the nature of steroid hormones, and
how their structure affects their function.
Chapter 10 The section on eukaryotic chromosome structure
was further updated to reflect current thinking on the
organization of chromatin in the nucleus. This update
includes a new figure on chromosome organization.
Part III Genetics and Molecular Biology
Chapter 11 Minor edits for clarity, including in several figures,
should improve student understanding. The
material on segregation without crossing over was
deleted as outside the scope of a majors course.
Chapter 12 The historical material in the beginning of the
chapter was streamlined, although not removed. The
material on extensions to Mendel was reorganized
and rewritten for both clarity and updated content.
The material on quantitative traits was updated to
provide a more modern view. A new figure was
added to replace an old one on epistasis with a more
approachable example for students.
Chapter 13 The chapter was reorganized to both provide a more
logical flow of topics, and integrate new material that
provides a more modern view of human genetics.
Chapter 14 The presentation of experiments underlying DNA
structure was edited for clarity. The descriptions
of DNA structure were edited to emphasize
structure–function relationships.
Chapter 15 The chapter was edited for clarity and some content
was updated. The discussion of the genetic code
was rewritten for clarity, and information on the
genome, transcriptome, and proteome was edited
for currency.
Chapter 16 The introduction on transcription factors was edited
for clarity and content. The influence of chromatin
structure on gene regulation was updated. Material
on the ubiquitin/proteasome regulation of protein
turnover was added, including one new figure.
Chapter 17 The section on genetic tools and modern medicine
was completely rewritten. New material on
molecular tests for infectious diseases, including
COVID-19, was added. This also clarifies the use of
different kinds of tests.
Chapter 18 The chapter was edited extensively to update
content on a very fast-moving field. The human
genome section was rewritten to update all
content. Discussion was added of reference
genomes, and current knowledge of the human
genome. The wheat genome project was also
rewritten to include new material on sequencing
complex genomes. The content on what makes us
human was also updated.
Part IV Evolution
Chapter 19 The section on genetic variation in populations
was revised reflecting new information based
on wide­spread genomic investigation. Genomic
variation in humans is now discussed in detail,
quantifying the extent of variation that exists
and how that variation is apportioned within and
between populations.
Part V The Diversity of Life
Chapter 22 Information on roots of the eukaryotic tree was
revised to reflect changes in the classification of
the protists.
Chapter 23 The chapter was edited to reflect new information
on the organization of prokaryotic cells, including
compartmentalization. The material on prokaryotic
genetics was compressed, and the figures were
edited. New material on SARS-CoV-2 was added,
including a figure showing the virus life cycle.
Chapter 24 Many changes were made to the chapter to
improve clarity and readability.
Chapter 25 The chapter was edited for clarity and readability.
Chapter 26 The chapter was edited for clarity and readability.
Chapter 27 The discussion of relationships at the base of the
phylogeny for all animals was revised to reflect
new understand­ing and debate about relationships
among sponges, ctenophores, and other animals.
Additional changes were made to reflect other
changes in understanding of phylogenetic
relationships among animal taxa, such as the
position of chaetognaths and relationships among
protostome taxa. Aspects of taxonomy and natural
history were updated in line with new findings.
Chapter 28 The discussion of human evolution was revised in
light of new discoveries.
Part VI Plant Form and Function
Chapter 29 Minor edits were made throughout for readability,
clarity, and accuracy.
Chapter 30 The chapter was edited for clarity and readability.
Chapter 31 The chapter was edited throughout for clarity,
accuracy, and readability.
Part VII Animal Form and Function
Chapter 32 The chapter was edited for clarity. The material on
bone structure and development was rewritten to
improve student understanding.
Chapter 33 The description of sensory systems, including
receptor types, was updated to make it more
consistent and to improve student understanding.
The material on the retina was edited extensively
for clarity, and one figure was added.
Chapter 34 The section on lung function was edited for clarity.
Information was added on how lung function is
assessed. The cardiac cycle was edited for clarity
and student understanding.
Chapter 35 The material on hormones was edited to provide
a better focus on structure–function relationships.
The material on fluid balance was edited to
emphasize homeostatic mechanisms. The section
on adaptive immunity was rewritten to include
more history of vaccination and a more worldwide
perspective. Material on vaccination and how
the immune system responds was added. This
includes a discussion of vaccination and the
COVID pandemic.
Chapter 36 The material on nuclear reprogramming was
edited to provide a better historical perspective.
The material on cloning in mammals was edited to
include more detail. The material on contraception
was edited for clarity.
Part VIII Ecology and Behavior
Chapter 37 Several Review of Concept questions were
replaced to better assess student understanding.
Chapter 38 Human population trends and other timely data
were updated to stay current. Also, a new section
(38.7) was added on pandemics and human health
that covers the general topic, but extensively
details the population biology of the COVID-19
pandemic.
Chapter 40 Data on biosphere impacts of humans were
updated to stay current.
changes to this edition vii
Acknowledgments
A revision of this scope relies on the talents and efforts
of many people working behind the scenes, and we have
benefited greatly from their assistance.
The copy editor, Beth Bulger, labored many hours and always
improved the clarity and consistency of the text. She has made
a tremendous contribution to the quality of the final product.
We were fortunate to work with MPS Limited to update the
art program and improve the layout of the pages. Our close
collaboration resulted in a text that is pedagogically effective as
well as more beautiful than any other biology text on the market.
We have the continued support of an excellent team at
McGraw Hill. Lora Neyens, the Portfolio Manager for Biology,
has been a steady leader during a time of change. Senior
Product Developer, Liz Sievers, provided support in so many
ways, it would be impossible to name them all. Kelly Hart, Lead
Content Project Manager, and David Hash, Designer, ensured
our text was elegantly designed. Kelly Brown, Senior Marketing
Manager, is always a sounding board for more than just
marketing, and many more people behind the scenes have all
contributed to the success of our text. This includes the digital
team, to whom we owe a great deal for their efforts to continue
improving our Connect assessment tools.
Throughout this edition we have had the support of spouses
who have seen less of us than they might have liked because
of the pressures of getting this revision completed. They
have adapted to the many hours this book draws us away
from them, and, even more than us, looked forward to its
completion.
In the end, the people we owe the most are the generations
of students who have passed through our lecture halls. They
have taught us at least as much as we have taught them, and
viii
their questions and suggestions continue to improve the text
and supplementary materials.
Finally, we need to thank instructors from across the country
who are continually sharing their knowledge and experience
with us through market feedback, reviews, and symposia. Their
feedback shaped this edition. All of these people took time to
share their ideas and opinions to help us build a better edition of
Understanding Biology for the next generation of introductory
biology students, and they have our heartfelt thanks.
Reviewers for Understanding Biology, 4th edition
Christopher Allen Lone Star College, University Park
Lauri Carey Illinois Valley Community College
Christopher Chamberlain St. Cloud Technical
& Community College
Mary Colon Seminole State College
Carolyn Danna Stevenson University
Jacqueline Dartley Bergen Community College
Sandra Fox-Moon Anne Arundel Community College
Robin Graham Dallas College
Linda Johnson University of Maryland Eastern Shore
Olga Ruiz Kopp Utah Valley University
Kimberly Maznicki Seminole State College
Rosa Moscarella University of Massachusetts
Terina Nusinov Seminole State College
Helene Peters Brewton-Parker College
Debra Rinne Seminole State College
Sherry Stewart Navarro College
viii
A Learning Path to Understanding Biology
Understanding Biology and its online assets have been carefully thought out and crafted to help students and professors work efficiently
and effectively through the material in the course, making the most of study time and furthering instructional goals.
7
Le arning Path
7.1
7.2
7.3
7.4
ErikAgar/Getty Images
C onc e pt Ove rvie w
This Concept Overview provides a diagram of the overarching concepts that are covered in this chapter. These concepts will be expanded upon
in the Concept Overview at the end of the chapter.
Organisms convert chemical energy to ATP through cellular respiration
Cellular respiration
extracts energy from
organic molecules
Glycolysis splits glucose
and yields a little ATP
Introduc tion
Life is driven by energy. All the activities carried out by organisms use energy—the swimming of bacteria, the purring of a cat, and even the thoughts
you are forming to process these words. In this chapter, we discuss the processes all cells use to obtain chemical energy from organic molecules,
which is used to synthesize ATP. Then, in chapter 8, we will examine photosynthesis, which uses light energy to make chemical energy. We consider
the conversion of chemical energy to ATP first because all organisms—including the plant, a photosynthesizer, and the caterpillar feeding on the
plant, pictured in the photo—are capable of harvesting energy from chemical bonds. Obtaining energy via respiration is an ancient and universal
process.
▲ the mitochondria.
Every chapter opens with a Learning Path that walks students
through the main concepts in the chapter to enable their
understanding of where the material fits in the context of
other concepts in the chapter. A new Concept Overview
provides a graphical presentation of the Learning Path,
highlighting the main concepts in the chapter.
This complex can be dissociated into two subportions: the
F0 membrane-bound complex, and the F1 complex composed of
the stalk and a knob, or head domain.
The F1 complex has enzymatic activity. The F0 complex con-
tains a channel through which protons move across the membrane
down their concentration gradient. As they do so, their movement
causes part of the F0 complex and the stalk to rotate relative to the
knob. The mechanical energy of this rotation is used to change the
conformation of the catalytic domain in the F1 complex.
Thus, the synthesis of ATP is achieved by a tiny rotary motor,
the rotation of which is driven directly by a gradient of protons.
The flow of protons is like that of water in a hydroelectric power
plant. As the flow of water driven by gravity causes a turbine to
rotate and generate electric current, the proton gradient produces
the energy that drives the rotation of the ATP synthase generator.
REVIEW OF CONCEPT 7.4
The electron transport chain receives electrons from NADH
and FADH2 and passes them down the chain to oxygen, using
the energy from electron transfer to pump protons across the
membrane, creating an electrochemical gradient. The enzyme
ATP synthase uses this gradient to drive the endergonic reac-
tion of phosphorylating ADP to ATP.
■ How would poking a small hole in the outer membrane
affect ATP synthesis?
▲ class, students might focus on the “Understand” level. As they
At the end of each section, Review of Concept questions
allow students to check their understanding before
moving on to the next concept.
How Cells Harvest Energy
Cells Harvest Energy from
Organic Compounds by
Oxidation
Glycolysis Splits Glucose and
Yields a Small Amount of ATP
The Citric Acid Cycle Is the
Oxidative Core of Cellular
Respiration
Electrons Removed by
Oxidation Pass Along an
Electron Transport Chain
The citric acid cycle
oxidizes 2-carbon
units
Oxidative phosphorylation
uses chemiosmosis to
make ATP
The Learning Path
glycolysis needs to be transported into the mitochondria by active
7.5 The Total Energy Yield of transport, which costs one ATP per NADH transported. This
reduces the predicted yield for eukaryotes to 30 ATP.
Aerobic Respiration Far Exceeds
That of Glycolysis
Calculation of P/O Ratios Has
Changed over Time
Ever since the link was discovered between electron transport
and the proton gradient used by ATP synthase, biochemists have LEARNING OBJECTIVE 7.5.2 Explain how our understanding
attempted to determine the number of ATPs produced per NADH of the P/O ratio has changed over time.
feeding electrons into electron transport. This number has proved
7.5 The Total Energy Yield of to be surprisingly elusive. Early estimates were based on errone- The value for the amount of ATP synthesized per O2 molecule
Aerobic Respiration Far
ous assumptions, but we now have both theoretical and calculated reduced is called the phosphate-to-oxygen ratio (P/O ratio).
Exceeds That of Glycolysis
values that are in agreement. Both theoretical calculations and direct measurement of this
7.6 Aerobic Respiration Is value have been contentious issues. When theoretical calcula-
Regulated by Feedback
Inhibition tions were first made, we lacked detailed knowledge of the
The Theoretical Yield for Eukaryotes Is respiratory chain and the mechanism for coupling electron
7.7 Some Forms of Energy
Metabolism Do Not Require O2 30 Molecules of ATP per Glucose transport to ATP synthesis. Because redox reactions occur
7.8 Carbohydrates Are Not the Molecule at three sites for NADH and two sites for FADH 2 , it was
Only Energy Source Used by assumed that three molecules of ATP were produced per
Heterotrophs
LEARNING OBJECTIVE 7.5.1 Calculate the number of ATP NADH and two per FADH2. We now know that assumption
produced by a cell via aerobic respiration. was overly simplistic.
Understanding that a proton gradient is the link between
The number of molecules of ATP produced by ATP synthase per electron transport and ATP synthesis changed the nature of the
molecule of glucose depends on the number of protons trans- calculations. We need to know the number of protons pumped
ported across the inner membrane and the number of protons during electron transport: 10 H+ per NADH, and 6 H+ per
▲ needed per ATP synthesized. The number of protons transported
per NADH and FADH2 is 10 and 6 H+, respectively. Each ATP
synthesized requires 4 H+, leading to 10/4 = 2.5 ATP/NADH,
FADH2. Then we need to know the number of protons needed per
ATP. Because ATP synthase is a rotary motor, this calculation
depends on the number of binding sites for ATP and the number
Every concept is broken down into sections that
and 6/4 = 1.5 ATP/FADH2.
Oxidizing glucose to pyruvate via glycolysis yields 2 ATP
of protons required for rotation. We know that ATP synthase has
three binding sites for ATP. If 12 protons are used per rotation,
directly, and 2 × 2.5 = 5 ATP from NADH. The oxidation of you get the value of 4 H+ per ATP used in the previous calcula-
Energy metabolism
varies across
living systems
cover skills or ideas students should master. Learning
pyruvate to acetyl-CoA yields another 2 × 2.5 = 5 ATP from
NADH. Finally, the citric acid cycle produces 2 ATP directly,
tion. Actual measurements of the P/O ratio have been problem-
atic, but they now appear to be at most 2.5.
objectives at the beginning of each section help identify
6 × 2.5 = 15 ATP from NADH, and 2 × 1.5 = 3 ATP from FADH2.
Summing all of these leads to 32 ATP for respiration (figure 7.16).
We can also calculate how efficiently respiration captures
the free energy released by the oxidation of glucose in the form of
This number is accurate for bacteria, but it does not hold ATP. The amount of free energy released by the oxidation of glu-
important concepts.
for eukaryotes, because the NADH produced in the cytoplasm by cose is 686 kcal/mol, and the free energy stored in each ATP is
Figure 7.16 Theoretical Glucose 2
ATP
ATP yield. The theoretical
yield of ATP harvested from 2 ATP Glycolysis
glucose by aerobic respiration
Pyruvate 2 NADH 5 ATP
totals 32 molecules. In
Assessing the Learning Path
eukaryotes this is reduced to
30 because it takes 1 ATP to
Chemiosmosis
Pyruvate oxidation 2 NADH 5 ATP
transport each molecule of
NADH that is generated by
Understand glycolysis in the cytoplasm into 4. An endergonic reaction has which of the following properties?
1. A covalent bond between a hydrogen atom and an oxygen
a. +ΔG and the reaction is spontaneous. 2
b. +ΔG and the reaction is not spontaneous.
ATP
atom represents what kind of energy? c. −ΔG and the reaction is spontaneous.
Citric Acid
a. Kinetic energy Cycle 6 NADH 15
d. −ΔG and the reaction is not spontaneous.
ATP
b. Potential energy Chemiosmosis
c. Mechanical energy 5. The molecule ATP is less stable than ADP + Pi because
d. Solar energy a. the negatively charged
2 FADH 2
phosphates 3 repel
ATP each other.
b. the positively charged phosphates repel each other.
2. During a redox reaction, the molecule that gains an electron is
a. reduced and now has a higher energy level.
b. oxidized and now has a lower energy level.
c. ATP is much larger than ADP and Pi.
d. the adenine inTotal
ATP netis charged.
ATP yield = 32
(30 in eukaryotes)
6. What is activation energy?
c. reduced and now has a lower energy level.
146
d. oxidized andPart
nowIIhas
Biology of the
a higher Cell level.
energy a. The thermal energy associated with random movements of
3. When a bear eats a salmon, some of the energy stored in the molecules
salmon is used by the bear for its activities and growth. Much
of the energy originally in the salmon is dissipated as heat.
b. The energy released through breaking chemical bonds
c. The difference in free energy between reactants and
products
This is an example of
a. the conversion of kinetic energy to potential energy. d. The energy required to initiate a chemical reaction
b. the Second Law of Thermodynamics. 7. Which of the following is NOT a property of an enzyme?
c. 100% efficient energy conversion. a. An enzyme reduces the activation energy of a reaction.
d. a conversion of potential energy to kinetic energy. b. An enzyme lowers the free energy of the reactants.
Apply
1. When a hibernating animal uses its stored fat to power basic Synthesize
body functions (for example, breathing), it is
1. Some people argue that evolution, which is generally
a. converting kinetic energy to potential energy.
associated with progressive increases in the complexity
b. converting kinetic energy to chemical energy.
c. converting potential energy to kinetic energy.
(order) of organisms, cannot occur because entropy (disorder)
is increasing in the universe. Is this argument valid? Explain.
d. converting chemical energy to potential energy.
2. On summer nights in many parts of the country, one can
2. During certain stages of cellular respiration, electrons are
transferred from glucose molecules to a molecule called
often see fireflies glowing briefly in the dark. Do you suppose
producing this light requires energy? If so, where might the
nicotinamide adenine dinucleotide (NAD+). During this reaction,
a. glucose is oxidized and NAD+ is reduced.
energy come from? How would you test your hypothesis?
3. Examine the graph shown here. Describe what happens
b. glucose is reduced and NAD+ is oxidized.
to this human enzyme protein’s structure when the body’s
c. both glucose and NAD+ have gained protons.
d. glucose has gained protons and NAD+ has lost protons. temperature is raised above 40°C.
3. Sodium ions (Na+) can move through channel proteins
across some biological membranes. If Na+ is present in a Optimum temperature
Rate of reaction
higher concentration on one side of a membrane, the ions for human enzyme
will tend to move across the membrane until they are equally
distributed on both sides of the membrane. This process
a. results in a gain of potential energy for the cell.
b. results in a decrease in entropy.
c. follows the Second Law of Thermodynamics. 30 40 50
d. All of the above Temperature of reaction
4. If the products of a chemical reaction have higher free energy
4. Phosphofructokinase functions to add a phosphate group
than the reactants, this reaction
a. will not proceed spontaneously. to fructose-6-phosphate. This enzyme functions early in
b. will proceed spontaneously. glycolysis (refer to chapter 7). The enzyme’s active site binds
c. must have increased the total energy in the universe. to fructose and ATP. An allosteric inhibitory site can also bind
d. must have decreased the total energy in the universe. ATP when ATP levels are high.
5. ATP can be used to drive an endergonic reaction if a. Predict the rate of the reaction if with low ATP levels.
b. Predict the rate of the reaction if with high ATP levels.
a. ATP hydrolysis lowers the activation energy for the reaction.
b. the sum of ΔG for ATP hydrolysis and ΔG for the reaction c. Describe what is happening to the enzyme with high levels
is negative. of ATP.
▲
Chapter concepts are assessed at three different levels at the
end of the chapter. On a first pass through the chapter prior to
gain greater mastery of the material, they should challenge
themselves with “Apply” and “Synthesize” questions that
require higher cognitive skills.
ix
 Think Like a Scientist

Scientific Thinking figures throughout Inquiry & Analysis

▲
the text walk the student through How Hemorrhagic E. coli
features at the ends of

Inquiry & Analysis

pH Sensitivity of the Glu-GABA Antiporter

a scientific experiment, laying out Resists the Acid Environment

of the Stomach
300
GadC
AdiC
all chapters explore a
250

the Hypothesis, Predictions, Test scientific investigation in

Substrate accumulation
(nmol per mg protein)
Recent years have been marked by a series of food 200
poisoning outbreaks involving hemorrhagic (producing

Procedures, Results, and Conclusion. more detail, presenting

internal bleeding) strains of the bacterium Escherichia coli 150
(E. coli). Bacteria are often a source of food poisoning,
typically milder infections caused by food-borne strepto- 100

Some also challenge the students to experimental results and

coccal bacteria. Less able to bear the extremely acidic
conditions encountered by food in the human stomach 50
(pH = 2), E. coli has not been as common a problem. The

devise further experiments. challenging students to

hemorrhagic strains of E. coli responsible for recent out- 0
5 6 7 8 9
breaks seem to have evolved more elaborate acid-
pH
▲ resistance systems.

interpret the data.

How do hemorrhagic E. coli bacteria survive in the
acid environment of the stomach? The problem they face,
in essence, is that they are submerged in a sea of hydro-
gen ions, many of which diffuse into their cells. To rid
themselves of these excess hydrogen ions, the E. coli Analysis
SCIENTIFIC THINKING use a clever system to pump hydrogen ions back out of
their cells. 1. Applying Concepts
Hypothesis: All wavelengths of light are equally effective in First, the hemorrhagic E. coli cells take up cellular a. Variable. In the graph, what is the dependent
hydrogen ions by using the enzyme glutamic acid decarbox- variable?
promoting photosynthesis.
ylase (GAD) to convert the amino acid glutamate to b. Substrate. What is a substrate? In this
Prediction: Illuminating plant cells with light broken into different investigation, what are the substrates that are
γ-aminobutyric acid (GABA), a decarboxylation reaction
wavelengths by a prism will produce the same amount of O2 for that consumes a hydrogen ion. accumulating?
Second, the hemorrhagic E. coli export this GABA c. pH. What is the difference in hydrogen
all wavelengths.
from their cell cytoplasm using a Glu-GABA antiporter ion concentration between pH 5 and pH 7?
Test: A filament of algae immobilized on a slide is illuminated by light How many times more (or less) is that? Explain.
called GadC (this transmembrane protein channel is called
that has passed through a prism. Motile bacteria that require O2 for an antiporter because it transports two molecules across the 2. Interpreting Data
membrane in opposite directions). a. Does the amount of amino acid transported
growth are added to the slide. in the 10-minute experimental interval
However, to survive elsewhere in the human body, it
is important that the Glu-GABA antiporter of hemorrhagic (expressed as substrate accumulation) vary
high with pH for the arginine-transporting AdiC
Oxygen-seeking bacteria E. coli not function, lest it short-circuit metabolism. To evalu-
ate if the GadC antiporter indeed functions only in acid antiporter? For the glutamate-transporting
GadC antiporter?
Absorption

environments, investigators compared its activity at a vari-

b. Compare the amount of substrate accumulated
Light

Filament of green algae ety of pHs with that of a different amino acid antiporter
called AdiC, which transports arginine out of cells under a by AdiC in 10 minutes at pH 9.0 with that
broad range of conditions. The results of monitoring trans- accumulated at pH 5.0. What fraction of the
port for 10 minutes are presented in the graph. low pH activity is observed at the higher pH?
low c. In a similar fashion, compare the amount of
substrate accumulated by GadC at pH 9.0
with that accumulated at pH 5.0. What
Result: The bacteria move to regions of high O2 , or regions of most fraction of the low pH activity is observed at
Outside Inside the higher pH?
active photosynthesis. These are in the purple/blue and red regions cell cell
3. Making Inferences Would you say that the GadC
of the spectrum. antiporter exhibits the same pH dependence as the
Conclusion: All wavelengths are not equally effective at promoting AdiC antiporter? If not, which antiporter is less
GABA active at nonacid pHs?
photosynthesis. The most effective constitute the action spectrum
4. Drawing Conclusions Is the glutamate-GABA
for photosynthesis. antiporter GadC active at nonacid pHs?
Further Experiments: How does the action spectrum relate to the 5. Further Analysis The GadC antiporter also
various absorption spectra in figure 8.5? Glutamate transports the amino acid glutamine (Gln). Do you
think this activity has any role to play in combating
low pH environments? How would you test this
hypothesis?
Figure 8.7 Determination of an action spectrum for
photosynthesis. Chapter 5 Membranes 111

Concept Overviews
Each Concept Overview, a new feature in Understanding Biology, 4th edition, is a graphic representation of the Learning Path that
guides students through the concepts in the chapter. Concept Overviews provide a conceptual framework of the chapter. When
concept statements are placed in the context of a flow diagram, students see relationships and connections between concepts.

Flowering plant reproductive strategies ensure their success and diversity Flowering plant reproductive strategies ensure their success and diversity

Flowering is the first step Self-pollination can be favored Flowering is the first step Double fertilization is followed Plant growth starts with Self-pollination can be favored
Double fertilization is followed Plant growth starts with
in plant reproduction by embryogenesis germination in stable environments
in plant reproduction by embryogenesis germination in stable environments

Flowering leads to gamete Fertilization Seeds play roles Germination Life spans vary Asexual reproduction

▲ production, fertilization, produces the in dispersal, requires water, reduces variation

embryogenesis, and adult endosperm and protection, and metabolism,
plant development one embryo food storage and Woody plants
environmental are generally In apomixis, seeds

The Concept Overview that appears at the beginning of the A root–shoot A food supply
cues perennial,
growing
form from cloned
cells in the ovule
Flowering is A complete axis and a develops, the every year

chapter contains three to five broad concept statements regulated by

internal
flower has
sepals,
petals,
radial axis are
formed
seed coat forms,
and a fruit forms
Roots anchor
the seedling Herbaceous
In vegetative
reproduction, new
developmental plants can be plants can arise
that reveal the overarching structure of the chapter. and
environmental
cues
stamens,
and a
carpel
Primary
meristems
Stored nutrients
are essential until
Stored starch,
fat, and
annual, biennial,
or perennial
from stolons,
rhizomes, suckers,
or plantlets
differentiate photosynthesis proteins
into protoderm, can occur fuel growth Biennial
ground plants grow Single plant
Female Pollination meristem, and protoplasts can be
gametophytes can occur Outer ovule one year and

Progressive Concept
procambium Emerging cloned in the lab
▲

are embryo by wind, develops into a reproduce the

tissues shoots next
sacs self-pollination, seed coat become
or pollinators Plant tissue culture
Flowering plant reproductive strategies ensure their success and diversity
Overviews are inserted Male
gametophytes Pollinators are
A fruit is the
photosynthetic
Annual plants
die after one
can be used for
genetic engineering
mature ovary of
Double fertilization is followed
by embryogenesis
throughout the chapter are pollen
grains
attracted by
odor or flower
characteristics
an angiosperm growing
season

in the eBook, at the end and disperse

pollen

Fertilization
produces the
endosperm and
Seeds play roles
in dispersal,
protection, and
of relevant sections. They include
bees,
butterflies,
one embryo food storage
These diagrams identify birds, and
bats

A root–shoot
axis and a
A food supply
develops, the concepts that support
radial axis are
formed
seed coat forms,
and a fruit forms
the overarching concept ▲
Primary
meristems
Stored nutrients
are essential until statements. The Each chapter’s end-of-chapter Concept Overview brings
photosynthesis
differentiate
into protoderm,
ground
can occur
Progressive Concept together all of the concept statements in the Progressive
meristem, and
procambium
Outer ovule
develops into a Overviews are available Overview diagrams to reveal a conceptual overview of the
tissues
seed coat

in the Instructor chapter. These Concept Overviews differ from “concept maps”
A fruit is the
mature ovary of
an angiosperm Resources for students in that there is a hierarchical aspect important in developing
using a printed text. a conceptual framework for the chapter.

x a learning path to understanding biology

 Strengthen Problem-Solving Skills with Connect®
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a learning path to understanding biology xi
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xii a learning path to understanding biology


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Contents
About the Authors iv PART II Biology
Changes to This Edition vi of the Cell 65
Acknowledgments viii Dr. Gopal Murti/Science Source

A Learning Path to Understanding Biology ix 4 Cell Structure 65

Concept 4.1 All Living Organisms Are Composed
of Cells 66
Concept 4.2 Prokaryotic Cells Are Relatively Simple 69
PART I The Molecular Concept 4.3 Eukaryotic Cells Are Highly Compartmentalized 71
Basis of Life 1 Concept 4.4 Membranes Organize the Cell Interior into Functional
Compartments 75
Soames Summerhays/Natural Visions
Concept 4.5 Mitochondria and Chloroplasts Are Energy-Processing
1 The Science of Biology 1 Organelles 79
Concept 1.1 The Diversity of Life Is Overwhelming 2 Concept 4.6 An Internal Skeleton Supports the Shape
of Cells 81
Concept 1.2 Biology Is the Science of Life 2
Concept 4.7 Extracellular Structures Protect Cells 84
Concept 1.3 Science Is Based on Both Observation and Reasoning 6
Concept 4.8 Cell-to-Cell Connections Determine How Adjacent
Concept 1.4 The Study of Evolution Is a Good Example Cells Interact 86
of Scientific Inquiry 9
Inquiry & Analysis 90
Concept 1.5 A Few Important Concepts Form the Core
of Biology 14 Retracing the Learning Path 91
Inquiry & Analysis 17 Assessing the Learning Path 92
Retracing the Learning Path 18
Assessing the Learning Path 19 5 Membranes 94
Concept 5.1 Membranes Are Phospholipid Bilayers with
2 The Nature of Molecules and Embedded Proteins 95
Concept 5.2 Phospholipids Provide a Membrane’s Structural
the Properties of Water 21 Foundation 98
Concept 2.1 All Matter Is Composed of Atoms 22 Concept 5.3 Membrane Proteins Enable a Broad Range of
Concept 2.2 The Elements in Living Systems Have Low Atomic Interactions with the Environment 99
Masses 25 Concept 5.4 Passive Transport Moves Molecules Across
Concept 2.3 Molecules Are Collections of Atoms Held Together Membranes by Diffusion 101
by Chemical Bonds 27 Concept 5.5 Active Transport Across Membranes Requires
Concept 2.4 The Properties of Water Result from Its Energy 105
Polar Nature 31 Concept 5.6 Bulky Materials Cross Membranes Within
Concept 2.5 Water Molecules Can Dissociate into Ions 34 Vesicles 108
Inquiry & Analysis 36 Inquiry & Analysis 111
Retracing the Learning Path 37 Retracing the Learning Path 112
Assessing the Learning Path 38 Assessing the Learning Path 113

3 The Chemical Building Blocks of Life 40 6 Energy and Metabolism 115

Concept 3.1 Carbon Provides the Framework of Biological Concept 6.1 Energy Flows Through Living Systems 116
Molecules 41 Concept 6.2 The Laws of Thermodynamics Govern All Energy
Concept 3.2 Carbohydrates Form Both Structural and Energy- Changes 117
Storing Molecules 43 Concept 6.3 ATP Is the Energy Currency of Cells 119
Concept 3.3 Proteins Are the Tools of the Cell 47 Concept 6.4 Enzymes Speed Up Reactions by Lowering Activation
Concept 3.4 Nucleic Acids Store and Express Genetic Energy 121
Information 55 Concept 6.5 Metabolism Is the Sum of a Cell’s Chemical
Concept 3.5 Hydrophobic Lipids Form Fats and Membranes 58 Activities 125
Inquiry & Analysis 61 Inquiry & Analysis 127
Retracing the Learning Path 62 Retracing the Learning Path 128
Assessing the Learning Path 63 Assessing the Learning Path 129

xv
 7 How Cells Harvest Energy 131 Concept 10.3 The Eukaryotic Cell Cycle Is Complex and Highly
Organized 203
Concept 7.1 Cells Harvest Energy from Organic Compounds Concept 10.4 During Interphase, Cells Grow and Prepare for
by Oxidation 132 Mitosis 204
Concept 7.2 Glycolysis Splits Glucose and Yields a Small Amount Concept 10.5 In Mitosis, Chromosomes Segregate 205
of ATP 136
Concept 10.6 Events of the Cell Cycle Are Carefully Regulated 210
Concept 7.3 The Citric Acid Cycle Is the Oxidative Core of
Cellular Respiration 139 Concept 10.7 Cancer Is a Failure of Cell-Cycle Control 214
Concept 7.4 Electrons Removed by Oxidation Pass Along an Inquiry & Analysis 217
Electron Transport Chain 142 Retracing the Learning Path 218
Concept 7.5 The Total Energy Yield of Aerobic Respiration Far Assessing the Learning Path 219
Exceeds That of Glycolysis 146
Concept 7.6 Aerobic Respiration Is Regulated by Feedback

Concept 7.7
Inhibition 147
Some Forms of Energy Metabolism Do Not
PART III Genetics and
Require O2 148 Molecular Biology 221
Concept 7.8 Carbohydrates Are Not the Only Energy Source Used Steven P. Lynch
by Heterotrophs 149
Inquiry & Analysis 152 11 Sexual Reproduction and Meiosis 221
Retracing the Learning Path 153 Concept 11.1 Sexual Reproduction Requires Meiosis 222
Assessing the Learning Path 154 Concept 11.2 Meiosis Consists of Two Divisions with One Round
of DNA Replication 223
8 Photosynthesis 156 Concept 11.3 The Process of Meiosis Involves Intimate Interactions
Between Homologs 224
Concept 8.1 Photosynthesis Uses Sunlight to Power the Synthesis
of Organic Molecules 157 Concept 11.4 Meiosis Has Four Distinct Features 229
Concept 8.2 Experiments Revealed That Photosynthesis Is a Concept 11.5 Genetic Variation Is the Evolutionary Consequence
Chemical Process 158 of Sex 231
Concept 8.3 Pigments Capture Energy from Sunlight 160 Inquiry & Analysis 232
Concept 8.4 Photosynthetic Pigments Are Organized into Retracing the Learning Path 233
Photosystems 163 Assessing the Learning Path 234
Concept 8.5 Energy from Sunlight Is Used to Produce a Proton
Gradient 165 12 Patterns of Inheritance 236
Concept 8.6 Carbon Fixation Incorporates CO2 into Organic
Concept 12.1 Experiments Carried Out by Mendel Explain
Molecules 170
Inheritance 237
Concept 8.7 Photorespiration Short-Circuits Photosynthesis 172
Concept 12.2 Mendel’s Principle of Segregation Accounts
Inquiry & Analysis 175 for 3:1 Phenotypic Ratios 238
Retracing the Learning Path 176 Concept 12.3 Mendel’s Principle of Independent Assortment
Assessing the Learning Path 177 Asserts That Genes Segregate Independently 241
Concept 12.4 Probability Allows Us to Predict the Results of
9 Cell Communication 179 Crosses 243
Concept 9.1 The Cells of Multicellular Organisms Concept 12.5 Extending Mendel’s Model Provides a Clearer View
Communicate 180 of Genetics in Action 245
Concept 9.2 Signal Transduction Begins with Cellular Receptors 182 Concept 12.6 Genotype Dictates Phenotype by Specifying Protein
Sequences 250
Concept 9.3 Intracellular Receptors Respond to Signals by
Regulating Gene Expression 184 Inquiry & Analysis 252
Concept 9.4 Protein Kinase Receptors Respond to Signals by Retracing the Learning Path 253
Phosphorylating Proteins 185 Assessing the Learning Path 254
Concept 9.5 G Protein–Coupled Receptors Respond to Signals
Through Effector Proteins 189 13 The Chromosomal Basis
Inquiry & Analysis 194
of Inheritance 256
Retracing the Learning Path 195
Concept 13.1 Sex Linkage and the Chromosomal Theory
Assessing the Learning Path 196
of Inheritance 257
10 How Cells Divide 198 Concept 13.2 There Are Two Major Exceptions to Chromosomal
Inheritance 259
Concept 10.1 Bacterial Cell Division Is Clonal 199 Concept 13.3 Some Genes Do Not Assort Independently:
Concept 10.2 Eukaryotes Have Large, Linear Chromosomes 200 Linkage 261

xvi contents
Concept 13.4 Genetic Crosses Provide Data for Genetic Maps 263 Inquiry & Analysis 350
Concept 13.5 Changes in Chromosome Number Can Have Drastic Retracing the Learning Path 351
Effects 265 Assessing the Learning Path 352
Concept 13.6 Inheritance in Humans Can Be Studied by Analyzing
Pedigrees 267
17 Biotechnology 354
Inquiry & Analysis 272
Concept 17.1 Enzymes Allow the Creation of Recombinant
Retracing the Learning Path 273
Molecules In Vitro 355
Assessing the Learning Path 274
Concept 17.2 The Polymerase Chain Reaction Is Used to Amplify
Specific DNA Sequences 359
14 DNA: The Genetic Material 276 Concept 17.3 Molecular Tools Allow Us to Analyze and Modify
Concept 14.1 DNA Is the Genetic Material 277 Genetic Variation 362
Concept 14.2 The DNA Molecule Is a Double Helix 279 Concept 17.4 Transgenic Organisms Are Used to Analyze Gene
Function 364
Concept 14.3 Both Strands Are Copied During DNA
Replication 283 Concept 17.5 Genetic Tools Are Changing Modern Medicine 366
Concept 14.4 Prokaryotes Organize the Enzymes Used Concept 17.6 Genetic Engineering Is Used in Industry and
to Duplicate DNA 286 Agriculture 370
Concept 14.5 Eukaryotic Chromosomes Are Large and Linear 291 Inquiry & Analysis 375
Concept 14.6 Cells Repair Damaged DNA 293 Retracing the Learning Path 376
Inquiry & Analysis 296 Assessing the Learning Path 377
Retracing the Learning Path 297
Assessing the Learning Path 298 18 Genomics 379
Concept 18.1 Mapping Identifies and Locates Functional Elements
in Genomes 380
15 Genes and How They Work 300
Concept 18.2 The Modernization of DNA Sequencing Has
Concept 15.1 Experiments Have Revealed the Nature of Genes 301 Accelerated Discovery 383
Concept 15.2 The Genetic Code Relates Information in DNA Concept 18.3 Genome Projects Reveal Insights into Medicine and
and Protein 303 Agriculture 386
Concept 15.3 Prokaryotes Exhibit All the Basic Features Concept 18.4 Genome Annotation Assigns Functional Information
of Transcription 306 to Genomes 388
Concept 15.4 Eukaryotes Use Three Polymerases and Extensively Concept 18.5 Genome Comparisons Provide Information About
Modify Transcripts 309 Genomic Structure and Function 391
Concept 15.5 Eukaryotic Genes May Contain Noncoding Concept 18.6 Comparative Genomics Informs Evolutionary
Sequences 311 Biology 396
Concept 15.6 The Ribosome Is the Machine of Protein Inquiry & Analysis 400
Synthesis 313
Retracing the Learning Path 401
Concept 15.7 The Process of Translation Is Complex and
Assessing the Learning Path 402
Energy-Expensive 315
Concept 15.8 Mutations Are Heritable Changes in Genetic
Material 321
Inquiry & Analysis 324
Retracing the Learning Path 325
PART IV Evolution 404
Assessing the Learning Path 326 Tetra Images/Getty Images

19 Genes Within Populations 404

16 Control of Gene Expression 328 Concept 19.1 Natural Populations Exhibit Genetic Variation 405
Concept 16.1 All Organisms Control Expression of Their Genes 329 Concept 19.2 Frequencies of Alleles Can Change 407
Concept 16.2 Regulatory Proteins Control Genes by Interacting with Concept 19.3 Five Agents Are Responsible for Evolutionary
Specific DNA Nucleotide Sequences 330 Change 409
Concept 16.3 Prokaryotes Regulate Their Genes in Clusters 331 Concept 19.4 Selection Can Act on Traits Affected by Many
Concept 16.4 Transcription Factors Control Gene Transcription Genes 414
in Eukaryotes 336 Concept 19.5 Natural Selection Can Be Studied Experimentally 415
Concept 16.5 Chromatin Structure Affects Gene Expression 339 Concept 19.6 Fitness Is a Measure of Evolutionary Success 418
Concept 16.6 Eukaryotic Genes Are Also Regulated After Concept 19.7 Evolutionary Processes Sometimes Maintain
Transcription 341 Variation 418
Concept 16.7 Gene Regulation Determines How Cells Will Concept 19.8 Sexual Selection Determines Reproductive
Develop 346 Success 421

contents xvii
Inquiry & Analysis 425 Concept 22.4 Taxonomy Attempts to Classify Organisms in an
Retracing the Learning Path 426 Evolutionary Context 482
Assessing the Learning Path 427 Concept 22.5 The Largest Taxa Are Domains 485
Inquiry & Analysis 490
20 The Evidence for Evolution 429 Retracing the Learning Path 491
Assessing the Learning Path 492
Concept 20.1 The Beaks of Darwin’s Finches Provide Evidence of
Natural Selection 430
Concept 20.2 Peppered Moths and Industrial Melanism Illustrate 23 Prokaryotes and Viruses 494
Natural Selection in Action 433
Concept 23.1 Prokaryotes Are the Most Ancient Organisms 495
Concept 20.3 Human-Initiated Artificial Selection Is Also a
Powerful Agent of Change 434 Concept 23.2 Prokaryotes Have an Organized but Simple
Structure 497
Concept 20.4 Fossils Provide Direct Evidence of Evolution 436
Concept 23.3 The Genetics of Prokaryotes Focuses on DNA
Concept 20.5 Anatomical Evidence for Evolution Is Extensive Transfer 501
and Persuasive 440
Concept 23.4 Prokaryotic Metabolism Is Diverse 504
Concept 20.6 Genes Carry a Molecular Record of the Evolutionary
Past 443 Concept 23.5 Bacteria Cause Important Human Diseases 505
Concept 20.7 Natural Selection Favors Convergent Evolution in Concept 23.6 Viruses Are Not Organisms 506
Similar Environments 444 Concept 23.7 Bacterial Viruses Infect by DNA Injection 510
Concept 20.8 Addressing Common Criticisms of Evolutionary Concept 23.8 Animal Viruses Infect by Endocytosis 511
Theory 446 Inquiry & Analysis 516
Inquiry & Analysis 448 Retracing the Learning Path 517
Retracing the Learning Path 449 Assessing the Learning Path 518
Assessing the Learning Path 450
24 Protists 520
21 The Origin of Species 452
Concept 24.1 Protists, the First Eukaryotes, Arose by
Concept 21.1 The Biological Species Concept Highlights Endosymbiosis 521
Reproductive Isolation 453 Concept 24.2 Protists Are a Very Diverse Group 523
Concept 21.2 Natural Selection May Reinforce Reproductive Concept 24.3 The Rough Outlines of Protist Phylogeny Are
Isolation 457 Becoming Clearer 525
Concept 21.3 Natural Selection and Genetic Drift Play Key Roles Concept 24.4 Excavata Are Flagellated Protists Lacking
in Speciation 459 Mitochondria 526
Concept 21.4 Speciation Is Influenced by Geography 460 Concept 24.5 SAR: Stramenopiles and Alveolates Exhibit Secondary
Concept 21.5 Adaptive Radiation Requires Both Speciation and Endosymbiosis 529
Habitat Diversity 462 Concept 24.6 SAR: Rhizaria Have Silicon Exoskeletons or
Concept 21.6 The Pace of Evolution Varies 467 Limestone Shells 535
Concept 21.7 Speciation and Extinction Have Molded Biodiversity Concept 24.7 Archaeplastida Are Descended from a Single
Through Time 468 Endosymbiosis Event 536
Inquiry & Analysis 470 Concept 24.8 Amoebozoa and Opisthokonta Are Closely
Retracing the Learning Path 471 Related 539
Assessing the Learning Path 472 Inquiry & Analysis 542
Retracing the Learning Path 543
Assessing the Learning Path 544

PART V The Diversity 25 Fungi 547

Concept 25.1 Fungi Have Unique Reproductive and Nutritional
of Life 474 Strategies 548
Imagemore Co, Ltd./Imagemore/Getty Images Concept 25.2 Fungi Have an Enormous Ecological Impact 550
22 Systematics and Phylogeny 474 Concept 25.3 Fungi Are Important Plant and Animal Pathogens 553
Concept 25.4 Fungi Are Taxonomically Diverse 554
Concept 22.1 Systematics Reconstructs Evolutionary
Relationships 475 Concept 25.5 Microsporidia Are Unicellular Parasites 556
Concept 22.2 Cladistics Focuses on Traits Derived from a Common Concept 25.6 Chytridiomycota and Relatives: Fungi with
Ancestor 476 Zoospores 556
Concept 22.3 Classification Is a Labeling Process, Not an Concept 25.7 Zygomycota Produce Zygotes 558
Evolutionary Reconstruction 480 Concept 25.8 Glomeromycota Are Asexual Plant Symbionts 559

xviii contents
Concept 25.9 Basidiomycota Are the Mushroom Fungi 560 Concept 28.8 Primates Include Lemurs, Monkeys, Apes, and
Concept 25.10 Ascomycota Are the Most Diverse Phylum Humans 643
of Fungi 560 Inquiry & Analysis 650
Inquiry & Analysis 564 Retracing the Learning Path 651
Retracing the Learning Path 565 Assessing the Learning Path 652
Assessing the Learning Path 566

26 Plants 568
Concept 26.1 Land Plants Are Multicellular Autotrophs Adapted PART VI Plant Form
to Terrestrial Life 569 and Function 655
Concept 26.2 Bryophytes Have a Dominant Gametophyte
Generation 571 Susan Singer

Concept 26.3 Seedless Vascular Plants Evolved Roots, Stems, 29 Plant Form 655
and Leaves 573
Concept 29.1 Meristems Articulate the Body Plan 656
Concept 26.4 Lycophytes Have a Dominant Sporophyte
Concept 29.2 Plants Contain Three Main Tissues 659
Generation 575
Concept 29.3 Roots Have Four Growth Zones 664
Concept 26.5 Pterophytes Are Ferns and Their Relatives 576
Concept 29.4 Stems Provide Support for Aboveground Organs 669
Concept 26.6 Seed Plants Were a Key Step in Plant Evolution 579
Concept 29.5 Leaves Are a Plant’s Photosynthetic Organs 673
Concept 26.7 Gymnosperms Are Plants with “Naked Seeds” 580
Inquiry & Analysis 676
Concept 26.8 Angiosperms Are Flowering Plants 584
Retracing the Learning Path 677
Inquiry & Analysis 588
Assessing the Learning Path 678
Retracing the Learning Path 589
Assessing the Learning Path 590
30 Flowering Plant Reproduction 680
27 Animal Diversity 593 Concept 30.1 Reproduction Starts with Flowering 681
Concept 30.2 Flowers Attract Pollinators 683
Concept 27.1 The Diversity of Animal Body Plans Arose by a Series
of Evolutionary Innovations 594 Concept 30.3 Fertilization Leads to Embryogenesis 688
Concept 27.2 Molecular Data Are Clarifying the Animal Concept 30.4 Seeds Protect Angiosperm Embryos 690
Phylogenetic Tree 601 Concept 30.5 Fruits Promote Seed Dispersal 692
Concept 27.3 True Tissue Evolved in Simple Animals 603 Concept 30.6 Germination Begins Seedling Growth 693
Concept 27.4 Flatworms and Rotifers Are Very Simple Concept 30.7 Plant Life Spans Vary Widely 696
Bilaterians 605 Concept 30.8 Asexual Reproduction Is Common Among Flowering
Concept 27.5 Mollusks and Annelids Are the Largest Groups Plants 697
of Lophotrochozoans 607 Inquiry & Analysis 700
Concept 27.6 Lophophorates Are Very Simple Marine Retracing the Learning Path 701
Organisms 611
Assessing the Learning Path 702
Concept 27.7 Nematodes and Arthropods Are Both Large Groups
of Ecdysozoans 612
Concept 27.8 Deuterostomes Are Composed of Echinoderms and 31 The Living Plant 705
Chordates 617 Concept 31.1 Water Moves Through Plants Based on Potential
Inquiry & Analysis 620 Differences 706
Retracing the Learning Path 621 Concept 31.2 Roots Absorb Minerals and Water 709
Assessing the Learning Path 623 Concept 31.3 Xylem Transports Water from Root to Shoot 711
Concept 31.4 Transpiration Rate Reflects Environmental
28 Vertebrates 625 Conditions 713
Concept 31.5 Plants Are Adapted to Water Stress 715
Concept 28.1 Nonvertebrate Chordates Do Not Form Bone 626
Concept 31.6 Phloem Transports Organic Molecules 716
Concept 28.2 Almost All Chordates Are Vertebrates 627
Concept 31.7 Plants Require a Variety of Nutrients 717
Concept 28.3 Fishes Are the Earliest and Most Diverse
Vertebrates 627 Concept 31.8 Plants Use Hormones to Regulate Growth 719
Concept 28.4 Amphibians Are Moist-Skinned Descendants of the Concept 31.9 Plant Growth Is Responsive to Light 724
Early Tetrapods 632 Concept 31.10 Plant Growth Is Sensitive to Gravity 727
Concept 28.5 Reptiles Are Fully Adapted to Terrestrial Living 633 Inquiry & Analysis 730
Concept 28.6 Birds Are Essentially Flying Reptiles 637 Retracing the Learning Path 731
Concept 28.7 Mammals Are the Least Diverse of Vertebrates 640 Assessing the Learning Path 733

contents xix
 PART VII Animal Form Concept 34.6 Lungs Are the Respiratory Organs of Terrestrial
Vertebrates 800
and Function 735 Concept 34.7 Oxygen and Carbon Dioxide Are Transported by
Fundamentally Different Mechanisms 803
Stockbyte/Getty Images
Concept 34.8 Circulating Blood Carries Metabolites and Gases
32 The Animal Body and How It Moves 735 to the Tissues 806
Concept 32.1 The Vertebrate Body Has a Hierarchical Concept 34.9 Vertebrate Circulatory Systems Put a Premium on
Organization 736 Efficient Circulation 808
Concept 32.2 Epithelial Tissue Covers Body Surfaces 738 Concept 34.10 The Four Chambers of the Heart Contract in
a Cycle 811
Concept 32.3 Nerve Tissue Conducts Signals Rapidly 740
Concept 34.11 The Circulatory Highway Is Composed of Arteries,
Concept 32.4 Connective Tissue Supports the Body 740
Capillaries, and Veins 814
Concept 32.5 Muscle Tissue Powers the Body’s Movements 742
Inquiry & Analysis 817
Concept 32.6 Skeletal Systems Anchor the Body’s Muscles 744
Retracing the Learning Path 818
Concept 32.7 Vertebrate Endoskeletons Are Made of Bone 745
Assessing the Learning Path 820
Concept 32.8 Muscles Contract Because Their Myofilaments
Slide 749
Concept 32.9 Animal Locomotion Takes Many Forms 753 35 Maintaining Homeostasis 822
Inquiry & Analysis 756 Concept 35.1 Homeostasis Maintains a Constant Internal
Retracing the Learning Path 757 Environment 823
Assessing the Learning Path 758 Concept 35.2 Hormones Are Chemical Messages That Direct Body
Processes 827
Concept 35.3 The Pituitary and the Hypothalamus Are the Body’s
33 The Nervous System 760 Control Centers 831
Concept 33.1 The Nervous System Directs the Body’s Actions 761 Concept 35.4 Peripheral Endocrine Glands Play Major Roles in
Concept 33.2 Neurons Maintain a Resting Potential Across the Homeostasis 834
Plasma Membrane 762 Concept 35.5 Animals Are Osmoconformers or Osmoregulators 838
Concept 33.3 Action Potentials Propagate Nerve Impulses 764 Concept 35.6 The Kidney Maintains Osmotic Homeostasis in
Concept 33.4 Synapses Are Where Neurons Communicate with Mammals 839
Other Cells 766 Concept 35.7 Hormones Control Osmoregulation 844
Concept 33.5 The Central Nervous System Includes the Brain Concept 35.8 The Immune System Defends the Body 845
and Spinal Cord 769 Concept 35.9 Cell-Mediated Immunity Involves Helper and Killer
Concept 33.6 The Peripheral Nervous System Consists of Both T Cells 850
Sensory and Motor Neurons 771 Concept 35.10 In Humoral Immunity, B Cells Produce Protective
Concept 33.7 Sensory Receptors Provide Information About the Antibodies 852
Body’s Environment 773 Inquiry & Analysis 856
Concept 33.8 Mechanoreceptors Sense Touch and Pressure 774 Retracing the Learning Path 857
Concept 33.9 Sounds and Body Position Are Sensed by Vibration Assessing the Learning Path 859
Detectors 775
Concept 33.10 Taste, Smell, and pH Senses Utilize
Chemoreceptors 778 36 Reproduction and Development 861
Concept 33.11 Vision Employs Photoreceptors to Perceive Objects Concept 36.1 Mammals Are Viviparous 862
at a Distance 779
Concept 36.2 The Human Male Reproductive System Is Typical
Inquiry & Analysis 785 of Mammals 863
Retracing the Learning Path 786 Concept 36.3 The Human Female Reproductive System Undergoes
Assessing the Learning Path 787 Cyclic Gamete Development 866
Concept 36.4 The First Step in Development Is Fertilization 871
34 Fueling the Body’s Metabolism 789 Concept 36.5 Cells of the Early Embryo Are Totipotent 874
Concept 36.6 Cleavage Leads to the Blastula Stage 878
Concept 34.1 Vertebrate Digestive Systems Are Tubular Tracts 790
Concept 36.7 Gastrulation Forms the Basic Body Plan
Concept 34.2 Food Is Processed as It Passes Through the Digestive of the Embryo 880
Tract 791
Concept 36.8 The Body’s Organs Form in Organogenesis 882
Concept 34.3 The Digestive Tract Is Regulated by the Nervous
System and Hormones 796 Concept 36.9 Human Development Takes Nine Months 885
Concept 34.4 Respiratory Systems Promote Efficient Exchange of Inquiry & Analysis 889
Gases 797 Retracing the Learning Path 890
Concept 34.5 Gills Provide for Efficient Gas Exchange in Water 798 Assessing the Learning Path 891

xx contents
 PART VIII Ecology 39 Community Ecology and Ecosystem
and Behavior 893 Dynamics 944
Concept 39.1 Competition Shapes How Species Live Together in
Mike Powles/Getty Images
Communities 945
37 Behavioral Biology 893 Concept 39.2 Predator–Prey Relationships Foster Coevolution 949
Concept 37.1 An Animal’s Genome Influences Its Behavior 894 Concept 39.3 Cooperation Among Species Can Lead to
Concept 37.2 Learning Also Influences Behavior 896 Coevolution 953
Concept 37.3 Thinking Directs the Behavior of Many Animals 897 Concept 39.4 Ecological Succession Is a Consequence of Habitat
Alteration 958
Concept 37.4 Migratory Behavior Is Both Innate and Learned 898
Concept 39.5 Chemical Elements Move Through Ecosystems in
Concept 37.5 Animal Communication Plays a Key Role in Biogeochemical Cycles 959
Ecological and Social Behavior 900
Concept 39.6 Energy Flows Through Ecosystems in One
Concept 37.6 Behavior Evolves Adaptively 903 Direction 965
Concept 37.7 Behavioral Ecology Examines the Adaptive Concept 39.7 Biodiversity May Increase Ecosystem Stability 969
Significance of Behaviors 904
Inquiry & Analysis 974
Concept 37.8 Behavioral Strategies Have Evolved to Maximize
Reproductive Success 907 Retracing the Learning Path 975
Concept 37.9 Some Behaviors Decrease Fitness to Benefit Other Assessing the Learning Path 977
Individuals 909
Concept 37.10 Group Living Has Evolved in Both Insects and 40 The Living World 979
Vertebrates 912
Concept 40.1 Ecosystems Are Shaped by Sun, Wind, and
Inquiry & Analysis 914 Water 980
Retracing the Learning Path 915 Concept 40.2 Earth Has 14 Major Terrestrial Ecosystems, Called
Assessing the Learning Path 916 Biomes 984
Concept 40.3 Freshwater Habitats Occupy Less Than 2% of Earth’s
38 Ecology of Individuals and Surface 986
Concept 40.4 Marine Habitats Dominate the Earth 987
Populations 919
Concept 40.5 Humanity’s Pollution Is Severely Impacting the
Concept 38.1 Populations Are Groups of a Single Species in Biosphere 989
One Place 920
Concept 40.6 Human Activity Is Altering Earth’s Climate 991
Concept 38.2 Population Growth Depends upon Members’ Age
Inquiry & Analysis 996
and Sex 923
Retracing the Learning Path 997
Concept 38.3 Evolution Favors Life Histories That Maximize
Lifetime Reproductive Success 926 Assessing the Learning Path 999
Concept 38.4 Environment Limits Population Growth 928
Concept 38.5 Resource Availability Regulates Population Appendix: Answer Key A-1
Growth 930
Concept 38.6 Earth’s Human Population Is Growing Index I-1
Explosively 933
Concept 38.7 Pandemics and Human Health 936
Inquiry & Analysis 939
Retracing the Learning Path 940
Assessing the Learning Path 942

contents xxi
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Part I The Molecular Basis of Life

1
The Science of Biology

Lea r ni ng Pa th
1.1 The Diversity of Life Is 1.4 The Study of Evolution Is a
Overwhelming Good Example of Scientific
Inquiry
1.2 Biology Is the Science of Life
1.5 A Few Important Concepts
1.3 Science Is Based on Both Form the Core of Biology
Observation and Reasoning

Soames Summerhays/Natural Visions

Concept Overview
This Concept Overview provides a diagram of the overarching concepts that are covered in this chapter. These concepts will be expanded upon
in the Concept Overview at the end of the chapter.

Biology is the study of living things

Biological organisms are diverse Science uses observation and reasoning Five concepts unify biology

In tr oduct ion
You are about to embark on a journey of discovery about the nature of life. Almost two centuries ago, a young English naturalist named Charles
Darwin set sail on a similar journey on board H.M.S. Beagle; the photo on this page shows a replica of this ship. Darwin’s observations during
the voyage of the Beagle influenced his development of the theory of evolution by natural selection, which has become the core of the science of
biology. Before we begin, however, let’s take a moment to think about what biology is and why it’s important.
 1.1 The Diversity of Life
Is Overwhelming
Biology is the study of living things—literally the science of life.
The living world is teeming with a breathtaking variety of creatures—
whales, butterflies, mushrooms, plants, bacteria—which despite
their obvious differences share features common to all living
­organisms. We will begin with a brief history of how we classify this
diversity.

Domain Archaea. This Domain Bacteria. This

Biological Diversity Can Be Organized prokaryotic domain includes prokaryotic domain includes
this methanogen, which this purple sulfur bacteria,
by Evolutionary History manufactures methane as a which can use light energy to
result of its metabolic activity. drive the synthesis of organic
compounds (false color).
LEARNING OBJECTIVE 1.1.1 Describe the three domains of life.
Figure 1.1 The two prokaryotic domains. Bacteria and
The amazing diversity of life inspired a long history of classifica-
archaea share the feature of lacking a membrane-bounded
tion systems to bring order to this diversity. This culminated in
nucleus. Organisms from both of these domains are single-celled.
the mid-1700s with the Linnaean classification system, which
(Archaea): Power and Syred/Science Source; (Bacteria): Alfred Pasieka/Science
used observable similarities to group organisms into hierarchical Source
groups. This system used seven levels (kingdom, phylum, class,
order, family, genus, species) to classify all organisms. One
kingdom, but clearly does not represent a single line of evolution-
change from previous systems was the introduction of two-part,
ary descent. This has led to attempts to find clear groups with
or ­binomial, names. For instance, humans are Homo sapiens,
shared evolutionary history. We will discuss this in detail in chap-
which is the genus and species description from the longer, com-
ters 22 and 24.
plete classification. Classification based on shared evolutionary
The work of biologists affects your everyday life: what you
history often, but not always, produces results similar to that
eat, what happens to you when you go to the hospital, and how
based on shared characteristics. Although the Linnaean system
our society will handle environmental issues such as climate
does not always reflect evolutionary relationships, it remains use-
change. Unifying the diverse systems studied by biologists are the
ful today because it is the only universal system, and its long
shared characteristics of all living things that have been shaped
history means that an extraordinary amount of biodiversity has
by the process of evolution by natural selection. Keeping this
been classified this way. These issues are considered in detail in
theme in mind will help you to manage the complexity and
chapter 22.
diversity of biology.
The highest level of organization in the Linnaean system is
the kingdom. With the advent of microscopes, we also defined
two basic cell types: those that have genetic material in a REVIEW OF CONCEPT 1.1
­membrane-bounded nucleus (eukaryotes), and those that lack this The living world is incredibly diverse. Various systems of
membrane-bounded nucleus (prokaryotes). This was accommo- classification have been proposed, which remains an area of
dated within the Linnaean system by creating a kingdom for active research. The oldest branching of the tree of life is into
prokaryotes, specifically Bacteria (once called Kingdom Monera). Bacteria, Archaea, and Eukarya.
This was later upended by the discovery that there were actually
two kinds of prokaryotes, called Bacteria (or Eubacteria) and ■■ What are some shared features of living systems?
Archaea (figure 1.1; discussed in detail in chapter 23).
This change led to the addition of a new taxonomic rank
above kingdom called a domain. All living organisms can be
divided into three domains: Eubacteria, Archaea, and Eukarya. 1.2 Biology Is the Science
Even this system has been challenged as there is evidence that of Life
eukaryotes arose from within Archaea, leading some to advocate
for a two-domain system. This remains an area of active research,
In its broadest sense, biology is the study of living things. So it
with many new prokaryotes being identified based on the ability
would seem that biologists would have no problem defining life.
to analyze genetic material without actually visualizing or cultur-
In fact, it is quite difficult to provide a simple definition of life.
ing organisms.
The kingdoms you are probably most familiar with contain
plants, animals, and fungi (figure 1.2), and these represent single Life Defies Simple Definition
lines of evolutionary descent. The group that has been problem-
LEARNING OBJECTIVE 1.2.1 Describe five fundamental
atic is the protists, which were grouped based on the shared simi-
properties of life.
larity of being eukaryotic and unicellular. This was formerly a
2 Part I The Molecular Basis of Life
Protists. Once considered a
kingdom, this term is still used
informally to refer to unicellular
eukaryotes.
Fungi. This kingdom contains
mostly multicellular
nonphotosynthetic organisms
that digest their food
externally, such as mushrooms.
Plantae. This kingdom contains
photosynthetic multicellular
organisms that are terrestrial,
such as the flowering plant
pictured here.
Animalia. Organisms in this
kingdom are nonphotosynthetic
multicellular organisms that
digest their food internally,
such as this ram.

Figure 1.2 The eukaryotic domain. Eukaryotes consist of most familiar organisms and many that are not. The eukaryotes can be
divided into three kingdoms: Fungi, Plantae, and Animalia. Protists are a diverse group of unicellular eukaryotic organisms with no
exclusive common ancestor.
(Protista): Dr. Stan Erlandsen and Dr. Dennis Feely/CDC; (Fungi): Russell Illig/Getty Images; (Plantae): Iconotec/Glow Images; (Animalia): Alan and Sandy Carey/
Getty Images

What does it mean to be alive? What properties define a living
organism? These questions are not as simple as they appear,
because some of the most obvious properties of living organisms
are also properties of many nonliving things—for example, com-
plexity (a computer is complex), movement (clouds move in the
sky), and response to stimulation (a soap bubble pops if you touch
it). To appreciate why these three properties, so common among
living things, do not help us to define life, imagine a mushroom
standing next to a television: the television seems more complex
than the mushroom, the picture on the television screen is mov-
ing but the mushroom just stands there, and the television
responds to a remote-control device but the mushroom continues
to just stand there—yet it is the mushroom that is alive.
All living things also share five more fundamental proper-
ties, passed down over millions of years from the first organisms
to evolve on Earth: cellular organization; energy utilization; homeo-
stasis; growth, development, and reproduction; and heredity.
1. Cellular organization. All living things are composed of one
or more cells. Often too tiny to see, cells carry out the
basic activities of living. Some cells have simple interiors,
whereas others have complex organizations, but all are able
to grow and reproduce. Many organisms possess only a
single cell, like the paramecium in figure 1.3; your body
contains about 10 trillion to 100 trillion cells (depending
on your size).
2. Energy utilization. All living things use energy. Moving,
growing, thinking—everything you do requires energy.
Where does all this energy come from? It is captured from
sunlight by plants and algae through photosynthesis. To get
the energy that powers our lives, we extract it from plants
or from plant-eating animals. That’s what the kingfisher is
doing in figure 1.4, eating a fish that ate algae.
3. Homeostasis. All living things maintain relatively con-
stant internal conditions so that their complex processes
can be better coordinated. Although the environment
often varies considerably, organisms act to keep their
interior conditions relatively constant, a process called
homeostasis. Your body acts to maintain an internal
temperature of 37˚C (98.6˚F), however hot or cold the
weather might be.
4. Growth, development, and reproduction. All living things
can grow and reproduce, although all members of a
species may not reproduce. Bacteria increase in size and
simply split into two, as often as every 15 minutes.
Multicellular organisms grow by increasing the number of
cells, and most produce different kinds of cells during
development.
5. Heredity. All organisms possess a genetic system that is based
on the replication and duplication of a long molecule called
DNA (deoxyribonucleic acid). The information that determines
Figure 1.3 Cellular organization. This paramecium is a
complex single-celled protist that has just ingested several yeast
cells. Like this paramecium, many organisms consist of just a
single cell, while others are composed of trillions of cells.
Melba Photo Agency/Alamy Stock Photo

Chapter 1 The Science of Biology 3


Figure 1.4 Energy utilization. This kingfisher obtains the
energy it needs to move, grow, and carry out its body processes
by eating fish. It harvests the energy from food using chemical
processes that occur within cells.
Armin Floreth/imagebroker/Alamy Stock Photo
what an individual organism will be like is contained in a code
dictated by the order of the subunits making up the DNA
molecule. Because DNA is passed from one generation to the
next, any change in a gene can be preserved and passed on to
future generations. The transmission of characteristics from
parent to offspring is a process called heredity. All organisms
interact with other organisms and the nonliving environment
in ways that influence their survival, and as a consequence,
organisms evolve adaptations to their environments.
CELLULAR LEVEL
1 Atoms 2 Molecule 3 Macromolecule 4 Organelle
H
N
H
N
C
O
0.2 µm
4 Part I The Molecular Basis of Life
Living Systems Show Hierarchical
Organization
LEARNING OBJECTIVE 1.2.2 Describe the hierarchical nature
of living systems.
Life’s organisms interact with each other at many levels, in
ways simple and complex. A key factor organizing these inter-
actions is their degree of complexity. The organization of the
biological world is hierarchical—that is, each level builds on
the level below it, from the very simplest level of individual
Figure 1.5 Hierarchical organization of living systems.
Life is highly organized, from the simplest atoms to complex,
multicellular organisms. Along this hierarchy of structure, atoms
form molecules, which are used to form organelles, which in turn
form the functional subsystems within cells. Cells are organized
into tissues, then into organs and organ systems such as the
nervous system. This organization extends beyond individual
organisms to populations, communities, ecosystems, and finally
the entire biosphere.
(organelle): Keith R. Porter/Science Source; (cell): STEVE GSCHMEISSNER/
SCIENCE PHOTO LIBRARY/Alamy Stock Photo; (tissue): Ed Reschke/Getty Images;
(organism): Russell Illig/Getty Images; (population): George Ostertaga/
gefotostock/Alamy Stock Photo; (species top): USDA Natural Resources
Conservation Service; (species bottom): U.S. Department of Agriculture (USDA);
(community): Ryan McGinnis/Alamy Stock Photo; (ecosystem): Steven P. Lynch/
McGraw Hill; (biosphere): Goddard Space Flight Center/NASA
5 Cell 6 Tissue 7 Organ
100 µm
atoms to the vastly complex level of interacting ecosystems and spinal cord, and a network of neurons that convey
(figure 1.5): signals between the brain and the other organs and tissues
of the body.
The Cellular Level. At the cellular level, atoms 1 , the fundamental
The Populational Level. Individual organisms 9 occupy several
elements of matter, are joined together by chemical bonds
hierarchical levels within the living world. The most basic
into stable assemblies called molecules 2 . Large, complex
of these is the population 10 —a group of organisms of the
molecules are called macromolecules 3 . DNA, which stores
same species living in the same place. All populations of a
the hereditary information, is a macromolecule. Complex
particular kind of organism together form a species 11 , its
biological molecules are assembled into tiny structures called
members are similar in appearance and able to interbreed.
organelles 4 , within which cellular activities are organized.
At a higher level of biological organization, a biological
A mitochondrion is an organelle within which the cell
community 12 consists of all the populations of different
extracts energy from food molecules. Membrane-bounded
species living together in one place.
units called cells 5 are the basic units of life. Bacteria are
The Ecosystem Level. At the highest tier of biological organization,
composed of single cells. Animals, plants, and many other
a biological community and the physical habitat (soil
organisms are multicellular—composed of many cells.
composition, available water, temperature range, wind, and
The Organismal Level. Cells of multicellular organisms
a host of other environmental influences) within which it
exhibit three levels of organization. The most basic
lives together constitute an ecological system, or ecosystem
level is that of tissues 6 , which are groups of similar
13 . The entire planet can be thought of as a global
cells that act as a functional unit. Nerve tissue is one
ecosystem we call the biosphere 14 .
kind of tissue, specialized to carry electrical signals.
Tissues, in turn, are grouped into organs 7 —body
structures composed of several different tissues that act Novel Properties Emerge from
as a structural and functional unit. Your brain is an
organ composed of nerve cells and cells that nourish
More Complex Organization
and support them, as well as a variety of associated LEARNING OBJECTIVE 1.2.3 Discuss how living systems
connective tissues that form both protective coverings display emergent properties.
and a network of blood vessels to bring oxygen and
nutrients to the brain. At the third level of organization, At each higher level in the living hierarchy, novel properties
organs are grouped into organ systems 8 . The nervous emerge—properties that were not present at the simpler level of
system, for example, consists of sensory organs, the brain organization. These emergent properties result from the way in

ORGANISMAL LEVEL POPULATIONAL LEVEL ECOSYSTEM LEVEL

8 Organ system 9 Organism 10 Population 11 Species 12 Community 13 Ecosystem 14 Biosphere

Chapter 1 The Science of Biology 5

which components interact and often cannot be guessed just by
looking at the parts themselves. You have the same array of cell
types as a giraffe, for example—so examining a collection of its
individual cells gives little clue about what your body is like.
The emergent properties of life are not magical or super-
natural. They are the natural consequences of the hierarchy or
structural organization that is the hallmark of life. Both water
(which makes up 50 to 75% of your body’s weight) and ice are
made of H2O molecules, but one is liquid and the other is solid,
because the H2O molecules in ice are more organized. Two
­proteins—long chains of amino acids—may contain the same num-
ber of each amino acid, yet one might act as an enzyme to pro-
mote a chemical reaction whereas the other might not; the
­enzymatic activity is an emergent property, reflecting the infor-
mation contained in the sequence of the amino acids.
Functional properties emerge from more complex organiza-
tion. Metabolism is an emergent property of life. The chemical
reactions within a cell arise from interactions between molecules
that are orchestrated by the orderly environment of the cell’s inte-
rior. Consciousness is an emergent property of the brain that
results from the interactions of many neurons in different parts of
the brain.
This description of the common features and organization
of living systems begins to get at the nature of what it is to be alive.
The rest of this book illustrates and expands on these basic ideas
to provide you with a more complete account of living systems.
REVIEW OF CONCEPT 1.2
Biology is a unifying science that brings together all branches
of science to study living systems. Life does not have a simple
definition, but living systems share a number of properties that
together describe life. Biologists organize living systems hier-
archically, from the subcellular level to the entire biosphere,
with emergent properties arising at each stage that cannot be
guessed from studying its parts.
■■ Can you name an emergent property at the population
level?
1.3 Science Is Based on Both
Observation and Reasoning
Much like life itself, the nature of science defies simple descrip-
tion. For many years scientists have written about the “scientific
method” as though there were a single way of doing science. This
oversimplification has contributed to nonscientists’ confusion
about the nature of science. Although there is no single scien-
tific method, we could say that there is a scientific mindset
that involves skepticism and the importance of objective, verifi-
able facts.
At its core, science is concerned with developing an increas-
ingly accurate understanding of the world by using observation and
reasoning. To begin with, we assume that natural forces acting now
have always acted, that the fundamental nature of the universe has
not changed since its inception, and that it is not changing now.
6 Part I The Molecular Basis of Life
There is a corollary to these assumptions about the action
of natural forces: science does not address the so-called super-
natural. This would seem to set science and religion in opposi-
tion, and there are some who hold this view. Others take the
position that science and religion address different questions or
shed light on different aspects of the human condition. Regard-
less of the position you take, questions of religion, the existence of
ghosts, astrology, and any other supernatural phenomena are out-
side the scope of this book.
The Scientific Process Involves Observation
and Both Deductive and Inductive Reasoning
LEARNING OBJECTIVE 1.3.1 Distinguish between deductive
and inductive reasoning.
Virtually all science begins with observation of the natural world.
This might also involve measurement or use technology that
extends our ability to make observations. The use of technology
has become increasingly important in biology, as we will learn
throughout our explorations.
Descriptive science
The common version of the scientific process is that observations
lead to hypotheses, which in turn make experimentally testable
predictions. In this way, scientists develop an increasingly accu-
rate understanding of nature. We discuss this way of doing sci-
ence later in this section, but it is important to understand that
much of science is purely descriptive: in order to understand any-
thing, the first step is to describe it completely. Much of biology
is concerned with arriving at an increasingly accurate description
of nature.
The study of biodiversity is an example of descriptive sci-
ence that has implications for other aspects of biology, in addition
to societal implications. Efforts are currently under way to clas-
sify all life on Earth. This ambitious project is purely descriptive,
but it will lead to a greater understanding of biodiversity and how
human activity affects biodiversity.
A new approach to biodiversity uses massive DNA sequenc-
ing of environmental samples, followed by using computer soft-
ware to identify organisms in the sample. This has revealed an
even greater than anticipated diversity of both prokaryotic and
eukaryotic organisms than was suspected based on culturing
organisms in the laboratory. At this point our ability to generate
these data is outrunning our ability to directly analyze any of
these organisms.
One of the most important accomplishments of molecular
biology at the dawn of the 21st century was completing the
sequencing of the human genome. Since the initial project was
completed, this descriptive work has been continuing as we refine
and extend our knowledge of the human genome. All of these
data are helping biologists to explore new areas of human biology
and to generate hypotheses about both basic human biology and
the understanding of disease states.
Logical reasoning
Scientists are often portrayed in popular culture as unemotional,
purely logical individuals. Popular television has a long history of
such characters, most recently Dr. Temperance Brennan in the
drama Bones and Dr. Sheldon Cooper in the sitcom The Big Bang
Theory. While entertaining, this is not an accurate portrayal of
most scientists. However, the use of logical reasoning is critical to
the scientific process.
The form of reasoning that applies general principles to pre-
dict specific results is called deductive reasoning. This form of rea-
soning arose from mathematics and philosophy and is used to test
the validity of general ideas in all branches of knowledge. For
example, if all mammals by definition have hair, and you find an
animal that does not have hair, then you may conclude that this
animal is not a mammal. A biologist uses deductive reasoning to
infer the species of an unknown specimen from its characteristics.
More than 2200 years ago, the Greek scientist Eratosthenes used
Euclidean geometry and deductive reasoning to accurately esti-
mate the circumference of the Earth. This is a powerful example
of how the use of observation and logic can help us understand the
natural world.
When logic flows in the opposite direction, from specific
cases to general principles, we call it inductive reasoning. Inductive
reasoning uses specific observations to construct general scientific
principles. For example, if poodles have hair and terriers have hair,
and every dog that you observe has hair, then you may conclude
that all dogs have hair. Inductive reasoning leads to generalizations
that can then be tested. Inductive reasoning first became important
to science in the 17th century in Europe, when Francis Bacon,
Isaac Newton, and others, including Margaret Cavendish and
Jeanne Dumée, began to use the results of particular experiments
to infer general principles about how the world operates.
Hypothesis-Driven Science Makes
and Tests Predictions
LEARNING OBJECTIVE 1.3.2 Illustrate how experimentation
can be used to test hypotheses.
Experimental scientists use a combination of inductive and
deductive reasoning. We use induction to go from a variety of
specific observations to form a general proposal to explain the
observations. We call the proposal that explains our initial obser-
vations a hypothesis. This hypothesis is more than an explanation;
to be useful, it must generate testable predictions. We use the pro-
cess of deduction to generate predictions that usually take an “if
X, then Y” form. These predictions from one or more alternative
hypotheses can be tested by experiments. This leads to a hypoth-
esis being either supported or rejected. A hypothesis that is sup-
ported has not been “proven” and is subject to replication and
additional testing. If it continues to be supported, our confidence
in this as an explanation increases. If a hypothesis must be
rejected, we need to either formulate a new hypothesis or modify
the existing one (figure 1.6).
Because experiments do not actively “prove” a hypothesis,
this process is ongoing as new data are generated. Even founda-
tional ideas in a field will evolve with new information. For
example, geneticists George Beadle and Edward Tatum studied
the nature of genetic information to arrive at the “one-gene/
one-enzyme” hypothesis (refer to chapter 15). This states that
a gene represents the genetic information necessary to make a
Problem
Collect observations
Induction
Generate hypotheses
Deduction
Modify
Generate testable predictions
hypothesis or
generate new
hypothesis
Experimental test of predictions
Falsification Hypothesis supported
Reject hypothesis Replication and new tests
Falsification Hypothesis supported
Modify hypothesis or generate
new hypothesis
Figure 1.6 How experimental science is done. This
provides a general flowchart for testing hypotheses by
experimentation. A problem of interest is identified and
observations are collected. Inductive reasoning leads to
development of one or more potential explanations (hypotheses).
Experimental results will either support or falsify a hypothesis. A
hypothesis that is supported is still subject to further replication
and testing, leading to either eventual rejection or further
support. Falsified hypotheses can be modified or rejected in
favor of a new hypothesis.
single enzyme. As investigators learned more about the molecu-
lar nature of genetic information, the hypothesis was refined to
“one-gene/one-polypeptide,” because enzymes can be made up
of more than one polypeptide. With still more discoveries about
the nature of genetic information, other investigators found that
a single gene can specify more than one polypeptide, and the
hypothesis was refined again.
Testing hypotheses
We call the test of a hypothesis an experiment. Suppose you enter
a dark room. To understand why it is dark, you propose several
hypotheses. The first might be “The room is dark because the
light switch is turned off.” Other hypotheses could involve
whether the switch is functional, whether the light bulb is burned
out, or even loss of power to the entire room. To evaluate these
hypotheses, you would conduct experiments designed to elimi-
nate one or more of the hypotheses.
For example, you might test your hypotheses by flipping the
light switch. If you do so and the room is still dark, you have dis-
proved the first hypothesis: something other than the setting of
the light switch must be the reason for the darkness. Note that a
test such as this does not prove that any of the other hypotheses
Chapter 1 The Science of Biology 7
are true; it merely demonstrates that the one being tested is not. A SCIENTIFIC THINKING
successful experiment is one in which one or more of the alterna-
tive hypotheses are demonstrated to be inconsistent with the Question: What is the source of contamination that occurs in a flask
results and are thus rejected. of nutrient broth left exposed to the air?
As you proceed through this text, you will encounter many Germ Hypothesis: Preexisting microorganisms present in the air
hypotheses that have withstood the test of experiment. Many contaminate nutrient broth.
will continue to do so; others will be revised as biologists make Prediction: Sterilized broth will remain sterile if microorganisms are
new observations. Biology, like all science, is in a constant state prevented from entering flask.
of change, with new ideas appearing and replacing or refining Spontaneous Generation Hypothesis: Living organisms will
old ones. spontaneously generate from nonliving organic molecules in broth.
Prediction: Organisms will spontaneously generate from organic
The importance of controls molecules in broth after sterilization.
Test: Use swan-necked flasks to prevent entry of microorganisms. To
Often scientists are interested in learning about processes that
ensure that broth can still support life, break swan-neck after sterilization.
are influenced by many factors, or variables. To evaluate alterna-
tive hypotheses, we keep the variable of interest constant in a Broken
neck
control group and alter it in the experimental treatment. This of flask
allows us to isolate the effects of a single variable in our experi-
ment, so any difference in the outcome is due to the influence of
this variable. The factors or conditions that you manipulate in
an experiment are called independent variables, and the factor
observed or measured is called the dependent variable. Flask is sterilized Unbroken flask Broken flask becomes
by boiling the broth. remains sterile. contaminated after
For example, in a trial to test the efficacy of a vaccine exposure to germ-laden air.
against a virus, the hypothesis being tested is whether the vaccine Result: No growth occurs in sterile swan-necked flasks. When the
prevents infection. The independent variable is whether a partici- neck is broken off and the broth is exposed to air, growth occurs.
pant received a placebo (control) or the vaccine (experimental), Conclusion: Growth in broth is of preexisting microorganisms.
and the dependent variable is the incidence of disease in the two
groups. Notice in this case there are a variety of possible “end
points” that could be analyzed, including any infection, a reduc- Figure 1.7 Experiment to test spontaneous generation
tion in severity of infection, hospitalizations, or deaths. versus germ hypothesis.

Summing up: An example

This result was predicted by the germ hypothesis—that
A hypothesis that does not make testable predictions will not do
when the sterile flask is exposed to air, airborne germs are depos-
much to advance our understanding. It is also true that a hypoth-
ited in the broth and grow. The spontaneous generation hypothe-
esis that explains everything you have observed is quite possibly
sis predicted no difference in results with exposure to air. This
dead wrong. Or, as Thomas Huxley, a contemporary and sup-
experiment disproved the hypothesis of spontaneous generation
porter of Charles Darwin, put it: “The great tragedy of science,
and supported the hypothesis of airborne germs under the
the slaying of a beautiful theory by an ugly fact.”
conditions tested.
Let’s put this together with an important example from the
early history of microbiology. It was known that nutrient broth
left sitting exposed to air becomes contaminated. Two hypothe-
ses were proposed to explain this observation: spontaneous gen-
Theories Are the Solid Conclusions
eration and the germ hypothesis. Spontaneous generation was an of Science
old idea that had been used to explain a variety of observations,
LEARNING OBJECTIVE 1.3.3 Discuss how scientists use
including why maggots appear in rotting meat. This theory held
models to describe, explain, and test theories.
that there was an inherent property in organic molecules that
could lead to the spontaneous generation of life. The more As we have discussed, a successful hypothesis should generate
recent, at the time, germ hypothesis proposed that preexisting predictions that allow for experimental tests. If an experiment
microorganisms that were present in the air could contaminate produces results inconsistent with predictions, the hypothesis
the nutrient broth. must be rejected or modified. In contrast, if the predictions are
These competing hypotheses were tested by a number of supported by experimental testing, the hypothesis is supported.
experiments that involved filtering air and boiling the broth to The more experimentally supported predictions a hypothesis
kill any contaminating germs. The definitive experiment was per- makes, the stronger our confidence in its validity.
formed by Louis Pasteur, who constructed flasks with curved
necks that could be exposed to air, but that would trap any con-
taminating germs. When such flasks were boiled to sterilize The nature of scientific theories
them, they remained sterile, but if the curved neck was broken off, Scientists use the word theory in two main ways. The first mean-
they became contaminated (figure 1.7). ing of theory is essentially deductive, a proposed explanation for

8 Part I The Molecular Basis of Life

some natural phenomenon, based on general principles. Thus, we
speak of the principle first proposed by Newton as the “theory of
gravity.” Such theories often bring together concepts that were
previously thought to be unrelated.
The second meaning of theory is essentially inductive: a
body of interconnected concepts, supported by inductive scien-
tific reasoning and experimental evidence, that explains the facts
in some area of study. For example, quantum theory in physics
brings together a set of ideas about the nature of the universe
derived from diverse experimental observations and serves as a
guide to further questions and experiments.
To scientists, theories are the solid ground of science,
expressing ideas about which they are most certain. By contrast,
to the general public the word theory usually implies the opposite—
a lack of knowledge or a guess (“it’s only a theory . . .”). Not
surprisingly, this difference often results in confusion. In this
text, theory will always be used in its scientific sense, in reference
to an accepted general principle or body of knowledge.
Reductionism
Scientists often use the philosophical approach known as
reductionism to understand a complex system by reducing it to its
working parts. Reductionism has limits when applied to living
systems, however—the complex interworking of many intercon-
nected functions leads to emergent properties that cannot be pre-
dicted based on the workings of the parts. For example,
ribosomes—complex cellular machines that make proteins—can
be disassembled into their constituent parts. However, examination
of the parts in isolation would not lead to predictions about the
nature of protein synthesis. On a higher level, understanding the
physiology of a single Canada goose would not lead to predictions
about flocking behavior. Biologists are just beginning to come to
grips with this problem and to think about ways of dealing with
the whole as well as the workings of the parts. The emerging field
of systems biology focuses on this different approach.
Biological models
Biologists construct models in many different ways for a variety
of uses. Geneticists construct models of interacting networks of
proteins that control gene expression, often even drawing cartoon
figures to represent that which we cannot see. Population biolo-
gists build models of how evolutionary change occurs. Cell biolo-
gists build models to explain cell communication and the events
leading from an external signal to internal events. Structural
biologists build models of the structure of proteins and macromo-
lecular complexes in cells.
Models provide a way to organize how we think about a
problem. Models can also get us closer to the larger picture and
away from the extreme reductionist approach. The working parts
are provided by the reductionist analysis, but the model shows
how they fit together. Often these models suggest other experi-
ments that can be performed to refine or test the model.
Science as a social construct
Research results are written up and submitted for publication in
scientific journals, where other scientists review the experiments
and conclusions. This process of evaluation of others’ work lies at
the heart of modern science. It reduces the incidence of faulty
research or false claims being given the authority of scientific
fact, and provides a starting point for examining the reproduc-
ibility of experimental results. Results that cannot be reproduced
are not taken seriously for long.
Even with evaluation by other scientists, there are issues
with the reproducibility of experiments in multiple branches of
science. Some of this has to do with misusing statistics, and some
has to do with other issues, including the pressure to publish and
the unwillingness of journals to publish negative results. There
have also been a number of well-publicized examples of outright
scientific fraud. While these are serious issues, that they are being
debated indicates that science can be self-correcting.
There are also examples of theories whose acceptance
led to resistance to alternative explanations. Thomas Kuhn
described the progress of science as accepted dogma that is
occasionally overturned by revolutionary new ideas. This is a
great oversimplification of his ideas, but in some instances this
appears to have been the case. The theory of relativity in physics
is a good example. Our understanding of gravity that goes back
to Newton was eventually replaced by Einstein’s theory of gen-
eral relativity.
This positive view of science and scientists should not
obscure the fact that, like all other areas of human endeavor, the
history of science includes examples of misuse of its perceived
authority by scientists and politicians. The eugenics movement in
the 20th century was used to justify forced sterilization of so-
called “mentally unfit” minorities, and the history of medical sci-
ence includes examples of terrible experiments on minorities
without consent. These abuses have led to more robust systems of
consent for experiments involving humans, but provide a warning
to be vigilant about abuses.
REVIEW OF CONCEPT 1.3
Much of science is descriptive, amassing observations to gain
an accurate view. Both deductive and inductive reasoning are
used in science. Scientific hypotheses are suggested explana-
tions for observed phenomena. When a hypothesis has been
extensively tested and no contradictory information has been
found, it becomes an accepted theory. Theories are coherent
explanations of observed data, and they may be modified by
new information.
■■ How does a scientific theory differ from a hypothesis?
1.4 The Study of Evolution Is a Good
Example of Scientific Inquiry
Darwin’s theory of evolution explains how organisms on Earth
have changed over time and acquired a diversity of new forms. We
devote three chapters to a detailed examination of evolution, but
it is worth considering this theory now as an example of how a
scientist develops a hypothesis and how a scientific theory is
tested and gains acceptance.
Chapter 1 The Science of Biology 9
The Idea of Evolution Existed
Prior to Darwin
LEARNING OBJECTIVE 1.4.1 Describe ideas about evolution
proposed before Darwin.
Charles Robert Darwin (1809–1882; figure 1.8) was an English
naturalist who, after 30 years of study and observation, wrote
one of the most famous and influential books of all time. This
book, On the Origin of Species by Means of Natural Selection, cre-
ated a sensation when it was published, and the ideas Darwin
expressed in it have played a central role in the development of
human thought.
Birth of the idea of evolution
In Darwin’s time, most people believed that the different kinds of
organisms and their individual structures resulted from direct
actions of a Creator (many people still believe this). Species were
thought to have been specially created and to be unchangeable
over the course of time. This was the view of Carolus Linnaeus
(1708–1778), the Swedish biologist who established the system of
naming organisms that is still in use.
By the first part of the 18th century, many more kinds of
organisms were being discovered than previously, as well as many
fossil animals and plants. These discoveries gradually began to
trigger discussions of evolution—the possibility that living things
have changed during the history of life on Earth. The great French
biologist Georges-Louis Leclerc, Comte de Buffon (1707–1788),
spoke explicitly, a century before Darwin, of natural affinities
between kinds of organisms, writing of “the universal kinship of
all generations born from a common mother.” He could see no
Figure 1.8 Charles Darwin. This newly rediscovered
photograph, taken in 1881, the year before Darwin died, appears
to be the last ever taken of the great biologist.
Huntington Library/Superstock
10 Part I The Molecular Basis of Life
explanation for the common features of all mammals except their
evolution from a common ancestor.
Within 50 years these ideas led Jean Baptiste de Lamarck
(1744–1829) to explicitly propose evolution as a theory to account
for the patterns observed in nature. In 1801 he suggested that all
species, including human beings, were descended from other spe-
cies. Lamarck thought of life as having evolved progressively from
simple to more complex forms, and he was the first to propose a
coherent theory of evolution.
Lamarck’s theory was based on the incorrect idea that
organs and structures became stronger through use, and that the
strengthened character was then passed on to offspring—the
­theory of inheritance of acquired characteristics. Although incor-
rect, Lamarck’s theory called wide attention to the possibility of
evolution and, by doing so, set the stage for the acceptance of the
correct, and much simpler, explanation proposed by Charles
Darwin half a century later.
Darwin attributed evolution to what he called natural selec-
tion, which he proposed as a coherent, logical explanation. His
book On the Origin of Species was a best seller in its day and
brought his ideas to wide public attention.
Darwin Gathered Information During
the Voyage of the Beagle
LEARNING OBJECTIVE 1.4.2 Identify important observations
made by Darwin on the Beagle.
The story of Darwin and his theory of evolution begins in 1831,
when Darwin was 22 years old. He was part of a five-year naviga-
tional mapping expedition around the coasts of South America
­(figure 1.9), aboard H.M.S. Beagle. During this long voyage,
­Darwin had the chance to study a wide variety of plants and
­animals on continents, islands, and distant seas. Repeatedly,
­Darwin saw that the characteristics of similar species varied
somewhat from place to place. These geographical patterns sug-
gested to him that lineages change gradually as species migrate
from one area to another. On the Galápagos Islands, 960 km
(600 miles) off the coast of Ecuador, Darwin encountered a vari-
ety of finches on the various islands. The 14 species, although
related, differed slightly in appearance, particularly in their
beaks (figure 1.10).
Darwin thought it was reasonable to assume that all these
birds had descended from a common ancestor that had arrived
from the South American mainland several million years ago.
Eating different foods on these islands, the finches’ beaks had
changed during their descent—“descent with modification,” or
evolution. (These finches are discussed in more detail in chapters 20
and 21.)
In a more general sense, Darwin was struck by the fact
that the plants and animals on these relatively young volcanic
islands resembled those on the nearby coast of South America.
If each one of these plants and animals had been created inde-
pendently and simply placed on the Galápagos Islands, why
didn’t they resemble the plants and animals of islands with
similar climates—such as those off the coast of Africa? Why did
they resemble those of the adjacent South American coast
instead?
 British Isles
EUROPE
Western ASIA
Isles
NORTH
AMERICA
NORTH NORTH NORTH
PAC I F I C AT L A N T I C Canary
OCEAN PAC I F I C
OCEAN Islands OCEAN
Cape Verde Philippine
Galápagos Islands AFRICA Islands
Islands INDIAN
SOUTH O C E A N Keeling Equator
AMERICA Ascension Madagascar Islands
Bahia Friendly
Marquesas St. Helena Mauritius Islands
Valparaiso Rio de Janeiro Bourbon Island AUSTRALIA

Society Sydney
Islands Montevideo Cape of
Buenos Aires Good Hope King George’s
Port Desire Sound
Hobart
New
Straits of Magellan SOUTH Zealand
Falkland AT L A N T I C
Islands OCEAN
Cape Horn Tierra del Fuego

Figure 1.9 The five-year voyage of H.M.S. Beagle. Most of the time was spent exploring the coasts and coastal islands of
South America, such as the Galápagos Islands. Darwin’s studies of the animals of the Galápagos Islands played a key role in his
eventual development of the concept of evolution by means of natural selection.

Woodpecker Finch (Cactospiza pallida) Large Ground Finch (Geospiza magnirostris) Cactus Finch (Geospiza scandens)

Figure 1.10 Three Galápagos finches and what they eat. On the Galápagos Islands, Darwin observed 14 species of finches
that differ mainly in their beaks and feeding habits. These three finches eat very different food items, and Darwin surmised that the
shapes of their bills are evolutionary adaptations that improve their ability to eat the foods available in their specific habitats.

Darwin Proposed Natural Selection Darwin and Malthus

as a Mechanism for Evolution
LEARNING OBJECTIVE 1.4.3 Describe Darwin’s theory
of evolution by natural selection.
Darwin’s two great achievements were to observe evidence that
evolution has occurred and to formulate a hypothesis that explains
this evolution as the consequence of natural selection.
Of key importance to the development of Darwin’s insight into
natural selection was his study of Thomas Malthus’s An Essay on
the Principle of Population (1798). In this book Malthus stated that
populations of plants and animals (including human beings) tend
to increase geometrically, but humans are able to increase their
food supply only arithmetically. Put another way, Malthus argued
that population increases by a multiplying factor; for ­example, in
Chapter 1 The Science of Biology 11

geometric progression
arithmetic progression 54
18
6
11
2 8
5
Figure 1.11 Geometric and arithmetic progressions.
A geometric progression increases by a constant factor (for
example, ×3 for each step), whereas an arithmetic progression
increases by a constant difference (for example, +3 for each step).
Malthus contended that the human growth curve is geometric,
but the human food production curve is only arithmetic.
the series 2, 6, 18, 54, the multiplying factor is 3. By contrast,
Malthus believed that a food supply increases by an additive fac-
tor; for example, the series 2, 5, 8, 11 adds 3 to each starting
number. Figure 1.11 shows the difference that these two types of
relationships produce.
Malthus pointed out that because populations increase geo-
metrically, virtually any kind of animal or plant, if it reproduced
unchecked, would cover the entire surface of the world surpris-
ingly quickly. This does not happen, Malthus argued, because
death limits population numbers so that populations of species
remain fairly constant year after year.
Sparked by Malthus’s ideas, Darwin saw that although
every organism has the potential to produce more offspring than
can ­survive, only a limited number actually do survive and pro-
duce further offspring. Combining this observation with what he
had seen on the voyage of the Beagle, as well as with his own
experiences in breeding domestic animals, Darwin made an
important association: individuals possessing physical, behav-
ioral, or other attributes that give them an advantage in their envi-
ronment are more likely to survive and reproduce than are those
with less advantageous traits. By surviving, these individuals gain
the opportunity to pass on their favorable characteristics to their
­offspring. As the frequency of these characteristics increases in
the population, the nature of the population as a whole gradually
changes. Darwin called this process selection.
Natural selection
Darwin was thoroughly familiar with variation in domesticated
animals, and he began On the Origin of Species with a detailed
discussion of pigeon breeding. He knew that animal breeders
12 Part I The Molecular Basis of Life
selected certain varieties of pigeons and other animals, such as
dogs, to produce certain characteristics, a process Darwin called
artificial selection.
Artificial selection often produces a great variation in traits.
Domestic pigeon breeds, for example, show much greater variety
than all of the wild species found throughout the world. Darwin
thought that this type of change could occur in nature, too. Surely if
pigeon breeders could foster variation by artificial selection, nature
could do the same—a process Darwin called natural selection.
Darwin drafts his argument
Darwin drafted the overall argument for evolution by natural
selection in a book-length preliminary manuscript in 1842. After
showing the manuscript to a few of his closest scientific friends,
however, Darwin put it in a drawer and for 16 years turned to
other research. No one knows for sure why Darwin did not pub-
lish his initial manuscript—it is very thorough and outlines his
ideas in detail.
The stimulus that finally brought Darwin’s hypothesis into
print was an essay he received in 1858 from Alfred Russel
Wallace (1823–1913), a young naturalist who was in Indonesia. It
concisely set forth the hypothesis of evolution by means of natu-
ral selection, a hypothesis Wallace had developed independently
of Darwin. After receiving Wallace’s essay, friends of Darwin
arranged for a joint presentation of the two men’s ideas at a semi-
nar in ­London. Darwin then completed his own book, expanding
the 1842 manuscript he had written so long ago and submitting it
for publication the following year.
The Predictions of Darwin’s Theory
Have Been Well Tested
LEARNING OBJECTIVE 1.4.4 Identify how evolution has been
tested over time.
More than 130 years have elapsed since Darwin’s death in 1882.
During this period, the evidence supporting his theory has grown
progressively stronger. We briefly explore some of this evidence
here; in chapter 20, we will return to the theory of evolution by
natural selection and examine the evidence in more detail.
The fossil record
Darwin predicted that the fossil record would yield intermediate
links between the great groups of organisms—for example,
between fishes and the amphibians thought to have arisen from
them, and between reptiles and birds. Furthermore, natural selec-
tion predicts the relative positions in time of such ­transitional
forms. We now know the fossil record to a degree unthinkable in
the 19th century, and paleontologists have found what appear to
be transitional forms at the predicted p ­ ositions in time.
Recent discoveries of microscopic fossils have extended the
known history of life on Earth back to about 3.5 billion years ago
(bya). The discovery of other fossils has supported Darwin’s
­predictions and has shed light on how organisms have, over this
enormous time span, evolved from the simple to the complex. For
vertebrate animals especially, the fossil record is rich and exhibits
a graded series of changes in form, where the evolutionary
sequence can be discerned (figure 1.12).
 Comparative anatomy
Comparative studies of animals have provided strong evidence
for Darwin’s theory. In many different types of vertebrates, for
­example, the same bones are present, indicating their shared
10
­evolutionary past. Thus, the forelimbs shown in figure 1.13 are all
constructed from the same basic array of bones, modified for
­different purposes.
Early modern whale These bones are said to be homologous in the different
­vertebrates; that is, they have the same evolutionary origin, but
20 they now differ in structure and function. They are contrasted with
analogous structures, such as the wings of birds and butterflies,
which have similar function but different evolutionary origins.

Molecular evidence
Millions of years ago

Evolutionary patterns are also revealed at the molecular level.

30
By comparing the genomes (that is, the sequences of all the genes)
of different groups of animals or plants, we can more precisely spec-
ify the degree of relationship among the groups. A series of evolu-
Basilosaurus tionary changes over time should involve a continual ­accumulation
of genetic changes in the DNA.
40 With the recent advances in genome sequencing, this pre-
diction is now subject to direct test (figure 1.14). The result is
clear: for a broad array of vertebrates, the more distantly related
Rodhocetus two organisms are, the greater their genomic distance.
This pattern of divergence over time is also apparent at the
protein level. Comparing the hemoglobin amino acid sequences
50 Ambulocetus of different species, the pattern is again clear. Rhesus monkeys,
which (like humans) are primates, have fewer differences from
humans in the 146-amino-acid hemoglobin β-chain than do more
Anthracotheres distantly related mammals, such as dogs. Nonmammalian verte-
brates, such as birds and frogs, differ even more.
The sequences of some genes, such as the ones specifying
the hemoglobin proteins, have been determined in many organ-
isms, and the entire time course of their evolution can be laid
Figure 1.12 The evolutionary history of whales. Over out with confidence by tracing the origins of particular nucleo-
a period of 35 million years, modern whales have evolved from tide changes in the gene sequence. The pattern of descent
the piglike ancestors of a hippopotamus, with intermediate steps obtained is called a phylogenetic tree. It represents the evolution-
preserved in the fossil record. ary history of the gene, its “family tree.” Molecular phylogenetic

Human Cat Bat Porpoise Horse

Figure 1.13 Homology among vertebrate limbs. The forelimbs of these five vertebrates show the ways in which the relative
proportions of the forelimb bones have changed in relation to the particular way of life of each organism.

Chapter 1 The Science of Biology 13

 100 91 77 79 72 53 38 36 51 37 31 20 7 7 4 3 2 2

Chimpanzee

Marmoset

Opossum

Zebrafish
Macaque

Tetradon
Elephant

Platypus
Baboon

Chicken
Human

Galago

Mouse

Fugu
Frog
Cow
Dog
Rat
Family

New World
Old World

Prosimians
Monkeys

Monkeys
Superfamily Apes

Pachyderms
Carnivores

Ungulates

Minnows
Rodents

Monotremes
Suborder Anthropoids

Puffers
Marsupials
Order Primates

Amphibians
Subclass Placentals

Birds

Fish
Class Mammals

Subphylum Vertebrates

Figure 1.14 Evolution of the vertebrate genome. Genomic scientists have recently investigated the similarity of 44 representative
regions scattered around the vertebrate genome (1% of the total genome). Moving from right to left, genomic similarity (expressed along
the top as % sequence similarity to humans) increases as taxonomic distance from humans decreases—just as Darwin’s theory predicts.

trees agree well with those derived from the fossil record, which each chapter, in the Learning Path, we use concepts that are
is strong, direct evidence of evolution. The pattern of accumulat- derived from these five most fundamental concepts.
ing DNA changes represents, in a real sense, the footprints of The five concepts are consistent with the past organization
evolutionary history. of this text and with national attempts—such as Vision and
Change—to elaborate critical biological concepts. The five funda-
REVIEW OF CONCEPT 1.4 mental concepts are (1) life is subject to chemical and physical
laws; (2) structure determines function; (3) living systems
Darwin observed differences in related organisms and pro- ­transform energy and matter; (4) living systems depend on infor-
posed the hypothesis of evolution by natural selection to mation transactions; and (5) evolution explains the unity and
explain these differences. The predictions generated by natural diversity of life.
selection have been tested and continue to be tested by analy-
sis of the fossil record, comparative anatomy, and even the DNA
Five Concepts Unify Biology
of living organisms.
■■ Does Darwin’s theory of evolution by natural selection LEARNING OBJECTIVE 1.5.1 Describe five unifying concepts
explain the origin of life? of biology as a science.

1. Life is subject to chemical and physical laws.

1.5 A Few Important Concepts
Form the Core of Biology
The depth and breadth of material in introductory biology create
challenges for students new to college. Consequently, in recent
years there have been initiatives to emphasize conceptual under-
standing over factual recall, and to find ways to actively engage
students in the learning process. In this book five major concepts
unify our discussion of diverse biological topics. We will intro-
duce these concepts here, but they form the underlying logic
behind the organization of the entire book. At the beginning of
It may seem obvious, but it is important to emphasize that liv-
ing systems operate according to known chemical and physical
principles. For this reason, almost all introductory textbooks,
including this one, begin with several sections on chemistry.
This is because biological systems are the ultimate application
of some very complex chemistry. However, no new chemical or
physical laws are found in biology, just the consistent applica-
tion of familiar chemical principles and laws. For the student,
this means that some knowledge of atomic structure, chemical
bonding, thermodynamics, kinetics, and many other topics from
basic chemistry and physics is crucial for understanding bio-
logical systems.

14 Part I The Molecular Basis of Life

 It may seem that some physics and chemistry would only
be relevant in the “cell and molecular” sections of the book, but
in fact, those principles are applied throughout the book. The
movement of water in plants depends on the basic chemistry of
water, the kidney is an osmotic machine, energy flow and nutrient
cycling in ecosystems are driven by thermodynamic laws, and
the cycling of many important elements involves biogeochemical
cycles.
2. Structure determines function.
A major unifying theme of biology is the relationship between
structure and function. We can think about this relationship in
several ways. The proper functioning of molecules, of cells, and
of tissues and organs arises from their three-dimensional struc-
ture. We can learn about the function of molecules by studying
their structure, and we will encounter many examples of this
throughout this book. We can also investigate the function of a
molecule by experimentally altering its structure, then analyz-
ing the effect on function.
We can also work in the opposite direction: if we know
the function of a protein and find a similar protein in a differ-
ent organism, we can infer that the function will be similar. For
example, suppose that we know the structure of a human cell’s
surface receptor for insulin, the hormone that controls the
uptake of glucose. We then find a similar molecule in the mem-
brane of a cell from a very different species, such as a worm.
We might conclude that this membrane molecule acts as a
receptor for an insulin-like molecule produced by the worm.
This would also imply an evolutionary relationship between
glucose uptake in worms and in humans.
3. Living systems transform energy and matter.
From single cells to the highest level of biological organization,
the biosphere, living systems have a constant need for energy. If
we trace this all the way back, the original energy source for the
biosphere is the Sun. Without this energy, living systems would
not exhibit their characteristic highly organized state. This
sounds simple, but it means that the basic nature of life is to con-
stantly transform both energy and matter. We break down “food”
molecules for energy, then use this energy to build up other
complex molecules.
The energy from the Sun is trapped by photosynthetic
organisms, which use this energy to reduce CO2 and produce
organic compounds. The rest of us, who need a constant source
of energy and carbon, can oxidize these organic compounds back
to CO2, releasing energy to drive the processes of life. As all of
these energy transactions are inefficient, a certain amount of
energy is also randomized as heat.
This constant input of energy allows living systems to
function far from thermodynamic equilibrium. At equilibrium,
you are a pool of amino acids, nucleotides, and other small
­molecules, and not the complex dynamic system reading this
sentence. Nonequilibrium systems also can exhibit the property
of self-organization not present in equilibrium systems. Macro­
mole­cular complexes, such as the spindle necessary for chromo-
some separation, can self-organize (figure 1.15). A flock of
birds, a school of fish, and the bacteria in a biofilm all also
Figure 1.15 The spindle. In this dividing cell, microtubules
have organized themselves into a spindle (stained red), pulling
each chromosome (stained blue) to the central plane of the
dividing cell.
Waheeb Heneen, the Swedish University of Agricultural Sciences
display self-organization, exhibiting properties not seen in the
individual parts alone.
4. Living systems depend on information
transactions.
The most obvious form of information in living systems is the
genetic information carried in every cell in the form of deoxyribo-
nucleic acid (DNA). Each DNA ­molecule is formed from two long
chains of building blocks, called nucleotides, wound around each
other (figure 1.16). Four different nucleotides are found in DNA,
and the sequence in which they occur encodes the information to
make and maintain a cell.
The continuity of life from one generation to the next—
heredity—depends on the faithful copying of a cell’s DNA into
daughter cells. The entire set of DNA instructions that speci-
fies a cell is called its genome. The sequence of the human
genome, 3 billion nucleotides long, was decoded in rough-draft
form in 2001.
The importance of information goes beyond genomes and
their inheritance. Cells are highly complex nanomachines that
receive, process, and respond to information. The information
stored in DNA is used to direct the synthesis of cellular compo-
nents, and the particular set of components can differ from cell to
cell. The way proteins fold in space is a form of information that
is three-dimensional, and interesting properties emerge from the
interaction of these shapes in macromolecular complexes. The
control of gene expression allows the differentiation of cell types
in time and space, leading to changes over developmental time
into different tissue types—even though all cells in an organism
carry the same genetic information.
Chapter 1 The Science of Biology 15

Figure 1.16 A model of DNA. All organisms store their
hereditary information as sequences of DNA subunits, much as
this textbook stores information as sequences of alphabet letters.
LAGUNA DESIGN/Science Photo Library/Alamy Stock Photo
Living systems are able to collect information about the
environment, both internal and external, and then respond to this
information. As you are reading this page, you are familiar with
this process, but it also occurs at the level of cells, in terms of
both single-celled organisms and the cells of multicellular organ-
isms. Cells acquire information about their environment, send
and receive signals, and respond to all of this information with
signal transduction systems that can change cell morphology,
behavior, or physiology (the subject of chapter 9).
5. Evolution explains the unity
and diversity of life.
Biologists agree that all organisms alive today on Earth descended
from a simple cellular organism that arose about 3.5 bya. Some of
the characteristics of that organism have been preserved through
evolutionary history into the present. The storage of hereditary
information in DNA, for example, is common to all living things.
The retention of conserved characteristics in a long line of
descent implies that they are important to the success of the
16 Part I The Molecular Basis of Life
Figure 1.17 Fossil trilobites. Fossil organisms preserved in
the Earth’s rocks provide a vivid record of evolution.
Dave Sangster/Getty Images
organism. We see this at the visible, morphological level, and also
at the level of molecules. A quick glance at figure 1.14 shows us
that all mammals have four limbs, and overall anterior to poste-
rior polarity. Regarding molecules, in the same figure it is clear
that more related species have more similar genomes. This is also
true of many important proteins, such as the oxygen carrier
hemoglobin, which is very similar in all organisms that use it to
carry oxygen.
The unity of life that we see in certain key characteristics
shared by many related life-forms contrasts with the incredible
diversity of living things in the varied environments of Earth.
The underlying unity of biochemistry and genetics argues that
all life evolved from the same origin event. The incredible diver-
sity of life apparent today arose by evolutionary change, much of
it visible in the fossil record (figure 1.17).
REVIEW OF CONCEPT 1.5
Understanding biology requires higher-level concepts. We are
using five unifying concepts throughout the book: life is ­subject
to chemical and physical laws, structure determines function,
living systems transform energy and matter, living systems
­
depend on information transactions, and evolution explains the
unity and diversity of life.
■■ Why are there no unique laws of chemistry found in
biology?
Does the Presence of One

Inquiry & Analysis

Species Limit the Population
Size of Others?
Implicit in Darwin’s theory of evolution is the idea that
­species in nature compete for limited resources. Does
this really happen? Some of the best evidence of competi-
tion between species comes from experimental field
­studies, studies conducted not in the laboratory but out
in natural populations. By setting up experiments in
which two ­species occur either alone or together, scientists
can ­determine whether the presence of one species has a
­negative impact on the size of the population of the other
species. The experiment discussed here concerns a variety
Rick & Nora Bowers/Alamy Stock Photo
of seed-eating rodents that live in North American des-
erts. In 1988, researchers set up a series of 50-m × 50-m
enclosures to investigate the effect of kangaroo rats on
smaller seed-eating rodents. Kangaroo rats were removed
from half of the enclosures but not from the other Effect of Kangaroo Rats on Smaller Rodents
­enclosures. The walls of all the enclosures had holes that
allowed rodents to come and go, but in plots without Kangaroo rats removed
15 Kangaroo rats present

Number of captures of
­kangaroo rats the holes were too small to allow the
­kangaroo rats to enter.

other rodents
The graph displays data collected over the course of
the next three years as researchers monitored the number
of the smaller rodents present in the enclosures. To estimate
the population sizes, researchers determined how many
small rodents could be captured in a fixed interval. Data
were ­collected for each enclosure immediately after the
­kangaroo rats were removed in 1988, as well as at ­three-month
intervals thereafter. The graph presents the relative popula-
tion size—that is, the total number of captures averaged over
the number of enclosures. (For example, if a total of 30 rats
were captured from three enclosures, the average would be
10 rats.) The data show the number of small rodents for
­several years after removal of the kangaroo rats.
Analysis
1. Applying Concepts
a. Variable. In the graph, what is the dependent
variable?
b. Relative magnitude. Which of the two kinds of
enclosures maintains the higher population of
small rodents? Does it have kangaroo rats, or have
they been removed?
2. Interpreting Data
a. What is the average number of small rodents
in each of the two plots immediately after
kangaroo rats were removed? After one year?
After two?
b. At what point is the difference between the two
kinds of enclosures the greatest?
10
5
0
1988 1989 1990 1991
3. Making Inferences
a. What precisely is the observed impact of
kangaroo rats on the population size of
small rodents?
b. Examine the magnitude of the difference
between the number of small rodents in the two
plots. Is there a trend?
4. Drawing Conclusions Do these results support the
hypothesis that kangaroo rats compete with other
small rodents to limit their population sizes?
5. Further Analysis
a. Can you think of any cause other than
competition that would explain these results?
Suggest an experiment that could eliminate or
confirm this alternative.
b. Do the populations of the two kinds of enclosures
change in synchrony (that is, grow and shrink at
the same times) over the course of a year? If so,
why might this happen? How would you test this
hypothesis?

Chapter 1 The Science of Biology 17

Retracing the Learning Path
CONCEPT 1.1 The Diversity of Life Is Overwhelming
1.1.1 Biological Diversity Can Be Organized by
Evolutionary History Life on Earth is very diverse but has
traditionally been grouped into six kingdoms based on shared
characteristics.
CONCEPT 1.2 Biology Is the Science of Life
1.2.1 Life Defies Simple Definition Although life is
difficult to define, living systems have eight characteristics in
common: they are capable of movement; are complex and highly
ordered; can respond to stimuli; are composed of one or more cells;
use energy to accomplish work; can maintain relatively constant
internal conditions (homeostasis); can grow, develop, and
reproduce; and can transmit genetic information to their
offspring, making them capable of evolutionary adaptation to
the environment.
1.2.2 Living Systems Show Hierarchical Organization
The hierarchical organization of living systems progresses from
atoms to complex organisms to the entire biosphere.
1.2.3 Novel Properties Emerge from More Complex
Organization As biological systems become more complex,
emergent properties arise that could not be predicted from the sum
of the parts.
CONCEPT 1.3 Science Is Based on Both
Observation and Reasoning
1.3.1 The Scientific Process Involves Observation and Both
Deductive and Inductive Reasoning Science is concerned
with developing an increasingly accurate description of nature
through observation and experimentation. Science uses deductive
reasoning, applying general principles to predict specific results, and
inductive reasoning, using specific observations to construct general
scientific principles.
1.3.2 Hypothesis-Driven Science Makes and Tests
Predictions A hypothesis is constructed based on observations,
and it must generate experimentally testable predictions.
Experiments involve a manipulated variable and a control.
Hypotheses are rejected if their predictions cannot be verified by
observation or experiment.
1.3.3 Theories Are the Solid Conclusions of Science
Reductionism attempts to understand a complex system by
breaking it down into its component parts, but parts may act
differently when isolated from the larger system. Biologists
construct models to explain living systems. A model provides a
different way to study a problem, and it may suggest experimental
approaches. Scientists use the word theory in deductive and
inductive ways: as a proposed explanation for a natural
phenomenon and as a body of concepts that explains facts in
an area of study.
18 Part I The Molecular Basis of Life
CONCEPT 1.4 The Study of Evolution Is a Good
Example of Scientific Inquiry
1.4.1 The Idea of Evolution Existed Prior to Darwin
A century before Darwin, naturalists suggested that living things had
evolved over the course of Earth’s history.
1.4.2 Darwin Gathered Information During the Voyage of
the Beagle During the voyage of H.M.S. Beagle, Darwin had an
opportunity to observe worldwide patterns of diversity.
1.4.3 Darwin Proposed Natural Selection as a Mechanism
for Evolution Darwin noted that species produce many offspring,
of which only a limited number survive and reproduce. He proposed
that individuals possessing traits that increase survival and
reproductive success become more numerous in populations over
time. This is the essence of descent with modification (natural
selection). Alfred Russel Wallace independently came to the same
conclusions from his own studies.
1.4.4 The Predictions of Darwin’s Theory Have Been
Well Tested Natural selection has been tested using data from
many fields. Among these are the fossil record; comparative anatomy
and the study of homologous structures; and molecular data that
provide evidence for changes in DNA and proteins over time. Taken
together, these findings strongly support evolution by natural
selection.
CONCEPT 1.5 A Few Important Concepts
Form the Core of Biology
1.5.1 Five Concepts Unify Biology
1. Life is subject to chemical and physical laws. All
living systems function based on the laws of chemistry and
physics.
2. Structure determines function. The function of
macromolecules is dictated by and dependent on their
structure. Similarity of structure and function may indicate an
evolutionary relationship.
3. Living systems transform energy and matter. Living
systems have a constant need for energy, which is ultimately
provided by the Sun. The nature of life is to constantly
transform energy. We break down food molecules to provide
energy to build up complex structures.
4. Living systems depend on information transactions.
Hereditary information found in the DNA molecule is passed
on from one generation to the next. This information is read
out to produce proteins, which themselves have information
in their structures. Living systems can also acquire
information about their environment.
5. Evolution explains the unity and diversity of life.
The underlying similarities in biochemistry and genetics
support the contention that all life evolved from a single
source. The diversity found in living systems arises by
evolutionary change.
Concept Overview
This Concept Overview diagrams the key concepts that were discussed in this chapter.

Biology is the study of living things

Biological organisms are diverse Science uses observation and reasoning Five concepts unify biology

Life is classified Shared properties Living systems show Hypotheses make The process of Organisms adhere
based on shared define living hierarchical predictions that science is to chemical
evolutionary history organisms organization are tested by exemplified by and physical laws
experiments Darwin’s work
The structure of
Levels go from biological
Organisms are Organisms are atoms to Theories are Evolution is components
divided into three composed of ecosystems supported by based on determines their
domains, one or more cells extensive data evidence properties and
Bacteria, Archaea, Cellular functions
and Eukarya Living things level
use energy Complex systems Biological energy
Organismal can be reduced transformations
level to simpler parts drive life
Organisms maintain processes
homeostasis
Population
level Organisms perceive,
Living things grow express, and respond
and reproduce Ecosystem to internal and
level external information
Genetic information
passes from parent Complex All organisms
to offspring organization can share a common
lead to emergent ancestor
properties

Assessing the Learning Path

Understand c. arise at each level of hierarchical organization.

d. All of the above
1. Humans and yeast, single-celled organisms with a nucleus, are
both members of 4. The process of inductive reasoning involves
a. Domain Archaea. a. the use of general principles to predict a specific result.
b. Kingdom Animalia. b. the generation of specific predictions based on a belief system.
c. Domain Eukarya. c. the use of specific observations to develop general principles.
d. None of the above d. the use of general principles to support a hypothesis.
2. Which of the following is NOT a property of life? 5. A hypothesis in biology is best described as
a. The use of energy that ultimately comes from the Sun a. a possible explanation of an observation.
b. Movement b. an observation that supports a theory.
c. The maintenance of relatively constant internal conditions c. a general principle that explains some aspect of life.
in a variable environment d. an unchanging statement that correctly predicts some
d. The ability to grow, develop, and reproduce using aspect of life.
instructions found in DNA 6. The idea of evolution by natural selection
3. Emergent properties a. is an example of how a scientist develops a hypothesis
a. result from the way components within a hierarchical level based on previous knowledge and observations.
interact. b. was developed independently by Charles Darwin and
b. explain why our cells are similar to those of a mountain Alfred Russel Wallace.
goat, despite very different body plans. c. is supported by modern-day molecular evidence.
d. All of the above

Chapter 1 The Science of Biology 19

 7. How is the process of natural selection different from that of
artificial selection?
a. Natural selection produces more variation.
b. Natural selection makes an individual better adapted.
c. Artificial selection is a result of human intervention.
d. Artificial selection results in better adaptations.
8. How does the fossil record help support the theory of
evolution by natural selection?
a. It demonstrates that complex organisms have become
simplified and more efficient over time.
b. It provides evidence that organisms have changed over time.
c. It shows that diversity existed millions of years ago.
d. It doesn’t support the theory of evolution, as no
transitional forms have been identified.
9. When we study biological systems, it is clear that
a. they operate by unique laws of chemistry.
b. their unique chemistry allows them to violate chemical
principles.
c. they operate by the laws of chemistry and physics.
d. they are made of unique forms of matter.
10. Your DNA
a. is identical to the DNA from organisms classified in other
kingdoms of life.
b. provides the instructions for your cells to function.
c. can be replicated faithfully and passed to your offspring.
d. Both b and c
Apply
1. Protists and bacteria are grouped into different domains
because
a. protists are multicellular organisms and bacteria are
unicellular organisms.
b. protists are composed of cells and bacteria are not.
c. protists have a nucleus and bacteria do not.
d. All of the above
2. A group of giant pandas inhabit a bamboo forest in China.
The male members of the group have testes; testes are
organs that are part of the reproductive system. The testes
are composed of several cell types, including Leydig cells,
which secrete testosterone molecules. Which of the following
is a correct representation of the hierarchy of biological
organization from least complex to most complex?
a. Testosterone, Leydig cells, testes, reproductive system,
panda, panda population
b. Forest community, panda population, panda, reproductive
system, testes, Leydig cells, testosterone
c. Testosterone, testes, Leydig cells, panda, reproductive
system, panda community
d. Reproductive system, testosterone, Leydig cells, testes,
panda, panda population
3. Birds are vertebrate animals that have feathers and reproduce
by laying eggs. While on a nature walk, you come across a
vertebrate animal that lays eggs but does not have feathers. You
conclude that this animal is not a bird. This is an example of
a. inductive reasoning. c. descriptive science.
b. deductive reasoning. d. reductionism.
4. You are conducting an experiment to test the hypothesis that
apple trees will produce larger fruit when exposed to lullabies at
night. You have 10 experimental trees that are exposed to music at
night. Which of the following would make the best control group?
a. The control group should be exposed to lullabies during
the day.
b. The control group should not be exposed to lullabies at all.
20 Part I The Molecular Basis of Life
c. The control group should be exposed to a different type of
music, such as rap.
d. The control group should be exposed to music all the time.
5. In which of the following situations could evolution by natural
selection occur?
a. A population of island finches possesses almost no genetic
variability.
b. Over several generations a population of snails has access
to unlimited resources.
c. Over generations almost every member of a population of
howler monkeys is able to produce offspring.
d. Over generations a population of mountain gorillas with
genes for thick body hair survives cold and cloudy weather
better than gorillas that have genes for less body hair.
6. Ubiquitin is a small protein (76 amino acids); it has been found
in almost all tissues of eukaryotic organisms and is involved
in protein degradation in the cell. The amino acid sequence of
ubiquitin found in humans has one amino acid difference from
the ubiquitin found in Caenorhabditis elegans (roundworm)
and three amino acid differences from the ubiquitin found
in Saccharomyces cerevisiae (brewer’s yeast). Based on this
information, which of the following statements is ACCURATE?
a. Humans are more closely related to yeast than they are to
roundworms.
b. Yeast, roundworms, and humans all share a common ancestor.
c. These data tell us nothing about evolutionary history,
because molecular evidence relies only on DNA.
d. Molecular data can tell us little about evolution, because it is
usually incompatible with data derived from the fossil record.
7. Aquaporins are water channel proteins that have been found
in membranes of prokaryotes, protists, fungi, plants, and
animals. These proteins have very similar structure in all these
organisms, and they function to allow water to move into and
out of cells. Based on this information, which of the following
statements are ACCURATE? (Select all that apply.)
a. All of these organisms are related.
b. Any alterations in the structure of aquaporins would not
affect their function in transporting water.
c. Water transport is important to life.
d. The DNA that contains the hereditary information for
aquaporins must be conserved.
Synthesize
1. Scientists have recently estimated that there are 1021 stars in
our universe. If 1 in 10 of the stars that are like our Sun has
planets, if 1 in 10,000 of these planets is capable of supporting
life, and if 1 of each million life-supporting planets evolves an
intelligent life-form, how many planets in the universe support
intelligent life? Can you think of a reasonable hypothesis to
explain why we haven’t heard from anyone out there?
2. Based on Apply question 2, which level of hierarchical
organization is the lowest level that carries out all of the
activities we associate with life? Explain your answer.
3. The Cape Verde Islands are as far from the coast of Africa as
the Galápagos Islands are from the coast of South America,
and both island groups have similar climates. Do you think
Darwin would have found animals on the Cape Verde Islands
to be similar to those on the Galápagos? Explain.
4. Exobiology is the study of life on other planets. In recent years
scientists have sent various spacecraft into the galaxy in search
of extraterrestrial life. Assuming that all life shares common
properties, what should exobiologists be looking for as they
explore other worlds?
Another random document with
no related content on Scribd:
juste perspective toute l’antiquité classique si bien dépassée et en
même temps conservée par l’esprit catholique. J’ai lu que Saint
Jérôme s’accusait de lire avec trop de plaisir les auteurs païens.
Ainsi le plus beau du passé revenait dans la pensée chrétienne, et
l’humanisait. C’est un beau et juste mouvement qui ramène ainsi nos
idées à l’enfance, et les mûrit de nouveau dans la moindre de nos
méditations.
Je dirais d’après cela que d’un côté ce sont les vues plus
étendues de l’histoire qui mettent en place l’antiquité classique et la
révolution chrétienne ; car ce grand drame humain est lui-même petit
dans l’immense suite du progrès. Mais, d’un autre côté, ces
antécédents proches font vivre les autres par une reconnaissance
qui soutient les différences, et nous fait historiens. Un Chinois est
trop différent de nous peut-être ; de même un fétichiste d’Amérique
ou de Polynésie. Tacite est un autre genre de sauvage, bien plus
près de nous ; nous ne pouvons méconnaître notre frère ; et c’est le
beau d’abord qui nous en préserve, le beau, bien plus puissant que
le vrai. Sans ce passage, qui d’Homère porté déjà par Platon, nous
conduit tout près de nos naïfs ancêtres et congénères, nous
pourrions bien connaître les Égyptiens, Chinois et Peaux-Rouges
comme on connaît les mœurs des fourmis, tout en restant inhumains
peut-être. Je ne sais rien des Jésuites que par ouï-dire ; peut-être
eussent-ils été plus fanatiques encore sans ce fort préjugé, fondé
sur le jugement esthétique, et qui les tirait hors du temps présent.
Par quoi ils formaient des esprits libres, bien contre leur intention.
Gardons-nous, au rebours, de former des esprits esclaves, bien
contre notre intention.
 VI
HUMANITÉS

Il n’y a point d’Humanités modernes, par la même raison qui fait

que coopération n’est pas société. Il faut que le passé éclaire le
présent, sans quoi nos contemporains sont à nos yeux des animaux
énigmatiques. Ils le sont pour nous, si nous manquons d’études ; ils
le sont en eux-mêmes, s’ils manquent d’études. L’homme qui
invente le téléphone sans fil n’est qu’un animal ingénieux ; ce qu’il
peut montrer d’esprit vient d’autre source.
J’ai observé un certain genre d’incrédulité qui ne suffit à rien. Les
dogmes de l’Église sont à première vue indémontrables et même
absurdes. Soit donc, et laissons-les. Mais celui qui regarde dans les
perspectives du temps aperçoit beaucoup d’autres dieux, d’autres
cérémonies, et des temples qui parlent humainement. Chaîne
d’énigmes qui détourne de s’ébahir parce qu’un polytechnicien va à
la messe. Les hommes ont suivi bien d’autres messes. Mais il faut
s’approcher ; il faut connaître un peu plus intimement le peuple du
Droit, qui est le Romain, et le peuple Sophiste, qui est le Grec ; sans
négliger le peuple adorant, qui est le Juif. Ici un sublime sauvage et
impossible ; ici, par une crainte sans mesure, les superstitions de la
main et du pied, du couteau de table et du pot à beurre. Dans les
deux autres peuples, si proches de nous aussi, mais par d’autres
côtés, des dieux en tout bois et sur toute colline, des oracles, des
augures et des haruspices. L’Égypte et l’Assyrie, incompréhensibles,
forment le fond lointain. L’Orient rêve encore derrière, et le
Polynésien danse. On ignorerait tout de l’homme si l’on n’avait, par
bonheur, familiarité avec les Juifs, avec les Grecs, avec les
Romains, qui ont tant avancé en diverses parties de la sagesse,
gardant avec cela d’étonnantes erreurs. Celui qui ignore cela est
sauvage encore, par une incrédulité mal assise ; dont Montaigne
nous peut guérir ; mais il nous renvoie aux anciens ; il y faut aller. Ou
bien considérer Pascal comme une sorte de fou, et même
Descartes, qui pélerina à Lorette. Ainsi le Moderne, j’entends sans
culture rétrospective, ne voit que fous ; mais je l’attends au
spiritisme, à la théosophie, à tous ces fruits de l’étonnement ; car ce
sont des moments dépassés ; mais il faut les avoir dépassés et
surmontés par une sorte de jeu. Les études classiques assurent le
pied sur cette planète ; l’homme s’y étudie à croire sans se jeter. Nos
folles guerres viennent certainement de trop croire, comme il arrive à
ceux qui n’ont rien vu.
Polynésiens téléphonant ; cela ne fait pas un homme. D’où ces
autels sanglants, et sans dieu. Mais tous les autels furent sanglants
et sans dieu. On ne remarque pas assez que l’humaniste, déjà avec
rosa la rose, se lave les mains de ce sang mêlé à l’eau de la source
Bandusienne. Les Bacchantes retournent à la frise de marbre.
Poésie guérit de frénésie. Les surprises du cœur sont disciplinées ;
un dieu balance l’autre. Le galop des Centaures ne jette plus dans la
charge panique. Déjà Socrate et Phèdre, leurs pieds nus dans l’eau,
s’amusaient à l’entendre. Ce sont nos travaux d’Hercule, et nos
voyages d’esprit, par quoi nous effaçons sur la médaille humaine ce
pli de fanatisme bas. D’où mûrira l’enthousiasme qui ne tue point.
Jaurès modèle. Modèle de tous, et du forgeron encore mieux ; car
toute force est redoutable, et à elle-même aussi. Les Belles-Lettres
donc pour tous ? Et pourquoi non ? Regardons cette idée en face.
 VII
DE LA THÉOLOGIE

Il y a quelque chose de mort dans toute Théologie, quelque

chose de mort aussi dans toute Géométrie. Ce sont des idées sous
clef ; nul n’y va plus voir, et l’on en fait le compte par des registres et
abrégés, comme font les teneurs de livres. Or ces provisions d’esprit
se corrompent encore plus vite que les provisions de bouche. Et
qu’est-ce qu’une idée à laquelle on ne pense point ? Bossuet prouve
Dieu par les vérités éternelles. « Une vérité ne peut cesser d’être
vérité. Descartes meurt, Bossuet meurt, la vérité ne meurt point.
Mais comme une vérité n’est rien aussi sans quelque pensant, il
existe donc un Pensant éternel ». Voilà une pensée de disciple et
une armoire aux idées. Descartes est bien plus difficile à suivre,
parce qu’il brise l’armoire aux idées et les idées mêmes, allant
jusqu’à dire qu’il n’y a point du tout de vérités éternelles et que la
volonté de Dieu en décide à chaque instant, même du triangle et du
cercle. Comprenne qui pourra. Toujours est-il qu’il y a ici du scandale
et une occasion de douter de l’indubitable, par quoi la théologie de
Descartes se trouve animée d’incrédulité. Au feu les idoles. Ainsi va
le vrai Géomètre, toujours doutant et défaisant, d’où les idées
naissent et renaissent. Car je tiens que si l’on veut savoir ce que
c’est qu’une ligne droite il faut y penser toujours, j’entends la vouloir
et maintenir toujours, ce qui est douter et croire ensemble. Quant à
la ligne droite qui tient d’elle-même, et qui est enfermée en quelque
Palais des Mesures, je sais qu’elle n’est point droite. Rien au monde
n’est droit.
Rien au monde n’est juste. Aucun objet n’est Dieu. Mais l’homme
juste est celui qui pense toujours au juste, et continuellement le
maintient et le veut, imitant le Dieu de Descartes en cette création
continuée. C’est ainsi que le juste fait justice de tout, comme le
géomètre fait géométrie de tout. Un tel homme ne se fie point à
l’ordre des choses, et la pointe de son jugement toujours attaque la
justice établie et vénérée, la redressant d’après le modèle qui
n’existe pas. Ce feu du jugement moral, cette ardeur à briser, ce
culte du Dieu seulement aimé, nu, et sans aucune puissance, voilà
par où la religion vit et revit. Plus religion dans ce Socialiste que
dans ce Thomiste. Mais il se peut bien que le socialisme soit
théologique maintenant, et que la Justice soit maintenant sous clef
dans quelque Pavillon des Justes Mesures. L’idée aura donc péri par
la Suffisance.
On doit appeler machine, dans le sens le plus étendu, toute idée
sans penseur. Je remarque que la téléphonie sans fil guérit de
comprendre et même d’essayer de comprendre. Et l’avion a tué
l’idée de l’avion, comme les ailes, en l’oiseau, ont tué le doute, âme
des formules de Newton et d’Euler. Car qui pensera, si tout est
pensé ? Qui réglera, si tout est réglé ? La violence est l’effet
inévitable, et souvent prochain, d’une pensée sans aucun doute ; et
c’est ce que l’on voit en gros chez les fous. Peut-être est-il dans la
destinée de toute théologie, aussitôt achevée, de rouler sur la terre
comme un char d’assaut, C’est ainsi que la puissance déshonore la
justice.
 VIII
DE L’ART DE PERSUADER

Pensant à Joseph de Maistre, je faisais une revue en moi-même

des hommes qui ont fait serment de croire ; et au premier rang
j’apercevais Socrate, tel que Platon le représente en son Gorgias, ou
bien dans sa République, faisant de la tête signe que non, à chaque
fois que les disputeurs l’accablent de leurs preuves d’expérience ; et,
comme dit justement Socrate, il n’est pas difficile de faire voir que la
force gouverne partout et que la justice est ce qui plaît aux plus
forts ; c’est le spectacle humain ; on n’entend que cela ; on ne voit
que cela. Suivez ces longues discussions en leurs détours, vous
verrez apparaître la justice, et soudain disparaître. On la saisit à la
fin ; il vient un moment heureux où toutes les parties de la nature
humaine sont rassemblées et comme pacifiées selon la loi interne
de justice, à laquelle les manifestations externes de la force sont de
loin subordonnées. Tout s’ordonne alors, et la vraie punition répond
à la vraie récompense. Mais, pour parvenir à cette vue, il faut autant
de patience au moins qu’en montre Socrate. Un lecteur pressé verra
partout l’injustice revenant toujours à la suite de la puissance, et la
justice autant démunie de preuves que de richesses. En quoi il n’y a
point de jeu ni d’artifice, mais au contraire la plus parfaite peinture de
ces tâtonnements et détours de pensée qui rebutent promptement
celui qui n’a pas juré. Il faut jurer d’abord, et dire non aux arguments
diaboliques avant de savoir comment on y répondra.
 Autre chose encore, et qui irrite toujours un peu. Vous lisez ; vous
pesez au passage les preuves Socratiques ; vous les rassemblez ;
vous saisissez l’idée ; vous la confiez comme un trésor au coffret de
la mémoire. Mais le diable guette encore par là. Quand vous ouvrez
le coffret de nouveau, vous ne trouvez plus qu’une pincée de
cendres ; éléments dissous et dispersés ; chaos. Il faut tout refaire ; il
faut s’aider de nouveau de l’art socratique ; de nouveau l’injustice est
brillante et forte ; de nouveau la clameur diabolique assourdit le
pauvre homme ; il faut passer par ce chemin-là. Si le courage
manque, tout est dit. C’est pourquoi on voit trébucher tant de
penseurs vieillissants, et s’asseoir au festin de la Force, où l’on boit
l’hydromel dans le crâne de l’ennemi. J’ai vu un noble penseur se
lever et marcher à grands pas, allant et revenant, et disant à moi :
« On devrait savoir une bonne fois. Quand on a passé le lieu difficile,
on devrait le laisser derrière soi pour toujours. Et quand on a formé
l’idée, on devrait la posséder. Tout sera donc toujours à
recommencer » ? C’est ce que Socrate demandait en ces termes
mêmes. En tout, on veut une charte ou un diplôme, et dormir
dessus. Mais ce n’est point permis.
D’après ces rudes expériences, il faut comprendre ces préjugés
invincibles et volontaires que l’on rencontre en tout homme un peu
composé, et qui rendent si pénible le travail de prouver et de
convaincre. Combat difficile, où les meilleurs coups, les plus
savants, les mieux dirigés, sont justement ceux qui ébranlent le
moins l’adversaire. J’ai observé comment un esprit vigoureux
esquive la preuve forte, et qu’il voit venir de loin, refusant attention à
ce que vous dites, non parce qu’il le juge faible, mais parce qu’il le
juge fort, et récitant en lui-même son serment de fidélité comme une
prière. L’homme est beau alors. Car, si difficile que soit notre
condition de pensant, songez qu’elle serait tout à fait misérable, si
nous devions abandonner une idée précieuse, et bien des fois
éprouvée, dès que nous n’avons rien à répondre à quelque
disputeur. Dans le fait, personne ne pense ainsi. Tout homme
pensant s’appuie sur une foi invincible ; c’est son réduit et donjon.
D’où je tire la règle des règles, qui est de ne point penser contre
l’autre, mais avec l’autre, et de prendre sa profonde et chère
pensée, autant que je la devine, comme humaine et mienne. Et
quand cette pensée est la Justice éternelle, qu’on l’appelle Dieu ou
comme on voudra, on peut s’y établir, et travailler en partant de là ;
prises de ce côté-là, les murailles tomberont.
 IX
PROPHÉTIES

En aucun temps je ne me fiai aux coteaux de l’Aisne comme en

l’été de l’an quatorze. Je me trouvais réconcilié à cette terre,
sauvage un peu par ses roches, et par les noms sinistres qui
rappelaient les guerres de l’autre siècle. De précieux amis
vieillissaient là. Vingt marches de pierre me conduisaient à leur
jardin fleuri ; vingt marches encore, et l’on était au paisible jardin des
morts, fleuri de marjolaine et d’hysope, autour de l’église paysanne.
De ce promontoire la vue s’étendait presque jusqu’à Soissons, par
une trouée fameuse. Sur le plateau à blé, presque à la hauteur du
coq indicateur des vents, passait la Route des Dames au nom
charmant. La falaise était riche de sureaux et de vignes et portait,
sur ses pentes arides, des genévriers et un rosier sauvage à odeur
musquée que je n’ai vu que là. On pouvait s’y plaire ; et, en ce mois
de Juillet, j’achevais une clôture durable autour d’une maison de
tisserand.
C’est là que j’entendis deux prophéties. Une première fois moi-
même je vaticinai, je ne sais pourquoi, en compagnie d’un
philosophe paysan, que l’on jugeait un peu fou. Cette sécurité des
travaux, dont l’image s’offrait partout, me parut d’un moment, comme
elle était. Il ne faut qu’une peste, disais-je, ou une querelle entre les
hommes, pour que cette sauvage écorce de la terre, que l’on voit par
places, recouvre le coteau, le plateau, la vallée, et les collines
éparses semblables à des îles. Le soir, qui effaçait les différences, et
la vue aussi de mon mélancolique compagnon, me faisaient penser
à ces choses ; mais il me semble maintenant que je déclamai un peu
plus que l’état présent ne le conseillait. L’avenir d’alors, maintenant
passé, donne trop de sens à ces paroles de hasard.
Un autre jour ce fut une sorte de sorcière qui prophétisa, courbée
en deux par les travaux, levant vers moi son regard bleu et son
visage couleur de brique. Elle me montrait, dans le jardin et dans les
vergers en terrasse, une quantité étonnante de taupinières, et elle
parla en ces termes : « Vous savez ce qu’on dit par ici et ce que je
sais ; autant de taupinières, autant de tombes. » Elle redit plusieurs
fois la même chose, en regardant à droite et à gauche, comme elle
avait coutume. Sur quoi je fermai mon imagination comme une
porte, admirant comment la ressemblance fait preuve en ces esprits
trop faibles pour soulever la métaphore. Or il y eut, partout par là,
comme on sait, encore plus de tombes que de taupinières.
De ces rencontres émouvantes, je ne pense rien. Il y a, à toute
minute, des rencontres aussi admirables que celles-là, si l’on voulait
admirer ; et tout est signe dès que l’on cherche des signes. Du
moins je comprends un peu mieux les temps homériques, et ces
présages continuellement tirés des oiseaux, des nuages, de la
foudre ; dont quelques-uns se trouvaient vérifiés par hasard, et
beaucoup réalisés par l’action de ceux-là même qui y croyaient ; car
souvent l’oracle conseille en même temps qu’il annonce ; et ce n’est
pas merveille si la mêlée devient terrible, selon la prédiction, du
moment qu’on y croit. Le monde n’a point changé, et notre sagesse
repose toute sur elle-même. Qui veut croire trouvera des preuves,
d’autant que le souvenir ne retrouve jamais le passé tel qu’il fut,
mais le recouvre de ce qui a suivi, suspendant au présage
l’accomplissement comme une couronne. Les dieux sont les
premiers nés du souvenir.
 X
DES MÉTAPHORES

Je n’approuve point ceux qui veulent changer la pantoufle de

verre, dans Cendrillon, disant que ce n’est point du verre, et qu’il n’y
eut jamais de pantoufles de verre, chose dure et cassante, mais qu’il
s’agit de vair qui est fourrure souple et chaude. Remarquez qu’il y a
bien d’autres choses impossibles, dans les autres contes et dans
celui-là. Mais l’érudit est assez content d’avoir remis une pantoufle
en place ; il attend l’occasion d’expliquer par la même méthode la
citrouille qui devient carrosse, ou cette ronde de petites filles, qui, à
force de tourner, devient motte de beurre. On peut rire du pédant ;
mais il faut quelquefois le prendre au sérieux. C’est la sottise armée.
Je range le Pédant dans la puissante classe des Détourneurs,
dans laquelle on trouve aussi des espèces non dépourvues
d’élégance. Et la chasse du Détourneur est une chasse aux Idées.
Dès qu’une idée s’envole ils la tuent, comme on tue les Idées, en
détournant de les chercher. L’Esprit se jette sur quelque pauvre
relation bien aisée à saisir et à redresser. Il rit de cette victoire facile,
et le Détourneur marque un point.
J’ai dit souvent que tous les contes sont vrais ; mais ce n’est pas
assez dire. La profonde sagesse populaire est plus rusée que nos
philosophes. Et, au rebours du Détourneur, elle nous met en garde
contre cette fausse Raison, qui n’est qu’imagination conforme à la
coutume. Et par un piquant moyen, aussi ancien que l’espèce
humaine, qui est de nous jeter l’absurde aux yeux, de grossir et de
redoubler l’impossible, par quoi l’imagination est définie, en même
temps qu’elle est éveillée, et rappelée à son rôle de Folle. A quoi
servent aussi ces comparaisons étranges que le génie poétique jette
comme un défi. J’admire la grandeur des enfants, qui ne discutent
jamais sur la Lettre. Non qu’ils saisissent d’abord l’Esprit ; mais ils
savent bien que l’Esprit ne vise pas ce maigre gibier. Ainsi en
s’amusant de l’absurde ils ne déshonorent pas l’Esprit, mais au
contraire ils l’honorent. Par la croissance qu’il sent à l’œuvre en lui,
et qui lui donne espoir et patience, ce bel âge voit grand. Il attend
quelque chose de mieux que des fictions cohérentes. Certes il y a de
la Majesté à laisser jouer l’Imagination en même temps que le corps,
et par les mêmes lois. Mais il y a quelque chose d’impérieux aussi à
vouloir que l’absurde soit conservé comme il est ; c’est refuser les
petites raisons. Shakespeare se moque de ceux qui voudraient
comprendre comment Othello ou Hamlet sont passés d’un lieu à
l’autre, invitant ainsi énergiquement le spectateur à comprendre
d’autres vérités, plus cachées et plus difficiles. Sur l’absurde même
l’Esprit rebondit, car il n’y peut rester. Cette apparence ne peut
tromper, il faut donc voir au delà. Ces signes nous délivrent des
signes. Au contraire, par des signes de raisonnable apparence, nous
venons à penser les signes, et la coutume nous tient. Telle est la
vieillesse de l’Esprit.
 XI
DES APPARENCES

Quand un Hindou se marque au visage de signes rouges ou

bleus selon sa caste, vous ne demandez point si cela est vrai ou
faux. Il vaudrait mieux se demander en quel sens et sous quel
rapport cela est vrai ; pour le faux, il n’est point dans les faits ni dans
les actions, ni dans les pensées ; il y a vérité de tout ; il faut
seulement dire que nous sommes bien loin de connaître la vérité de
tout ; mais la privation n’est rien. Attendez les exemples. Il y a une
apparence du mouvement du ciel autour de son axe ; cette
apparence n’est qu’apparence, c’est dire qu’elle n’est point vraie ;
mais je ne dirai pas non plus qu’elle est fausse ; car, placés comme
nous sommes sur cette terre qui tourne, nous ne pouvons la voir
tourner. Pour mieux dire, je ne vois aucune chose comme elle est.
Je vois à l’angle de mon plafond trois angles joints dont je sais qu’ils
sont d’équerre tous les trois, mais je les vois obtus tous les trois, et
la perspective m’apprend que je dois les voir ainsi. Si je change de
place, je les verrai obtus autrement ; mais ce sont toujours trois
angles droits. Ceux qui retournent dans leur tête les paradoxes
d’Einstein croient souvent qu’ils ont à choisir entre plusieurs
apparences du temps et un temps unique ; je les invite à réfléchir sur
l’objet unique, qui donne pourtant d’innombrables perspectives. Je
dirai volontiers que cet Hindou qui se peint le visage se règle sur
quelque perspective de l’existence physiologique et politique ; et,
autant que je connais le vrai de la chose, il faut que je comprenne
cette perspective qui est sienne ; et aussi bien cette perspective d’un
autre qui se fait moine, et de moi-même aussi qui mets une cravate.
Si vous me proposez une religion, je l’examine, non point avec
l’idée qu’elle est fausse, mais au contraire avec l’idée qu’elle est
vraie. D’où vient donc que je passerai pour irréligieux ? C’est que je
pense la même chose de toutes les religions. Chacune d’elles n’est
qu’une perspective plus ou moins déformée dans laquelle il faut que
je retrouve l’objet unique. Travail copernicien. Difficile assurément,
mais considérez ce qui arrive quand on me montre des tours de
passe-passe ou des jeux de miroirs. Ce sont alors des apparences
étranges ; mais je sais sans le moindre doute que si je connaissais
bien les objets dont ces apparences sont les apparences, je ne
verrais plus rien d’étrange dans ce spectacle.
Lorsque Galilée disait que la terre tourne, c’était parce qu’il avait
deviné le secret d’une apparence, et vu, en quelque sorte, le double
fond de la boîte. Ainsi, bien loin qu’il pensât que les autres se
trompaient, au contraire, il comprenait leur erreur même comme
vérité, et se trouvait ainsi plus assuré de ce qu’ils disaient qu’eux-
mêmes. Mais eux voulaient le ramener aux apparences, et lui faire
jurer qu’il voyait les apparences. Aussi lui, qui voyait le soleil tourner,
comme voit n’importe quel astronome, ne trouva sans doute point
autant de difficulté qu’on voudrait croire à dire comme ils disaient ; et
peut-être comprit-il aussi le vrai de leur colère, et l’éternel objet
politique sous ces menaçantes apparences. Marc-Aurèle a dit là-
dessus le dernier mot peut-être : « Instruis-les, si tu peux ; si tu ne
peux les instruire, supporte-les. » Quand le roi David chante :
« L’Éternel est mon rocher », je lui donne raison, mais non pas
comme il voudrait : on peut parier qu’avant la fin de mon discours il
m’aurait fait pendre. Il faut être bien intolérant pour se laisser
pendre.
 XII
SCIENCE ET RELIGION

A ceux qui voudraient dire que l’homme est arrivé à la sagesse

par la prudence, je conte souvent ce que j’ai lu dans les journaux au
lendemain d’un attentat politique. Deux bombes avaient été lancées ;
une seule éclata ; l’autre fut portée au commissaire peut-être deux
jours après par un homme qui l’avait trouvée et mise dans sa poche.
J’ai supposé d’abord que cet homme téméraire n’arrivait pas à croire
que ce morceau de fonte en forme de pomme de pin pouvait éclater
au moindre choc et mettre un corps vivant en charpie ; et il faut bien
supposer cela, mais il ne faut pas expliquer cette action seulement
par l’ignorance ; un chimiste peut bien être téméraire aussi.
J’aimerais mieux dire que l’homme en solitude, et attentif surtout à
ses actions, n’arrive jamais à imaginer un événement redoutable. Et,
à bien regarder, un corps sain, vigoureux et intact ne peut point
témoigner du tout, par ses affections, que la griffe du lion peut le
déchirer, mais au contraire il exclut naturellement une telle image.
Ainsi le danger serait trop tard connu toujours, et l’expérience
n’instruirait guère.
Chose digne de remarque, la cérémonie modifie bien plus
énergiquement les sentiments de chacun. Cela vient de ce que,
dans la cérémonie, les actions sont prévues et faciles, et que notre
corps est principalement occupé à imiter les mouvements d’autrui ;
nous sommes tous alors des tragédiens de bonne foi. Par exemple
la peur nous est alors directement communiquée, à la manière d’une
maladie ; l’objet quel qu’il soit, même absent ou invisible, reçoit de
cette peur une puissance sans mesure. On peut comprendre ainsi
que des hommes très résolus en leurs actions soient comme des
enfants lorsqu’ils pensent aux revenants, aux lutins, aux diables,
quoiqu’ils ne les connaissent que par des récits. Un récit est bien
plus puissant que la chose même, par l’effervescence commune au
récitant et à la foule des auditeurs.
J’irais jusqu’à dire que les choses ne savent point toucher l’esprit
de l’homme ; elles ne passent point jusque-là ; cet animal a l’esprit
cuirassé contre tout expérience. Mais le cri d’un enfant, le soupir
d’une femme, le léger vent d’un geste humain, tous les signes
entrent librement dans la citadelle. Encore mieux les armées de
signes qui s’envolent d’une assemblée. D’où il arrive que l’homme
croit moins ce qu’il a vu que ce qu’on lui raconte, ce qui éclaire
toutes nos passions. Mais je veux retenir seulement ceci, que
l’histoire des Sciences expose sans l’expliquer, que l’homme est
venu à la prudence par le respect, et à la Science par la Religion ;
autrement dit que nos premières connaissances, et les seules que
nous prenions naturellement au sérieux, sont de ouï-dire, et non
d’expérience. La crainte de Dieu serait donc la première des
craintes, et le modèle de toutes.
 XIII
LE TEMPLE

Une des idées de l’Eupalinos est que le temple meut l’homme ;

par quoi l’architecture ressemble à la musique. Mais il faut voir
comment le temple meut l’homme. Par ceci que les grands reliefs
s’aplatissent dès que l’on s’arrête, comme si l’air des profondeurs en
était chassé. Au contraire, dès que le spectateur se met en
mouvement, si peu que ce soit, le temple aussitôt déplace ses
perspectives, et d’autant plus que les parties en apparence
juxtaposées sont réellement plus éloignées les unes des autres ;
mais c’est ce qu’il faut essayer, car on ne le croit jamais assez, en se
déplaçant seulement d’un pas devant l’entrecoupement des arceaux
et des flèches, ou devant les éclipses des colonnades. C’est
explorer la profondeur, et se rendre sensible la solidité de la chose.
C’est donc la mort de la chose dans son apparence qu’il faut vaincre
par le mouvement. Ainsi le monument nous appelle ; mais d’une
certaine manière, selon sa structure. Et Hegel a bien su dire que la
cathédrale gothique forme le plus énergique appel, par l’opposition
du dehors et du dedans, par l’énigme des contreforts et par la
promesse des portes et cette foule pressée des statues qui nous
font un chemin. Une colonnade grecque nous meut d’autre façon.
Ainsi la Madeleine serait plus politique que Notre-Dame. Mais de
toute façon il faut se mouvoir ; et c’est par le mouvement que le
monument nous fait penser.
 La peinture au contraire nous tient immobile. Supposons un
monument peint dans le fond d’un tableau ; cette image ne répond
point à nos mouvements, et l’on n’observe point ce glissement des
colonnes qui se montrent et se cachent selon nos pas ; ce qui
rabaisse aussitôt le monument peint au niveau des accessoires ; ce
qui, surtout, nous avertit que cette apparence doit rester à l’état
d’apparence, et qu’ici le vrai est de nous et non de l’objet. Il est donc
profondément vrai que l’architecture et la peinture ne sont point du
même âge. Car le puissant objet de pierre nous tire à une pensée
commune de manière à effacer toute méditation de soi sur soi ; il
nous soumet à la doctrine. Mais la peinture au contraire, par cette
apparence désormais fixée, nie l’être et divinise l’existence. Le
miracle de la peinture c’est de donner être à ce qui passe ; c’est
pourquoi tout peut plaire en peinture, un arbre, un nuage, un reflet.
Par quoi nous voilà immobiles, en quelque sorte, à la seconde
puissance ; car nous savons bien que le mouvement est un adieu à
l’apparence et à soi, mais plutôt une sorte d’hymne à ce monde
solide, et un massacre des apparences.
Un arc de triomphe est peut-être l’objet le plus éloquent qui soit.
Ce n’est pourtant qu’une porte de ville, et séparée des murs. Le petit
arc du Carrousel est posé sur cette place comme un signe ; on voit
d’autres choses par l’ouverture ; ce n’est qu’un passage ; mais il faut
passer. Entrer et sortir, ce n’est qu’un. Ainsi s’éveille le pas militaire ;
ainsi, par cette porte qui n’est que porte, l’entreprise qui ne promet
rien, ce qui donne vie au grenadier de pierre. Il attend de partir. Et
au-dessous, les ombres sur le sable stérile font oubli et désert. Car,
par la vertu de ce monument, qui signifie absolument l’en-dehors de
soi, il y faut passer, mais on n’y peut rester.
 XIV
IDOLES

L’esprit dans la chose, voilà le dieu. Une horloge en ses rouages

et accrochages me raconte l’idée de l’horloger ; mais il n’y a point de
merveilleux là-dedans ; chaque roue ne dit qu’une chose. Au lieu
que la Joconde en dit bien plus que le peintre ne savait. Une belle
statue signifie sans fin ; les arceaux d’un cloître ont des milliers
d’aspects, tous parents de nous-même. Un quatuor de Beethoven
prend plus de sens d’année en année. Toutes ces œuvres, outre
l’immense pensée qui leur est propre, et qui nous dépasse toujours,
renvoient aussi tout ce culte et tous ces hommages qu’elles ont
reçus, comme ces autels plus vénérables par les couronnes. Le
temps n’épuisera point cet avenir de gloire. J’ai lu l’Iliade une fois de
plus ; c’est comme si j’avais apporté encore une pierre à ce grand
tombeau.
Quand le sauvage eut ébauché des tronçons basaltiques selon la
forme humaine, il ne put juger son œuvre ; mais au contraire c’est lui
qui fut jugé. Ces yeux de pierre furent plus forts que lui. Cette
immobile armée le tint en respect mieux qu’un despote ; car un
despote change d’attitude et de lieu et désire enfin quelque chose ;
mais les statues n’ont pas besoin de nous, ni de rien. Ainsi la statue
fut un dieu. Je dois appeler prière cette méditation devant le signe,
cette offrande qui est due, et dont le dieu n’a pas besoin, ce muet
dialogue où, d’un côté, toutes les réponses sont faites d’avance, et
toutes les demandes d’avance devinées. Ainsi la pensée sait où elle
va, et le vrai se montre dans l’immobile.
On voudrait dire que l’homme a fait des idoles parce qu’il était
religieux ; c’est comme si l’on disait qu’il a fait des outils parce qu’il
était savant ; mais au contraire la science n’est que l’observation des
outils et du travail par les outils. De même je dirais plutôt que la
première contemplation eut pour objet l’idole, et que l’homme fut
religieux parce qu’il fit des idoles. Il fallait rendre compte de cette
puissance du signe, et inventer la mythologie pour expliquer le beau.
L’Imitation de Jésus-Christ n’est que la traduction abstraite de cette
imitation du signe, qui est cérémonie. La réflexion sur l’idole arrive à
nier l’idole, par les perfections mêmes que l’on y devine ; mais c’est
déjà impiété. L’Iconoclaste doit se trouver sans dieu finalement. De
ce côté est la perfection sans objet ; ce néant nous renvoie à l’idole,
objet alors d’une adoration purifiée ; tel est l’art en notre temps,
moment dépassé et conservé, comme dit Hegel.
Les moyens de ce penseur, qui avance toujours par position,
négation et solution, seraient donc les instruments de l’histoire. Ceux
qui ont méprisé trop vite cette dialectique devraient bien considérer
que Comte, qui la méconnut aussi, est pourtant arrivé à faire
entendre, par d’autres mots, les mêmes relations. Car selon ses
vues, chaque jour mieux vérifiées, l’ancien fétichisme est bien la
religion essentielle, tandis que la religion pensée et purifiée n’est que
la négation de la religion, qui, sous le nom de théologie et de
métaphysique, tire le dieu hors du signe, et même hors du temple,
lui-même signe, et nous jette dans l’infini sans matière, d’où nous
devons aussitôt revenir, C’est alors que, selon l’esprit positif, l’ancien
fétichisme, sous le nom de contemplation esthétique, doit orner
l’existence coopérative, qui est elle-même négation de négation.