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 [f o u r t h e d i t i o n]

Principles of

Biolog y
Robert J. Brooker
University of Minnesota–Minneapolis

Eric P. Widmaier
Boston University

Linda E. Graham
University of Wisconsin–Madison

Peter D. Stiling
University of South Florida

bro23659_fm_i-xxviii.indd 1 30/09/22 8:54 PM

 PRINCIPLES OF BIOLOGY

Published by McGraw Hill LLC, 1325 Avenue of the Americas, New York, NY 10019. Copyright ©2024 by McGraw
Hill LLC. All rights reserved. Printed in the United States of America. No part of this publication may be reproduced
or distributed in any form or by any means, or stored in a database or retrieval system, without the prior written
consent of McGraw Hill LLC, including, but not limited to, in any network or other electronic storage or transmission,
or broadcast for distance learning.

Some ancillaries, including electronic and print components, may not be available to customers outside the
United States.

This book is printed on acid-free paper.

1 2 3 4 5 6 7 8 9 LWI 28 27 26 25 24 23

ISBN 978-1-266-13582-8
MHID 1-266-13582-0

Cover Image: Leigh Gregg/Shutterstock

All credits appearing on page or at the end of the book are considered to be an extension of the copyright page.

The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website does not
indicate an endorsement by the authors or McGraw Hill LLC, and McGraw Hill LLC does not guarantee the accuracy
of the information presented at these sites.

mheducation.com/highered

bro35820_fm_ISE_ii.indd 2 23/09/22 4:23 PM

 Brief Table of Contents

1 An Introduction to Biology 1 24 Microbiomes: Microbial Systems on and Around

Us 521
UNIT I Chemistry 20 25 Plant Evolution: How Plant Diversification Changed
Planet Earth 538
2 The Chemical Basis of Life I: Atoms, Molecules, and
Water 21 26 Invertebrates: The Vast Array of Animal Life Without a
Backbone 562
3 The Chemical Basis of Life II: Organic Molecules 37
27 Vertebrates: Fishes, Amphibians, Reptiles, and
UNIT II Cells 58 Mammals 596

4 Evolutionary Origin of Cells and Their General UNIT VI Flowering Plants 618
Features 59
5 Membranes: The Interface Between Cells and Their 28 An Introduction to Flowering Plant Form and
Environment 98 Function 619
6 How Cells Utilize Energy 121 29 How Flowering Plants Sense and Interact with Their
Environments 640
7 How Cells Capture Light Energy via Photosynthesis 150
30 How Flowering Plants Obtain and Transport Water,
8 How Cells Communicate with Each Other and with the Mineral Nutrients, and Organic Compounds 658
Environment 169
31 How Flowering Plants Reproduce and Develop 682
UNIT III Genetics 188 UNIT VII Animals 700
9 The Information of Life: DNA and RNA Structure, DNA
32 General Features of Animal Bodies, and Homeostasis as
Replication, and Chromosome Structure 189
a Key Principle of Animal Biology 701
10 The Expression of Genetic Information via Genes I:
33 Neuroscience I: The Structure, Function, and Evolution
Transcription and Translation 211
of Nervous Systems 722
11 The Expression of Genetic Information via Genes II:
34 Neuroscience II: How Sensory Systems Allow Animals
Non-coding RNAs 231
to Interact with the Environment 752
12 The Control of Genetic Information via Gene
35 How Muscles and Skeletons Are Adaptations for
Regulation 250
Movement, Support, and Protection 774
13 Altering the Genetic Material: Mutation, DNA Repair,
36 Circulatory and Respiratory Systems: Transporting
and Cancer 271
Solutes and Exchanging Gases 789
14 How Eukaryotic Cells Sort and Transmit Chromosomes:
37 Digestive and Excretory Systems: Maintaining Nutrient,
Mitosis and Meiosis 289
Water, and Energy Balance and Removing Waste 822
15 Transmission of Genetic Information from Parents to
38 How Endocrine Systems Influence the Activities of All
Offspring I: Patterns That Follow Mendel’s Laws 313
Other Organ Systems 857
16 Transmission of Genetic Information from Parents to
39 The Production of Offspring: Reproduction and
Offspring II: Epigenetics, Linkage, and Extranuclear
Inheritance 331 Development 879
17 The Simpler Genetic Systems of Viruses, Bacteria, and 40 Immune Systems: How Animals Defend Against
Archaea 351 Pathogens and Other Dangers 905
18 Genetic Technologies: How Biologists Study Genes and 41 An Example of a System-Wide Response to a Challenge
Genomes 370 to Homeostasis 927

UNIT IV Evolution 396 UNIT VIII Ecology 942

19 Evolution of Life I: How Populations Change from 42 Behavioral Ecology: The Struggle to Find Food and
Generation to Generation 397 Mates and to Pass on Genes 943
20 Evolution of Life II: The Emergence of New Species 426 43 Population Growth and Species Interactions 960
21 How Biologists Classify Species and Study Their 44 Communities and Ecosystems: Ecological Organization
Evolutionary Relationships 445 at Large Scales 980
22 The History of Life on Earth and Human Evolution 463 45 Biomes: How Climate Affects the Distribution of Species
on Earth 1003
UNIT V Diversity 488 46 The Age of Humans 1017
47 Biodiversity and Conservation Biology 1043
23 Diversity of Microbial Life: Archaea, Bacteria, Protists,
and Fungi 489

Brief Table of Contents iii

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 About the Authors

Robert J. Brooker
Rob Brooker received his Ph.D. in genetics from Yale University in
1983. At Harvard, he studied lactose permease, the product of the lacY
gene of the lac operon. He continued working on transporters at the
University of Minnesota, where he is a professor in the Department
of Genetics, Cell Biology, and Development, and in the Department
of Biology Teaching and Learning. At the University of Minnesota,
Dr. Brooker teaches undergraduate courses in biology and genetics.
In addition to many other publications, he has written two undergrad-
uate genetics texts: Genetics: Analysis & Principles, seventh edition,
copyright 2021, and Concepts of Genetics, fourth edition, copyright
2022; and he is the lead author of Biology, sixth edition, copyright
2023, all published by McGraw Hill Education.

Eric P. Widmaier
Eric Widmaier received his Ph.D. in 1984 in endocrinology from the
University of California at San Francisco. His research focused on the
control of body mass and metabolism in mammals, the hormonal cor-
relates of obesity, and the effects of high-fat diets on intestinal cell
function. Dr. Widmaier is currently professor emeritus of biology at
Boston University, where he has taught undergraduate and graduate
courses in human physiology, comparative physiology, and endocrinol-
ogy and received the university’s highest honor for excellence in teach-
ing. Among other publications, he is lead author of Vander’s Human
Physiology: The Mechanisms of Body Function, sixteenth edition,
copyright 2023, and coauthor of Biology, sixth edition, copyright 2023,
both published by McGraw Hill Education.
Linda E. Graham
Linda Graham earned an undergraduate degree from Washington
University (St. Louis), a master’s degree from the University of
Texas, and a Ph.D. from the University of Michigan, Ann Arbor,
where she also did postdoctoral research. She is a past department
chair of the Department of Botany at the University of Wisconsin−
Madison, where her research explores the biology of algae and plants,
particularly their evolution and microbial associations. She is an
AAAS Fellow, past President of the Botanical Society of America,
and past President of the Phycological Society of America. She has
taught undergraduate courses in general biology, microbiology, and
Left to right: Peter Stiling, Linda Graham, Eric Widmaier, and Rob Brooker
©Ian Townsend
plant biology. She is the lead author of Algae, third edition, copy-
right 2016, a textbook on algal biology, and Plant Biology, third edi-
tion, copyright 2015, both published by LJLM Press, and coauthor
of Biology, sixth edition, copyright 2023, published by McGraw Hill
Education.
Peter D. Stiling
Peter Stiling obtained his Ph.D. from University College, Cardiff,
United Kingdom. Subsequently, he became a postdoctoral fellow
at Florida State University and later spent two years as a lecturer at
the University of the West Indies, Trinidad. Dr. Stiling, Professor of
Biology in the Department of Integrative Biology at the University of
South Florida (USF) at Tampa, was formerly Chair of the Department
and later Assistant Vice Provost for Strategic Initiatives. His research
interests include plant-animal relationships and invasive species.
He also taught biology to students in the summer program USF in
London, from 2015 to 2019, after establishing the program in 2015.
Dr. Stiling was elected an AAAS Fellow in 2012. He is also the author
of Ecology: Global Insights and Investigations, second edition, copy-
right 2015, and coauthor of Biology, sixth edition, copyright 2023,
both published by McGraw Hill Education.

iv About the Authors

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 Principles of Biology: Helping Students Learn
Critical-Thinking Skills
The fourth edition of Principles of Biology, along with previous editions, has been crafted with a very important goal in mind: Give students
an opportunity to develop critical-thinking skills. Critical thinking is the mental process of actively and skillfully conceptualizing, applying,
analyzing, synthesizing, and/or evaluating information gathered from, or generated by, various experiences including reading, observation,
experimentation, reflection, and/or oral communication. It provides a guide to belief and action. A person who is skillful at critical thinking can
draw reasonable conclusions from a collection of information and can discriminate between useful and less useful details to solve problems
and to make decisions.
If a primary goal for your students is to understand the principles of biology and to improve their critical-thinking skills, Principles of
Biology, fourth edition, by Brooker, Widmaier, Graham, and Stiling is an excellent choice. We expect that their journey through this textbook
will help your students to think like scientists and to develop skills that are needed in many different careers in biology.

EMPHASIZING SKILLS DEVELOPMENT AND PROBLEM SOLVING

Skills! Skills! Skills! That’s what instructors want their students to activation energy for a chemical reaction and thereby increases its
develop and that’s what students often find the most challenging. The rate, the author team has developed pedagogical features that should
pedagogical features of Principles of Biology, fourth edition, are aimed lower the barriers that prevent skill development and thereby allow
at achieving this goal. Much like the way that an enzyme lowers the students to develop their critical-thinking skills more quickly.

Critical-Thinking Questions at the End of Each Chapter

In the fourth edition of Principles of Biology, the end-of-chapter questions have a category of questions called Critical-Thinking Skills.
These questions are primarily at Bloom’s levels 3 (applying) and 4 (analyzing).

Assessing Your Knowledge and Skills

Foundational Knowledge
Multiple Choice
1. Which of the following statements best describes the chemical
composition of biological membranes?
a. Biological membranes are bilayers of proteins with associated
lipids and carbohydrates.
b. Biological membranes are composed of two layers: one layer of
phospholipids and one layer of proteins.
c. Biological membranes are bilayers of phospholipids with
associated proteins and carbohydrates.
d. Biological membranes are composed of equal numbers of
phospholipids, proteins, and carbohydrates.
e. Biological membranes are composed of lipids with proteins
attached to the outer surface.
2. Which of the following events in a biological membrane would
not be energetically favorable and therefore would not occur
spontaneously?
Critical-Thinking Skills
Multiple Choice
1. Let’s suppose an insect, which doesn’t maintain a constant body
temperature, was exposed to a shift in temperature from 60°F
to 80°F. Which of the following types of membrane changes
would be the most beneficial to help this animal cope with the
temperature shift?
a. an increase in the number of double bonds in the nonpolar tails Biology Principle
of phospholipids Modeling
b. an increase in the length of the nonpolar tails of Biologists use models and simulations to test experimental predictions and
phospholipids convey their ideas. This figure is a model of plant biologists’ understand-
c. a decrease in the amount of cholesterol in the membrane ing of physical and biological features of plant environments that influence
plant growth responses. This understanding is key to the design of experi-
d. a decrease in the amount of carbohydrate attached to membrane
ments conducted on Earth and in space. The Modeling Challenge in the
proteins figure legend asks you to consider which features are most likely to be the
e. a decrease in the amount of carbohydrate attached to subjects of experiments conducted in space.
phospholipids Physical stimuli Biological stimuli
2. When placed in a solution of water, a human skin cell bursts. Internal:
Which of the following statements best explains why this Circadian
rhythms
happens? Hormones

a. the rotation of phospholipids a. The inside of the cell is hypertonic to the outside. Environmental: Environmental:
b. the lateral movement of phospholipids b. The inside of the cell is hypotonic to the outside.
Light
c. the flip-flop of phospholipids to the opposite leaflet c. The membrane contains transporters that allow the rapid Herbivores

d. the rotation of membrane proteins movement of ions.

e. the lateral movement of membrane proteins d. The membrane contains channels that allow the rapid movement Atmospheric
Agricultural
gases including
3. Which of the following movements is not an example of passive of ions. CO2 hormone
applications
Modeling Challenges
transport?
a. the movement of water through aquaporin
e. Both c and d are correct explanations.
3. The solute concentration inside the cells of a plant is 0.3 M; outside
Humidity

Temperature Pathogens
b. the intercellular transport of molecules via gap junctions it is 0.2 M. If we assume that the solutes do not readily cross the
A growing trend is the use of models in biology education. Students are asked to Touch, wind
Gravity

interpret models and to create models based on data or a scenario. Furthermore, Soil water
Organic
chemicals
emitted by
other plants
using models and simulations is one of the core skills that is emphasized by Rocks and
other barriers

“Vision and Change.” The author team has added a new feature called a Modeling Soil
minerals
Soil
microorganisms

Challenge, which asks students to create a model or to interpret a model they are
Figure 29.2 Types of plant stimuli. Plants respond to both physical
given. Possible answers to the Modeling Challenges are provided to the instructor and biological stimuli. Stimuli may be internal to the plant or come
from the environment.

in Connect. Modeling Challenge: Figure 29.2 depicts diverse physical

and biological stimuli that typically affect plant growth
responses. Earth-bound biologists keep this model of
plant responses in mind when designing experiments
to be conducted in the field or in a lab. Sketch a parallel
model showing physical and biological stimuli that plant
biologists might consider when designing experiments to
be conducted in space, such as in an orbiting laboratory. In
such a location, dirt, rocks, and other biological organisms
are unlikely to be welcome, unless well contained.

Principles of Biology: Helping Students Learn Critical-Thinking Skills v

bro23659_fm_i-xxviii.indd 5 30/09/22 8:54 PM

 SCISKILLS

The beginning of each section of every chapter contains a set
of Learning Outcomes that inform students of concepts they
should understand. Many sections contain skills-based Learning
Outcomes, labeled as SCISKILLS. These Learning Outcomes are
specific to the skills that students will acquire when mastering the
material and provide a specific understanding of how such skills
may be assessed. SCISKILLS are mental actions such as analyzing
data, forming hypotheses, making predictions, or performing
calculations. These are skills that scientists generally rely on and
students should practice.
Feature Investigations
The emphasis on skills development continues in the Feature Investigations.will
including data analysis, so that students can understand how experimentation leads to an understanding of biological concepts.
5.2 Fluidity of Membranes
Learning Outcomes
1. Describe the fluidity of membranes.
2. SCISKILLS ⊲ Predict how fluidity will be affected by
changes in lipid composition.
3. SCISKILLS ⊲ Analyze the results of experiments indicating
that certain membrane proteins can diffuse laterally within
the membrane.
Let’s now turn our attention to the dynamic properties of membranes.
Although a membrane provides a critical interface between a cell
or an organelle and its environment, it is not a solid, rigid structure.
Rather, biological membranes exhibit the property of fluidity, which
means that individual molecules remain in close association yet have
the ability to readily move within the membrane. In this section, we
These provide
examine a complete
the fluid properties ofdescription of experiments,
biological membranes.

FEATURE INVESTIGATION
Agre Discovered That Osmosis Occurs More Quickly
in Cells with a Channel That Allows the Facilitated
Diffusion of Water
Figure 5.13 The discovery of water channels (aquaporins) by Agre and colleagues.
As discussed earlier in this chapter, osmosis is the movement of frog oocytes, chosen because these oocytes are large, are easy to
water to balance solute concentrations. Water can cross biological inject, and lack pre-existingCHIP28
HYPOTHESIS proteins
may in their asplasma
function a watermembranes
channel.
membranes slowly by simple diffusion through the phospholipid that allow theKEY rapid movement of water. Following injection,
MATERIALS Prior to this work, a protein called CHIP28 was identified that is abundant in red blood cells and kidney cells. The gene
bilayer. However, in the 1980s, researchers discovered that certain the mRNA was translated to make CHIP28
that encodes this proteins,
protein waswhich
cloned,were
which means that many copies of the gene were made in a test tube.
cell types allow water to move across the plasma membrane at inserted into the plasma membrane of the oocytes. (Note: The pro-
a much faster rate than would be predicted by simple diffusion cess of translation is described in Chapter 10.) After sufficient Experimental level Conceptual level

alone. For example, water moves very quickly across the mem- time had been Add allowed for this
an enzyme (RNAtopolymerase)
occur, theand oocytes were placed CHIP28 mRNA RNA polymerase
1
brane of red blood cells, which causes them to shrink and swell in a hypotonicnucleotides
medium. to Asa test
a control,
tube thatoocytes
contains that had not been
many copies of the CHIP28 gene.exposed
This Enzymes
in response to changes in extracellular solute concentrations (see injected with CHIP28 mRNA were also to a hypotonic
and nucleotides
results in the synthesis of many copies
Figure 5.10a). Likewise, bladder and kidney cells, which play a medium. of CHIP28 mRNA.
key role in regulating water balance in the bodies of vertebrates, As you can see in the data, a striking difference was observed
allow the rapid movement of water across their membranes. Based between oocytes that expressed CHIP28 versus the control oocytes. CHIP28
DNA
on these observations, researchers speculated that certain cell Within minutes, oocytes that contained the CHIP28 protein were
types might have channels in their plasma membranes that enable seen to swell due to the rapid uptake of water. Three to five minutes
the rapid movement of water. after being placed in a hypotonic medium, they actually lysed! By
Inject the CHIP28 mRNA into frog eggs
One approach to characterizing a new protein is to first iden- comparison,2 the controlWait
(oocytes). oocytes
severaldid nottoswell
hours allow as rapidly, and they
tify a protein based on its relative abundance in a particular cell did not rupture,timeeven
for after 1 hour.
the mRNA to beTaken together,
translated into these results are
type and then attempt to determine the protein’s function. This CHIP28 protein at the ER membrane
consistent with the hypothesis that CHIP28 functions and as a channel
then moved via vesicles to the plasma CHIP28 protein is
rationale was applied to the discovery of proteins that allow the that allows themembrane.
facilitated diffusion of water across the membrane. CHIP28 inserted into the
rapid movement of water across membranes. Peter Agre and his Many subsequent studies confirmed this observation. Later, CHIP28 mRNA plasma membrane.
colleagues first identified a protein that was abundant in red blood was renamed aquaporin to indicate its newly identified function of CHIP28 protein
Frog oocyte
cells and kidney cells but not found in high amounts in many other allowing water to diffuse through a channel in the membrane. More
Nucleus Cytosol Ribosome
cell types. Though they initially did not know the function of the recently, the three-dimensional structure of aquaporin was determined
protein, its physical structure was similar to other proteins that (see chapter-opening figure). In 2003, Agre was awarded the Nobel
were already known to function as channels. They named this Prize in Chemistry for this work.
Place oocytes into a hypotonic medium
3
protein CHIP28, which stands for channel-forming integral mem- and observe under a light microscope.
brane protein with a molecular mass of 28,000 daltons. During the As a control, also place oocytes that
Control
have not been injected with CHIP28
course of their studies, they also identified and isolated the gene ExperimentalmRNA
Questions
into a hypotonic medium and
that encodes CHIP28. observe by microscopy.
1. What observations about particular cell types in the human body
In 1992, Agre and his colleagues conducted experiments to
led to the experimental strategy in Figure 5.13?
determine if CHIP28 functions in the transport of water across
membranes (Figure 5.13). Because they had already isolated the 2. What characteristics of CHIP28 made Agre and associates
gene that encodes CHIP28, they could make many copies of speculate that it may transport water? In your own words,
this gene in a test tube (in vitro) using gene cloning techniques briefly explain how they tested the hypothesis that CHIP28 has
4 THE DATA
(see Chapter 18). Starting with many copies of the gene in vitro, this function.
they added an enzyme to transcribe the gene into mRNA that 3. SCISKILLS ⊲ Explain how the results of the experiment in
Oocyte Oocyte rupturing
encodes the CHIP28 protein. This mRNA was then injected into Figure 5.13 support the proposed hypothesis.

3–5 minutes
CHIP28 protein

Control CHIP28 Control CHIP28

5 CONCLUSION The CHIP28 protein, now called aquaporin, allows the rapid movement of water across the membrane.

6 SOURCE Preston, G. M., Carroll, T. P., Guggino, W. B., and Agre, P. 1992. Appearance of Water Channels in Xenopus Oocytes Expressing Red Cell
CHIP28 Protein. Science 256: 385–387.

(micrographs) ©Dr. Peter Agre

vi Principles of Biology: Helping Students Learn Critical-Thinking Skills

bro23659_fm_i-xxviii.indd 6 30/09/22 8:54 PM

 Quantitative Analysis BioTIPS provide students with practice at applying these 11
problem-solving strategies.
The Quantitative Analysis feature helps students develop their
analytical skills. This feature walks them through biological Here’s how BioTIPS is organized: A biological question related
concepts that have a quantitative component. The concluding to chapter content is posed. The BioTIPS then walk the student
Crunching the Numbers provides a sample problem to test through the process of answering the question. First, they help
students’ understanding. The answers to the sample problems are the student identify the topic of the question—what is really
provided in Connect, the digital partner to this textbook. being asked in the question? Then they help the student collect
962 CHAPTER 43
information that was presented in the chapter that is related to
the question. Finally, they help the student settle on one or more
can catch species such as spiders, lizards, or beetles wandering
over the surface (Figure 43.1b). Sweep nets can be passed over grass-
land vegetation to dislodge and capture the insects feeding there. Mist
strategies that can be followed to answer the question. The
nets—very fine netting spread between trees—can entangle flying birds
and bats (Figure 43.1c). Baited live traps can capture terrestrial animals
answers are provided to complete the problem-solving process.
(Figure 43.1d). Population density can thus be estimated as the number
of animals caught per trap or per unit area where a given number of
traps are set—for example, 10 traps per 100 m2 of habitat.
For some larger terrestrial or marine species, captured animals
can be fitted with radio collars and followed remotely, using an
antennal tracking device. Their home ranges can be determined and
population estimates developed based on the area of available habi-
tat. Unmanned aircraft systems, or drones, have emerged as a safe,
low-cost method to document wildlife abundance and have been used
to provide animal counts in relatively inaccessible places, including
counts of walrus and stellar sea lions on the Aleutian Islands along
the coast of Alaska.
In the case of very mobile animals, such as fish in the ocean or
large, terrestrial animals with a large range, it may not be feasible
to measure population density directly. As an alternative, ecologists
may use different methods to estimate population size. For example, Figure 43.2 The mark-recapture technique for estimating population
they can estimate relative population size by examining catch per unit size. An ear tag identifies this Rocky Mountain goat (Oreamnos
effort, which is especially valuable in commercial fisheries. They americanus) in Olympic National Park, Washington. Recapture of
can’t easily expect to count the number of fish in an area of ocean, but such marked animals permits estimates of population size.
they can count the number caught—say, per 100 hours of trawling. W. Wayne Lockwood, M.D./Getty Images

Using our data,

Quantitative Analysis: Mark-Recapture Can Be
Used to Estimate Population Size
Sometimes, population biologists capture animals and then
tag and release them (Figure 43.2). The rationale behind
this mark-recapture technique is that after the tagged animals
are released, they mix freely with unmarked individuals and within
a short time are randomly mixed within the population. The popu-
lation is then resampled, and the numbers of marked and unmarked
individuals are recorded. It is assumed that the ratio of marked to
unmarked individuals in the second sample is the same as the ratio
of marked individuals in the first sample to the total population
size. Thus,
Number of individuals Number of marked recaptures
marked in first catch = _________________________
____________________ in second catch
Total population size, N Total number of second catch
Let’s say we catch 50 largemouth bass in a lake and mark
them with colored fin tags. A week later, we return to the lake and
catch 40 fish; 5 of them are previously tagged fish. If we assume
no immigration or emigration has occurred, which is quite likely in
a closed system such as a lake, and we assume there have been no
births or deaths of fish, then the total population size is given by
rearranging the equation:
Number of marked individuals in first catch
Total population ____________________________________
× Total number of second catch
=
size, N Number of marked
recaptures in second catch
50 × 40 = 400
N = ______
5
From this calculation, we estimate that the lake has a total popula-
tion size of 400 largemouth bass. This might be useful information
for game and fish personnel who wish to know the total size of a
fish population in order to set catch limits.
However, the mark-recapture technique can have drawbacks.
Some animals that have been marked may learn to avoid the
traps. Recapture rates will then be low, resulting in an overes-
timate of population size. Imagine that instead of 5 tagged fish
out of 40 recaptured fish, we get only 2 tagged fish. Now our
population size estimate is 2000/2 = 1000, a dramatic increase
in our population size estimate. On the other hand, some animals
can become “trap-happy,” particularly if the traps are baited with
food. This effect would result in an underestimate of the popula-
tion size.
Crunching the Numbers: In a study of rodent popula-
tions, you catch 60 mice, mark them with plastic ear tags, and
recatch 40, of which 12 are marked. What is the population size?
You later realize that half of the ear tags fell off the mice prior
to being recaptured. By how much have you misestimated the
population size?

BioTIPS

In Connect, we have a feature called BioTIPS, which is intended

to help students refine problem-solving skills. Most of the BioTIPS
are called out with icons in the textbook, but additional BioTIPS
are included in SmartBook. The BioTIPS themselves are accessed
through links in SmartBook. BioTIPS focus on 11 strategies that will Skills! Skills! Skills!
help students solve problems:
Overall, Principles of Biology, fourth edition, has six pedagogical
1. Make a drawing. features that foster skill development.
2. Compare and contrast.
1. Critical-Thinking Questions
3. Relate structure and function.
2. Modeling Challenges
4. Sort out the steps in a complicated process.
3. SCISKILLS Learning Outcomes
5. Propose a hypothesis.
4. Feature Investigations
6. Design an experiment.
5. Quantitative Analyses
7. Predict the outcome.
6. BioTIPS
8. Interpret data.
9. Use statistics. We expect that these features will help to lower the “activation
10. Make a calculation. energy barrier,” so that students can succeed in their skill
11. Search the literature. development.

Principles of Biology: Helping Students Learn Critical-Thinking Skills vii

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 Principles of Biology: Helping Students Learn
Cutting-Edge Content
We can all agree that biology textbooks cover a lot of material. To help students see the “forest and not just the trees” and to learn
concepts that are essential to their future careers, the author team has developed pedagogical features to help students navigate the
content in Principles of Biology, fourth edition.

WAYS TO HELP STUDENTS NAVIGATE AND LEARN THE MATERIAL IN THIS TEXTBOOK

Focus on Core Principles In addition, a question at the end of each chapter is directly aimed at
exploring a particular biology principle related to the content of the
Although it is intended for majors in the biological sciences, chapter. The answers to these questions are provided in Appendix B.
Principles of Biology is a shorter textbook that emphasizes core
principles. Twelve principles of biology are enunciated in Chapter 1 Short Answer
(see Figure 1.3), and those principles are emphasized throughout
1. Explain the functions of the crop and gizzard in birds. Propose a
the textbook. This emphasis is achieved in two ways. First, the reason for why humans did not evolve a crop or gizzard. Smooth,
principles are highlighted in selected figures in which a specific polished stones have been found in the stomach region of fossilized
skeletons of ancient sauropod dinosaurs. What does this suggest
principle is illustrated. about the alimentary canal of such animals?
2. PRINCIPLES: HOMEOSTASIS A principle of biology is that
Biology Principle living organisms maintain homeostasis. The mammalian kidneys
are an excellent example of this principle. Briefly, how many
Genetic Material homeostatic processes can you describe in which the kidneys play
The genetic material provides a blueprint that allows organisms to grow, a role?
develop, and reproduce. The covalent linkage of a sequence of bases
allows DNA (the genetic material) to store and transmit information.
Learning Outcomes

Each section of every chapter begins with a set of learning outcomes.

These outcomes help students understand what they should be able
Backbone Bases
O to do if they have mastered the material in that section.
CH3 C H
C N

C C
Thymine (T) Formative Assessment
O– H N O
5′ O P O CH2

O–
5′
4′ C H
O

H
C 1′ Many instructors are devoting more class time to active learning
H
and spending less time on directly discussing the textbook
H
C C 2′ NH2
3′
H C
N

O H C
C N
Adenine (A) material. When students are expected to learn more textbook
C C
O P O
5′
CH2
O
N N H
material on their own, it is imperative that they be given regular
O– 4′ C H C 1′
H
C
H
H
C 2′ H
NH2
C
formative assessments so they can gauge whether they are
3′
H
C

C
N

C
Cytosine (C) mastering the material. Formative assessment is a major feature of
Phosphodiester
linkage O
O

P O CH2
H N O
this textbook and is bolstered by McGraw Hill Connect—a state-of-
the-art digital assignment and assessment platform. In Principles of
5′ O
O– 4′ C H C 1′
H Guanine (G)
H H
3′
C
H
C 2′ O
C H
Biology, formative assessment is provided in multiple ways.
N
C N
H C

O
O

P O CH2
N
C
N
C
NH2
1. Each section of every chapter ends with a set of multiple-
Single O
choice questions.
5′
nucleotide O– 4′ C H C 1′
H
Phosphate H C H
3′
OH
C 2′
H
2. Most figures have Concept Check questions so students can
Sugar (deoxyribose)
determine if they understand the key points in the figure.
3′
3. End-of-chapter questions continue to provide students with
feedback regarding their mastery of the material. Note: The
answers to the end-of-section and end-of-chapter questions
Figure 9.5 The structure of a DNA strand. Nucleotides are are provided in Appendix B.
covalently bonded to each other in a linear manner. Notice
the directionality of the strand and that it carries a particular 4. Further assessment tools are available in Connect. Question
sequence of bases. An RNA strand has a very similar structure,
except the sugar is ribose rather than deoxyribose and uracil is banks, test banks, quantitative questions, and graphing
substituted for thymine.
interactive banks can be assigned by the professor. McGraw
Concept Check: What is the difference between a
phosphoester bond and a phosphodiester linkage?
Hill SmartBook provides students with real-time assessment
and feedback of their learning as they are moving through
each chapter, page by page.

viii Principles of Biology: Helping Students Learn Cutting-Edge Content

bro23659_fm_i-xxviii.indd 8 30/09/22 8:54 PM

 BioConnections and Evolutionary Connections may influence the sizes and numbers of these organelles. For example,
Animals, fungi, and protists
Plants and algae
(contain mitochondria
when plants are exposed to more sunlight, the number of chloroplasts (contain mitochondria) and chloroplasts)
in leaf cells increases.
To help students broaden their understanding of biology, 0

EVOLUTIONARY CONNECTIONS
two recurring features are BioConnections and Evolutionary
Mitochondria and Chloroplasts Are Derived from Ancient
Connections. BioConnections questions are placed in key figure Symbiotic Relationships

Billions of years ago (bya)

Primordial
legends in most chapters and help students relate a topic they The observation that mitochondria and chloroplasts contain their own
genetic material may seem puzzling. Perhaps you might think that it
Evolution eukaryotic
cells
Evolution

would be simpler for a eukaryotic cell to have all of its genetic mate-
are currently learning to another topic elsewhere in the textbook, rial in one place—the nucleus. The distinct genomes of mitochon- 1

dria and chloroplasts can be traced to their evolutionary origin, which

often in a different unit. (The answers are provided in Appendix B.)
Evolutionary Connections provide a framework for understanding
how a topic in a given chapter relates to evolution, the core
unifying theme in biology.
involved an ancient symbiotic association.
A symbiotic relationship occurs when two different species
live in direct contact with each other. Endosymbiosis is a symbi-
otic relationship in which the smaller species (the symbiont) actually
lives inside the larger species (the host). In 1883, Andreas Schimper
proposed that chloroplasts evolved from an endosymbiotic relation- Cyanobacterium
ship between cyanobacteria (bacteria capable of photosynthesis) and
eukaryotic cells. In 1922, Ivan Wallin also hypothesized an endo- 2 Proteobacterium

symbiotic origin for mitochondria. Researchers now hypothesize

that mitochondria evolved from an ancient species related to a group
of modern bacteria called proteobacteria, which are described in
Chapter 23.
CAP site In spite of these interesting proposals, the question of whether
endosymbiosis gave rise to mitochondria and chloroplasts was largely
ignored until the discovery that these organelles contain their own
genetic material. In 1970, the idea of endosymbiosis as the origin of
mitochondria and chloroplasts was revived by Lynn Margulis in her
book Origin of Eukaryotic Cells. During the 1970s and 1980s, the
advent of molecular genetic techniques allowed researchers to ana-
lyze genes from mitochondria, chloroplasts, bacteria, and eukaryotic
nuclear genomes. Researchers discovered that genes in mitochondria
and chloroplasts are very similar to bacterial genes. Likewise, mito-
chondria and chloroplasts are strikingly similar in size and shape to
certain bacterial species.
These observations provided strong support for the
endosymbiosis theory, which proposes that mitochondria and chlo-
roplasts originated from an endosymbiotic relationship that gave rise
to eukaryotic cells (Figure 4.29). Over the next 2 billion years, the
characteristics of these intracellular bacterial cells gradually changed
to those of mitochondria or chloroplasts. The origin of eukaryotic
cells is discussed in more detail in Chapter 22.
Three- A symbiotic relationship is beneficial to one or both species.
According to the endosymbiosis theory, such a relationship provided
dimensional DNA eukaryotic cells with useful cellular characteristics. Chloroplasts,
structure of which were derived from cyanobacteria, have the ability to carry
out photosynthesis. This benefits plant cells by giving them the abil-
CAP bound ity to use the energy from sunlight. By comparison, mitochondria
to the CAP cAMP are thought to have been derived from a different type of bacteria
site known as proteobacteria. In this case, the endosymbiotic relationship
enabled eukaryotic cells to synthesize greater amounts of ATP. How
the relationship would have been beneficial to a cyanobacterium or
(a) Mitochondria originated (b) Chloroplasts originated
from endosymbiotic from endosymbiotic
proteobacteria. cyanobacteria.
Figure 4.29 A simplified view of the endosymbiosis theory.
(a) According to the endosymbiosis theory, modern mitochondria
were derived from a type of bacteria called proteobacteria. Over
the course of evolution, their characteristics changed into those
found in mitochondria today. (b) A similar phenomenon occurred for
chloroplasts, which were derived from cyanobacteria (blue-green
bacteria), bacteria that are capable of photosynthesis.
Concept Check: Discuss the similarities and differences
between modern bacteria and mitochondria.
proteobacterium is less clear, though the cytosol of a eukaryotic cell
may have provided a stable environment with an adequate supply of
nutrients.
During the evolution of eukaryotic species, many genes that
were originally found in the genomes of the primordial proteobac-
teria and cyanobacteria have been transferred from the organelles to
the nucleus. This has occurred many times throughout evolution, so
modern mitochondria and chloroplasts have lost most of the genes
that still exist in present-day proteobacteria and cyanobacteria. Some
researchers speculate that the movement of genes into the nucleus
makes it easier for the cell to control the structure, function, and divi-
sion of mitochondria and chloroplasts. In modern cells, hundreds of
different proteins that make up these organelles are coded by genes
that have been transferred to the nucleus. These proteins are made in
the cytosol and then taken up into mitochondria or chloroplasts. We
will discuss this topic next.

CAP dimer

CAP site

Promoter Operator
CAP cAMP Transcription
occurs.

Binding of RNA polymerase

to promoter is enhanced mRNA
by CAP binding. RNA polymerase

Figure 12.8 Positive control of the lac operon by the catabolite activator
protein (CAP). When cAMP is bound to CAP, CAP binds to the
major groove of the DNA and causes the DNA to bend. This bend
facilitates the binding of RNA polymerase.
Image from the RCSB PDB (www.rcsb.org; H.M. Berman, et al. (2000), “The
Protein Data Bank,” Nucleic Acids Research, 28: 235–242) of PDB ID 1CGP
(Schultz, S.C., Shields, G.C., Steitz, T.A.) (1991) Science, 253: 1001–1007.

BioConnections: Refer back to Figure 8.13. What is the

function of cAMP in eukaryotic cells?

Principles of Biology: Helping Students Learn Cutting-Edge Content ix

bro23659_fm_i-xxviii.indd 9 30/09/22 8:54 PM

 New! Additional eBook Questions

A new feature in the eBook of the

fourth edition of Principles of Biology
is the addition of eBook Questions
that support the Biology Principles
and BioConnections figures. Many
of the Biology Principles figures and
figures that have BioConnections
now have a link to an additional
question. These questions are
expected to bolster a student’s
appreciation and understanding of
these important concepts and skills.

Unit Openers

Unit openers serve two purposes. They allow the student to

9
see the big picture of the unit. In addition, the unit openers draw UNIT III
attention to the principles of biology that will be emphasized in that 10 GENETICS
unit.
Genetics is the branch of biology that deals with inheritance—the transmission
of characteristics from parent to offspring. We will begin this unit by examining
the structure of the genetic material, namely DNA, at the molecular and cellular
11 levels. We will explore the structure and replication of DNA and examine how

Overall, the pedagogy of Principles of Biology has been designed
to foster student learning. Instead of being a collection of “facts
0.1 µm the DNA is packaged into chromosomes (Chapter 9). We will then consider
how segments of DNA are organized into units called genes and explore how
12 genes are used to make products such as mRNAs, proteins, and noncoding
1 mm
RNAs (Chapters 10 and 11). In Chapter 12, we will consider how the expression
of genes is regulated. Gene expression is largely responsible for the

and figures,” Principles of Biology is intended to be an engaging 13

characteristics of living organisms. We will also examine how mutations can alter
the properties of genes and even lead to diseases such as cancer (Chapter 13).
In Chapter 14, we will turn our attention to the mechanisms by which
and motivating textbook in which formative assessment and many genes are transmitted from parent to offspring, beginning with a discussion of
how chromosomes are sorted and transmitted during cell division. Chapters
14
other pedagogical features allow students to move ahead and 15 and 16 explore the relationships between the transmission of genes and
the outcome of an offspring’s traits. We will look at genetic patterns called
Mendelian inheritance, named after Gregor Mendel, the 19th-century biologist
learn the material in a productive way. who discovered them, as well as more complex patterns that could not have
15 been predicted from Mendel’s work.
Chapters 9 through 16 focus on the fundamental properties of the
5 µm

genetic material and heredity. The remaining chapters explore additional top-
ics that are of importance to biologists. In Chapter 17, we will examine some of
16 the unique genetic properties of bacteria and viruses. Chapter 18 describes
genetic technologies that are used by researchers, clinicians, and biotech-
nologists to unlock the mysteries of genes and that provide tools and applica-
tions that benefit humans, and the chapter also explores the entire genomes
17 of bacteria, archaea, and eukaryotes.

The following biology principles will be emphasized

18 in this unit:

● The genetic material provides a blueprint that allows organisms

(9) Prof. Kenneth Seddon & Dr. Timothy Evans,
Queen’s Univ. Belfast/SPL/Science Source; (10) Elena
Kiseleva/Science Source; (11) ©Mauro Giacca,
Ana Eulalio, Miguel Mano; (12) Image from the
RCSB PDB (www.rcsb.org; H.M. Berman,
et al. (2000), “The Protein Data Bank,” Nucleic
Acids Research, 28: 235—242) of PDB ID 1CGP
(Schultz, S.C., Shields, G.C., Steitz, T.A. (1991)
Science, 253, 1001–1007); (13) Pikselstock/
Shutterstock; (14) Biophoto Associates/Science
Source; (15) Tomas Rodriguez/Getty Images;
(16) Thiriet/Andia/Alamy Stock Photo; (17) Norm
Thomas/Science Source; (18) ©Fumihiro Sugiyama
to grow, develop, and reproduce. Throughout this unit, we will see
how the genetic material carries the information for reproduction and
sustaining life.
● The structural features of living organisms determine their functions.
In Chapters 9 through 14, we will examine how the structure of DNA,
RNA, genes, and chromosomes underlies their functions.
● Living organisms interact with their environment. In Chapters 15 and
16, we will explore the interactions between an organism’s genes and
its environment.
● Biology affects our society. In Chapter 18, we will examine genetic
technologies that have many applications in our society.
● Biology is an experimental science. Most chapters in this unit have a
Feature Investigation that describes a pivotal experiment that provided
insights into our understanding of genetics.
● Biologists use models and simulations to test experimental predic-
tions and convey their ideas. Every chapter has a Modeling Challenge
to help you refine this important skill.

x Principles of Biology: Helping Students Learn Cutting-Edge Content

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 Content Changes to the Fourth Edition
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 for colors, placement of
colors, and other contrast issues to meet accessibility standards.
The following are the major content changes that have occurred
in the fourth edition.
Chapter 1 An Introduction to Biology. This chapter has a new
subsection on tuskless elephants (see Figure 1.7). Section 1.2, Unity
and Diversity of Life, has been split into two sections: Biological
Evolution and Classification of Living Things. The concept of organ
systems was inserted into Figure 1.4.
Chapter 2 The Chemical Basis of Life I: Atoms, Molecules, and
Water. A new figure describes the electronegativity differences
among nonpolar covalent, polar covalent, and ionic bonds (see
Figure 2.5). A new illustration in Section 2.2 compares the polarity
of water and carbon dioxide. In Section 2.4, the Quantitative
Analysis subsection has been revised to include a question on the
calculation of insulin levels before and after a meal.
Chapter 3 The Chemical Basis of Life II: Organic Molecules.
Figure 3.5, the synthesis of sucrose, has been revised for clarity.
Figure 3.6 now shows the overall arrangements of polysaccharides
in starch, glycogen, and cellulose. Figure 3.13a has been revised to
more accurately show how peptide bonds are formed.
Chapter 4 Evolutionary Origin of Cells and Their General
Features. With regard to the origin of living cells, new information
about alkaline hydrothermal deep sea vents has been added.
With regard to microscopy, the difference between resolution and
contrast has been clarified. Figure 4.19 has been revised to better
illustrate the structure of nuclear pores. In Table 4.1, the scales of
the drawings of microtubules, intermediate filaments, and actin
filaments were made equivalent so that students can readily
compare their relative sizes.
Chapter 5 Membranes: The Interface Between Cells and Their
Environment. Figure 5.9 has been revised to better depict the
direction of water movement during osmosis.
Chapter 6 How Cells Utilize Energy. Figure 6.19 has been revised
to reflect structural studies showing that H+ ions move through two
half-channels as they pass through ATP synthase. New information
has been added indicating that citric acid intermediates are used
for the biosynthesis of important cellular molecules, such as amino
acids. The overview of cellular respiration is now in its own section
(see Section 6.4).
Chapter 7 How Cells Capture Light Energy via Photosynthesis.
A subsection shows how photosynthesis involves the reduction
bro23659_fm_i-xxviii.indd 11
of carbon atoms to form carbohydrates; it includes a new figure
(see Figure 7.13). The description of the light reactions has been
modified to clarify the path of the electrons that move from PSII to
PSI. A leaf vein has been added to Figure 7.2 for clarity.
Chapter 8 How Cells Communicate with Each Other and with the
Environment. A new summary table compares the different types
of cellular receptors (see Table 8.1). A new section describes the
topic of crosstalk among signal transduction pathways; it includes a
new figure (see Figure 8.17 in Section 8.7).
Chapter 9 The information of Life: DNA and RNA Structure, DNA
Replication, and Chromosome Structure. A new figure shows
the formation of loop domains in eukaryotic chromosomes (see
Figure 9.21).
Chapter 10 The Expression of Genetic Information via Genes I:
Transcription and Translation. New information has been added
about the metalloribozyme in the spliceosome.
Chapter 11 The Expression of Genetic Information via Genes II:
Non-coding RNAs. The illustration of a non-coding RNA acting as
blocker has been revised to a two-part figure (see Figure 11.3).
Chapter 12 The Control of Genetic Information via Gene
Regulation. A new subsection discusses the similarities of
transcription in eukaryotes and archaea. Another new subsection
describes how the formation of facultative heterochromatin is a
way to regulate genes in a tissue-specific manner; it includes a new
figure (see Figure 12.17).
Chapter 13 Altering the Genetic Material: Mutation, DNA Repair,
and Cancer. A new subsection describes how mutations in the
coding sequences of genes can cause human disease; it includes
a new table (see Table 13.2). The figure describing the interaction
between E2F and Rb has been moved to Chapter 13 (from
Chapter 14). It is now Figure 13.14.
Chapter 14 How Eukaryotic Cells Sort and Transmit
Chromosomes: Mitosis and Meiosis. The topic of sexual
reproduction has now been placed in its own section (now Section
14.4), which includes a new subsection on how sexual reproduction
can foster genetic diversity and provide disease resistance.
Chapter 15 Transmission of Genetic Information from Parents to
Offspring I: Patterns That Follow Mendel’s Laws. The topic of how
temperature affects sex determination in alligators has been updated.
Chapter 16 Transmission of Genetic Information from Parents to
Offspring II: Epigenetics, Linkage, and Extranuclear Inheritance.
A new subsection describes how a chi square analysis can be
used to determine if two genes are linked (see Table 16.5). A study
by NASA regarding epigenetic changes associated with space
flight is discussed.
Content Changes to the Fourth Edition xi
30/09/22 8:54 PM
 Chapter 17 The Simpler Genetic Systems of Viruses, Bacteria,
and Archaea. New information has been added about COVID-19
(see Table 17.1 and Figure 17.3).
Chapter 18 Genetic Technologies: How Biologists Study Genes
and Genomes. This chapter now contains the technique of RNA
sequencing (see Figure 18.10). A new table describes applications
of CRISPR-Cas technology (see Table 18.3). A new subsection on
biotechnology provides examples of transgenic organisms (see
Figures 18.12, 18.13).
Chapter 19 Evolution of Life I: How Populations Change from
Generation to Generation. The contributions of Alfred Wallace to
the theory of evolution have been expanded. A new example of
convergent evolution, involving hummingbirds and hummingbird
moths, has been added (see Figure 19.6a).
Chapter 20 Evolution of Life II: The Emergence of New Species.
A new subsection describes hybrid zones, which includes three
new figures (see Figures 20.10–20.12). A new example of sympatric
speciation, namely the formation of the Big Bird lineage on a
Galápagos Island, has been added (see Figure 20.13).
Chapter 21 How Biologists Classify Species and Study Their
Evolutionary Relationships. Using giraffes as an example, a new
subsection describes how new information can result in changes in
taxonomy (see Figure 21.3). This chapter contains a new figure on
horizontal gene transfer (see Figure 21.13).
Chapter 22 The History of Life on Earth and Human Evolution.
Additional information regarding the effects of the ozone
layer on the history of life has been added. Based on research
in 2019, the possible effects of the K-T event during the
Cretaceous period are described. Additional possible reasons
for bipedalism in humans are discussed. A description of new
evidence that Neanderthals and Denisovans interbred has been
added.
Chapter 23 Diversity of Microbial Life: Archaea, Bacteria,
Protists, and Fungi. Phylogenetic diagrams were reviewed for
consistency with recent findings, and the fungal relationship
diagram revised accordingly.
Chapter 24 Microbiomes: Microbial Systems on and Around Us. A
new Feature Investigation illustrates modern efforts to understand
how environment and heredity influence the microbiomes of animal
hosts, and how research design, specifically sample sizes, influence
research results. A new example of animal microbiome engineering
is explained and accompanied by a new figure.
Chapter 25 Plant Evolution: How Plant Diversification Changed
Planet Earth. A phylogenetic illustration of plant evolution has
been updated according to new findings. This update, showing
bryophyte monophyly, is key to understanding the process by
which vascular plants arose.
Chapter 26 Invertebrates: The Vast Array of Animal Life Without
a Backbone. In Section 26.2, Animal Classification, new material
has been added to the discussion of animal phylogeny, illustrated
by Figure 26.2.
xii Content Changes to the Fourth Edition
bro23659_fm_i-xxviii.indd 12
Chapter 28 An Introduction to Flowering Plant Form and
Function. Information about the roles of genes and proteins
involved in plant development has been updated.
Chapter 29 How Flowering Plants Sense and Interact with Their
Environments. An evolutionary connection feature about the
evolutionary history of gibberellin responses has been revised,
based on new research findings, and illustrated by a revised figure.
New information has been added about the important role of the
mammalian neurotransmitter glutamate in plant signaling.
Chapter 30 How Flowering Plants Obtain and Transport Water,
Mineral Nutrients, and Organic Compounds. A new image showing
a biological soil crust effectively illustrates nitrogen-fixing microbes
and lichens that play key roles in soil fertility, particularly in arid lands.
Chapter 31 How Flowering Plants Reproduce and Develop. This
chapter, which covers diverse aspects of flower and fruit structure
and development, has been trimmed for more concise delivery.
Chapter 32 General Features of Animal Bodies, and Homeostasis
as a Key Principle of Animal Biology. The characteristics of
conformers have been clarified. A new table (Table 32.2) that lists
and describes common homeostatic variables in animals has been
added. A new photo of an animal responding to a homeostatic
challenge has been included.
Chapter 33 Neuroscience I: The Structure, Function, and
Evolution of Nervous Systems. The quantitative description of the
utility of the Nernst equation has been simplified.
Chapter 34 Neuroscience II: How Sensory Systems Allow
Animals to Interact with the Environment. Comparative aspects
of mechanoreception including hearing and balance in different
animals have been added. Recent Nobel Prize research by
David Julius and Ardem Patapoutian on the mechanisms of
thermoreception has been described. A new micrograph of a
section through a vertebrate retina has been added to Figure 34.17.
Chapter 35 How Muscles and Skeletons Are Adaptations
for Movement, Support, and Protection. The text has been
streamlined for clarity by the introduction of numerous bulleted lists
and shorter paragraphs. Several figures have been simplified and
re-drawn for clarity.
Chapter 36 Circulatory and Respiratory Systems: Transporting
Solutes and Exchanging Gases. A new light micrograph that
compares the structures of an artery and vein has been included.
A new photograph of an animal with external gills has been added.
The values in Table 36.2 have been updated.
Chapter 37 Digestive and Excretory Systems: Maintaining
Nutrient, Water, and Energy Balance and Removing Waste. The
major dietary categories of animals have been presented as a
bulleted list with additional examples of animals. The discussion
of the importance of salivary digestion of carbohydrates in some
animals has been expanded. Long segments of text have been
carefully edited and broken into manageable segments for
improved readability.
30/09/22 8:54 PM
 Chapter 38 How Endocrine Systems Influence the Activities of
All Other Organ Systems. Additional introductory material has
been added to several sections, emphasizing comparative and
evolutionary features of hormones in animals.
Chapter 39 The Production of Offspring: Reproduction and
Development. A new image of copulation in blue damselflies has
been added (Figure 39.5). A new illustration of the stages of labor
in a mammal (cow) has been added (Figure 39.13). Numerous
bulleted lists and paragraph breaks have been added to improve
readability and focus.
Chapter 40 Immune Systems: How Animals Defend Against
Pathogens and Other Dangers. Several illustrations have been
modified for clarity. SARS-CoV-2 and COVID-19 are introduced
and discussed in several places throughout the chapter, including
methods of detecting the virus and how it affects the immune system.
Edward Jenner’s innovations regarding vaccinations are discussed.
Chapter 41 An Example of a System-Wide Response to a
Challenge to Homeostasis. Additional examples of major
homeostatic challenges are described.
Chapter 42 Behavioral Ecology: The Struggle to Find Food and
Mates and to Pass On Genes. Section 42.3, Living in Groups and
Game Theory, has been re-written and expanded, including a
new table summarizing the costs and benefits of group living (see
Table 42.1) and a new figure illustrating an advantage of living in
groups (see Figure 42.8).
Chapter 43 Population Growth and Species Interactions. A
completely new Section 43.5, The Spread of Pathogens, discusses
the spread of diseases and includes up-to-date information on the
basic reproduction number, R0, the factors that affect the rates of
bro23659_fm_i-xxviii.indd 13
disease spread, how pathogens may arise from mutations in existing
human pathogens or via spreading from animals, how vaccination
can decrease the rate of spread, and the concept of herd immunity.
Chapter 44 Communities and Ecosystems: Ecological
Organization of Large Scales. A new chapter opener discusses
the recovery of life after volcanic eruptions on islands. In Section
44.3, Succession: Community Change, a new figure shows how
primary succession on sand dunes supports the facilitation model
(see Figure 44.5).
Chapter 46 The Age of Humans. A new feature of this chapter is
that each section ends with a short paragraph offering solutions
to some of the ecological effects of humans. Much of the data
in Section 46.1, Human Population Growth, has been updated,
including Figure 46.2, The Age Structure of Human Populations,
and Figure 46.3, Total Fertility Rates. We have also updated
Figure 46.5, which illustrates the ecological footprints of different
countries. The data on global warming and greenhouse gases
has also been updated. In Section 46.5, Habitat Destruction,
new examples of the adverse ecological changes brought about
by agriculture are provided, including the extinction of North
America’s only locust species in the 19th century. Finally, following
a year (2019) in which “murder hornets” from Asia were found in
the U.S., expanded coverage of invasive species is provided.
Chapter 47 Biodiversity and Conservation Biology. A new
chapter opener discusses the effects of the world’s most
destructive fungal pathogen, Batrachochytrium dendrobatidis, on
the worldwide decline of amphibian species, especially frogs. The
chapter ends with a new discussion on sustainable development,
an organizing principle that may help the survival of species now
and in the future.
Content Changes to the Fourth Edition xiii
30/09/22 8:54 PM
 Strengthen Critical-Thinking Skills with Connect®
Critical-Thinking Skills
Detailed Feedback in Connect®
Learning is a process of iterative development, of making mistakes,
reflecting, and adjusting over time. The question and test banks in
Connect® for Principles of Biology, fourth edition, are more than
direct assessments; they are self-contained learning experiences that
systematically build student learning over time.
For many students, choosing the right answer is not necessarily
based on applying content correctly; it is more a matter of increasing
their statistical odds of guessing. A major fault with this approach is
that students don’t learn how to process the questions correctly, mostly
because they are repeating and reinforcing their mistakes rather than
reflecting and learning from them. To help students develop problem-
solving skills, all higher-level Bloom’s questions in Connect® are sup-
ported with hints, to help students focus on important information
for answering the question. After submitting an answer, the student
is given detailed feedback that walks through the problem-solving
process, using Socratic questions in a decision tree–style framework
xiv Strengthen Critical-Thinking Skills with Connect®
bro23659_fm_i-xxviii.indd 14
to scaffold learning. Each step models and reinforces the learning
process.
The feedback for each higher-level Bloom’s question (Apply,
Analyze, Evaluate) follows a similar process: Clarify Question,
Gather Content, Choose Answer, Reflect on Process.
Unpacking the Concept
We’ve taken problem solving a step further. In each chapter, two
higher-level Bloom’s questions from the question and test banks are
broken down according to the steps in the detailed feedback.
Rather than leaving it up to the student to work through the
detailed feedback, we present a second version of the question in
a stepwise format. Following the problem-solving steps, in the
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 Acknowledgments
The lives of most science-textbook authors do not revolve around an Reviewers for Principles of Biology, fourth edition
analysis of writing techniques. Instead, we are people who understand
Sarah Arrington Appalachian State University
science and are inspired by it, and we want to communicate that infor-
mation to our students. Simply put, we need a lot of help to get it Anna Cunningham Arizona State University
right.
Gloria J. DeWalt Northeastern University
Editors are a key component that help the authors modify the
content of their book so it is logical, easy to read, and inspiring. The Ann M. Findley University of Louisiana at Monroe
editorial team for Principles of Biology, fourth edition, has been a
Donna-Marie Gardner Middlesex College
catalyst that kept this project rolling. The members played various
roles in the editorial process. Ian Townsend and Lora Neyens most Devyn Gillette Bowie State University
recently, Brand Manager for Majors Biology, did an outstanding job
Cheri Jones University of Colorado Denver
of overseeing the development of this new text. Elizabeth Sievers,
Senior Product Developer, has been the master organizer. Liz is the Jonathan Karpel Southern Utah University
glue that keeps each new edition moving forward. The author team is
Yevgeniya Lapik Harold Washington College
deeply indebted to her dedication to this textbook.
We would also like to acknowledge our copy editor, Carey Zachary Long University of North Carolina Wilmington
Lange, for keeping our grammar on track, as well as Susan Gall for
Kristen Miller University of Georgia
her excellent efforts at proofreading.
Another important aspect of the editorial process is the actual Aude Amelie Picard University of Nevada Las Vegas
design, presentation, and layout of materials. It’s confusing if the text
Laura Rusche University of Buffalo SUNY
and art aren’t near each other or if a figure is too large or too small.
We are indebted to the tireless efforts of Jessica Portz, Senior Content Lucia Santacruz Bowie State University
Project Manager, and David Hash, Senior Designer, at McGraw Hill.
Thomas Sasek University of Louisiana Monroe
Likewise, our production company, MPS Limited, did an excellent
job with the paging and art revisions. Manar Soliman Middlesex College
We would like to acknowledge the ongoing efforts of the superb
Chad Wayne University of Houston
marketing staff at McGraw Hill. Special thanks to Kelly Brown,
Executive Marketing Manager, Life Sciences, for his ideas and enthu- Elizabeth Zulick Northeastern University
siasm for this book.
We would like to thank the digital authors and subject mat-
ter experts who helped in the development of the digital assets in
Connect® that support Principles of Biology, fourth edition.
The authors are grateful for the help, support, and patience of their
families, friends, and students: Deb, Dan, Nate, and Sarah Brooker;
Maria, Caroline, and Richard Widmaier; Jim, Michael, and Melissa
Graham; and Jacqui, Zoe, Leah, and Jenna Stiling.

xx Acknowledgments

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 Detailed Table of Contents

Feature Investigation: Anfinsen Showed That the Primary

CHAPTER 1 Structure of Ribonuclease Determines Its Three-Dimensional
Structure 52
An Introduction to Biology 1 Evolutionary Connections: Proteins Contain Functional
Domains 54
1.1 Principles of Biology and the Levels of Biological
3.7 Nucleic Acids 55
Organization 2
1.2 Biological Evolution 7
Evolutionary Connections: The Frequency of the Tuskless UNIT II Cells
Elephant Appears to Be Increasing in Elephant Populations Due to
Poaching 9
1.3 Classification of Living Things 10
1.4 Biology as a Scientific Discipline 13

UNIT I Chemistry

Steve Gschmeissner/Science Source

CHAPTER 4

Evolutionary Origin of Cells

Vladislav Gajic/Shutterstock
and Their General Features 59
4.1 Origin of Living Cells on Earth 60
4.2 Microscopy 66
CHAPTER 2 4.3 Overview of Cell Structure and Function 69
4.4 The Cytosol 74
The Chemical Basis of Life I: 4.5 The Nucleus and Endomembrane System 79
Atoms, Molecules, and Water 21 4.6 Semiautonomous Organelles 85
Evolutionary Connections: Mitochondria and Chloroplasts Are
2.1 Atoms 22
Derived from Ancient Symbiotic Relationships 88
2.2 Chemical Bonds and Molecules 25
4.7 Protein Sorting to Organelles 89
2.3 Chemical Reactions 29
4.8 Extracellular Matrix and Plant Cell Walls 89
2.4 Properties of Water 30
4.9 Systems Biology of Cells: A Summary 94
Quantitative Analysis: Concentrations of Molecules in Solution
Can Be Defined by Mass and Moles 31
2.5 pH and Buffers 34 CHAPTER 5

CHAPTER 3 Membranes: The Interface Between Cells

and Their Environment 98
The Chemical Basis of Life II: 5.1 Membrane Structure 99
Organic Molecules 37 5.2 Fluidity of Membranes 100
3.1 The Carbon Atom and Carbon-Containing 5.3 Overview of Membrane Transport 103
Molecules 38 5.4 Proteins That Carry Out
3.2 Synthesis and Breakdown of Organic Molecules 40 Membrane Transport 107
Feature Investigation: Agre Discovered That Osmosis Occurs
3.3 Overview of the Four Major Classes of Organic
More Quickly in Cells with a Channel That Allows the Facilitated
Molecules Found in Living Cells 40
Diffusion of Water 108
3.4 Carbohydrates 41
5.5 Intercellular Channels 113
3.5 Lipids 44
5.6 Exocytosis and Endocytosis 114
3.6 Proteins 47
5.7 Cell Junctions 117

xxi

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 xxii Detailed Table of Contents

CHAPTER 6 UNIT III Genetics

How Cells Utilize Energy 121
6.1 Energy and Chemical Reactions 122
6.2 Enzymes 124
Quantitative Analysis: Enzyme Function Is Influenced by
Substrate Concentration and by Inhibitors 126
6.3 Metabolic Pathways 129
6.4 Overview of Cellular Respiration 132
6.5 Glycolysis 134
6.6 Breakdown of Pyruvate 137
Prof. Kenneth Seddon & Dr. Timothy Evans, Queen’s Univ. Belfast/SPL/Science
6.7 Citric Acid Cycle 138 Source
6.8 Oxidative Phosphorylation 139
Feature Investigation: Yoshida and Kinosita Demonstrated
That the γ Subunit of ATP Synthase Spins 143 CHAPTER 9
6.9 Connections Among Carbohydrate, Protein, and Fat
Metabolism 145 The Information of Life: DNA and RNA Structure, DNA
6.10 Anaerobic Respiration and Fermentation 146
Replication, and Chromosome Structure 189
9.1 Properties and Identification of the
CHAPTER 7
Genetic Material 190
Feature Investigation: Avery, MacLeod, and McCarty
How Cells Capture Light Energy Used Purification Methods to Reveal That DNA Is the
via Photosynthesis 150 Genetic Material 191
9.2 Nucleic Acid Structure 193
7.1 Overview of Photosynthesis 151
9.3 Discovery of the Double-Helix Structure of DNA 197
7.2 Reactions That Harness Light Energy 153
9.4 Overview of DNA Replication 199
7.3 Molecular Features of Photosystems 159
9.5 Molecular Mechanism of DNA Replication 202
7.4 Synthesizing Carbohydrates via the Calvin Cycle 161
Feature Investigation: The Calvin Cycle Was Determined
9.6 Molecular Structure of Eukaryotic Chromosomes 206
by Isotope-Labeling Methods 163
7.5 Variations in Photosynthesis 165 CHAPTER 10
Evolutionary Connections: C4 and CAM Plants Have Evolved a
Mechanism to Minimize Photorespiration 165
The Expression of Genetic Information via Genes I:
Transcription and Translation 211
CHAPTER 8
10.1 Overview of Gene Expression 212
10.2 Transcription 213
How Cells Communicate with Each Other
10.3 RNA Modifications in Eukaryotes 215
and with the Environment 169 10.4 Translation and the Genetic Code 218
8.1 General Features of Cell Communication 170 Feature Investigation: Nirenberg and Leder Found That an RNA
8.2 Receptor Activation 173 Triplet Can Promote the Binding of a tRNA to a Ribosome 220
Quantitative Analysis: Receptors Have a Measurable Affinity for 10.5 The Machinery of Translation 222
Their Ligands 174 Evolutionary Connections: Comparisons of Small Subunit
8.3 Cell Surface Receptors 175 rRNAs Among Different Species Provide a Basis for Establishing
Evolutionary Relationships 226
8.4 Intracellular Receptors 177
8.5 Signal Transduction and Cellular Response via an 10.6 The Stages of Translation 227
Enzyme-Linked Receptor 178
Evolutionary Connections: Receptor Tyrosine Kinases Are Found CHAPTER 11
in Choanoflagellates and Animals 180
8.6 Signal Transduction and Cellular Response via a The Expression of Genetic Information via Genes II:
G-Protein-Coupled Receptor 181
Non-coding RNAs 231
8.7 Crosstalk Among Signal Transduction Pathways 184
11.1 Overview of Non-coding RNAs 232

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 Detailed Table of Contents xxiii

11.2 Role of Non-coding RNAs in Eukaryotic DNA 14.3 Meiosis 299

Replication 236
11.3 Effects of Non-coding RNAs on Chromatin Structure and
Transcription 238
11.4 Effects of Non-coding RNAs on Translation and mRNA
Degradation 239
Feature Investigation: Fire and Mello Showed That Double-
Stranded RNA Is More Potent Than Antisense RNA in Silencing
mRNA 239
11.5 Non-coding RNAs and Protein Sorting 243
11.6 Non-coding RNAs and Genome Defense 244
11.7 Roles of Non-coding RNAs in Human Disease and Plant
Health 246
CHAPTER 12
The Control of Genetic Information via Gene
Regulation 250
12.1 Overview of Gene Regulation 251
12.2 Regulation of Transcription in Bacteria 254
12.3 Regulation of Transcription in Eukaryotes: Roles of
Transcription Factors 258
Evolutionary Connections: Transcription in Archaea Is a
Simplified Version of Eukaryotic Transcription 261
12.4 Regulation of Transcription in Eukaryotes: Changes in
Chromatin Structure and DNA Methylation 262
12.5 Regulation of RNA Splicing and Translation in
Eukaryotes 266
Quantitative Analysis: Alternative Splicing Is More Prevalent in
Complex Eukaryotic Species 267
CHAPTER 13
Altering the Genetic Material: Mutation, DNA Repair,
and Cancer 271
13.1 Consequences of Mutations 272
13.2 Causes of Mutations 275
Feature Investigation: The Lederbergs Used Replica Plating to
Show That Mutations Are Random Events 275
Quantitative Analysis: Testing Methods Determine If an Agent Is
a Mutagen 278
13.3 DNA Repair 280
13.4 Cancer 282
CHAPTER 14
How Eukaryotic Cells Sort and Transmit Chromosomes:
Mitosis and Meiosis 289
14.1 The Eukaryotic Cell Cycle 290
14.2 Mitotic Cell Division 294
Evolutionary Connections: Cell Division in Bacteria Involves
FtsZ, a Protein Related to Eukaryotic Tubulin 297
14.4 Sexual Reproduction 304
Quantitative Analysis: Meiosis Enhances Genetic Diversity 305
14.5 Variation in Chromosome Structure and Number 307
CHAPTER 15
Transmission of Genetic Information from Parents to
Offspring I: Patterns That Follow Mendel’s Laws 313
15.1 Mendel’s Laws of Inheritance 314
Quantitative Analysis: A Punnett Square Is Used to Predict the
Outcome of Crosses 318
15.2 Chromosome Theory of Inheritance 320
15.3 Pedigree Analysis of Human Traits 323
15.4 Variations in Inheritance Patterns and Their Molecular
Basis 324
15.5 Sex Chromosomes and X-Linked Inheritance
Patterns 328
CHAPTER 16
Transmission of Genetic Information from Parents to
Offspring II: Epigenetics, Linkage, and Extranuclear
Inheritance 331
16.1 Overview of Epigenetics 332
16.2 Epigenetics: Genomic Imprinting 333
16.3 Epigenetics: X-Chromosome Inactivation 335
16.4 Epigenetics: Effects of Environmental Agents 338
16.5 Extranuclear Inheritance: Organelle Genomes 341
Evolutionary Connections: Chloroplast and Mitochondrial
Genomes Are Relatively Small but Contain Genes That Code
Important Proteins 341
16.6 Linkage of Genes on the Same Chromosome 344
Feature Investigation: Bateson and Punnett’s Crosses of Sweet Peas
Showed That Genes Do Not Always Assort Independently 344
Quantitative Analysis: A Chi Square Test Can Be Used to
Distinguish Between Linkage and Independent Assortment 347
CHAPTER 17
The Simpler Genetic Systems of Viruses,
Bacteria, and Archaea 351
17.1 General Properties of Viruses 352
17.2 Viral Reproductive Cycles 353
17.3 Genetic Properties of Bacteria and Archaea 360
17.4 Gene Transfer Between Prokaryotic Cells 364
Evolutionary Connections: Horizontal Gene Transfer Is the
Transfer of Genes Between the Same or Different Species 367

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 xxiv Detailed Table of Contents

20.4 Evo-Devo: Evolutionary Developmental Biology 439

CHAPTER 18 Evolutionary Connections: The Hox Genes Have Been Important
in the Evolution of a Variety of Body Patterns 441
Genetic Technologies: How Biologists Study
Genes and Genomes 370 CHAPTER 21
18.1 Gene Cloning 371
18.2 Genomics: Techniques for Studying and Altering How Biologists Classify Species and Study Their
Genomes 377 Evolutionary Relationships 445
18.3 Bacterial and Archaeal Genomes 384
18.4 Eukaryotic Genomes 386 21.1 Taxonomy 446
Evolutionary Connections: Every Species Is Placed into a
Evolutionary Connections: Gene Duplications Provide Additional
Taxonomic Hierarchy 446
Material for Genome Evolution, Sometimes Leading to the
Formation of Gene Families 387 21.2 Phylogenetic Trees 449
18.5 Repetitive Sequences and Transposable Elements 389 21.3 Cladistics 453
Quantitative Analysis: The Principle of Parsimony Is Used to
Choose from Among Possible Cladograms 455
UNIT IV Evolution 21.4 Molecular Clocks 456
21.5 Horizontal Gene Transfer 459

CHAPTER 22

The History of Life on Earth and Human

Evolution 463
22.1 The Fossil Record 464
Quantitative Analysis: Radioisotopes Provide a Way to Date
Fossils 465
Mark Dadswell/Getty Images 22.2 History of Life on Earth 466
Evolutionary Connections: The Origin of Eukaryotic Cells
Involved a Union Between Bacterial and Archaeal Cells 470
CHAPTER 19
22.3 Human Evolution 476
Evolutionary Connections: Comparing the Genomes of Humans
Evolution of Life I: How Populations Change from and Chimpanzees 477
Generation to Generation 397
19.1 Overview of Evolution 398 UNIT V Diversity
19.2 Evidence of Evolutionary Change 401
19.3 Genes in Populations 409
Evolutionary Connections: Genes Are Usually Polymorphic 409
Quantitative Analysis: The Hardy-Weinberg Equation Relates
Allele and Genotype Frequencies in a Population 410
19.4 Natural Selection 412
Feature Investigation: The Grants Observed Natural Selection in
Galápagos Finches 416
19.5 Genetic Drift 419
19.6 Migration and Nonrandom Mating 423

David M. Phillips/Science Source

CHAPTER 20 2 µm
Evolution of Life II: The Emergence of CHAPTER 23
New Species 426
Diversity of Microbial Life: Archaea, Bacteria, Protists,
20.1 Identification of Species 427
20.2 Reproductive Isolating Mechanisms 429
and Fungi 489
Feature Investigation: Podos Found That an Adaptation for Feeding 23.1 Introduction to Microorganisms 490
May Have Promoted Reproductive Isolation in Finches 432 23.2 Diversity and Ecological Importance of Archaea 493
20.3 Mechanisms of Speciation 434 23.3 Diversity and Ecological Importance of Bacteria 494

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 Detailed Table of Contents xxv

23.4 Diversity in Bacterial Cell Structure and

Metabolism 497 CHAPTER 27
23.5 Diversity and Ecological Importance of Protists 501
Evolutionary Connections: Primary Plastids and Primary Vertebrates: Fishes, Amphibians, Reptiles, and
Endosymbiosis 506 Mammals 596
23.6 Diversity and Ecological Importance of Fungi 511
27.1 Vertebrates: Chordates with a Backbone 597
23.7 Technological Applications of Microorganisms 518
27.2 Cyclostomes: Jawless Fishes 597
27.3 Gnathostomes: Jawed Vertebrates 600
CHAPTER 24 27.4 Tetrapods: Gnathostomes with Four Limbs 604
Feature Investigation: Davis and Colleagues Provided a Genetic-
Microbiomes: Microbial Systems on and Around Us 521 Developmental Explanation for Limb Length in Tetrapods 605
27.5 Amniotes: Tetrapods with a Desiccation-Resistant
24.1 Microbiomes: Diversity of Microbes and Functions 522 Egg 608
24.2 Microbiomes of Physical Systems 525 27.6 Mammals: Milk-Producing Amniotes 613
24.3 Host-Associated Microbiomes 527
Feature Investigation: Grieneisen and Colleagues Discovered That
Detecting Microbiome Heritability Requires Large Samples Taken Over UNIT VI Flowering Plants
Time 533
24.4 Engineering Animal and Plant Microbiomes 535

CHAPTER 25

Plant Evolution: How Plant Diversification Changed Planet

Earth 538
25.1 Ancestry and Diversity of Land Plants 539
25.2 An Evolutionary History of Land Plants 546
25.3 Diversity of Modern Gymnosperms 549 George Grall/Getty Images
25.4 Diversity of Modern Angiosperms 552
Evolutionary Connections: Flower Organs Evolved from Leaflike
Structures 554 CHAPTER 28
Feature Investigation: Hillig and Mahlberg Analyzed Secondary
Metabolites to Explore Species Diversification in the Genus An Introduction to Flowering Plant Form and
Cannabis 557
25.5 Human Influences on Angiosperm Diversification 560
Function 619
28.1 From Seed to Seed: The Life of a Flowering
CHAPTER 26 Plant 620
28.2 Plant Growth and Development 624
28.3 The Shoot System: Stem and Leaf Adaptations 628
Invertebrates: The Vast Array of Animal Life Feature Investigation: Lawren Sack and Colleagues Showed That
Without a Backbone 562 Palmate Venation Confers Tolerance of Leaf Vein Breakage 631
26.1 Characteristics of Animals 563 28.4 Root System Adaptations 636
26.2 Animal Classification 564
Evolutionary Connections: The Protostomes Consist of Two Major CHAPTER 29
Clades—the Ecdysozoa and the Lophotrochozoa 567
26.3 Ctenophores: The Earliest Animals 570 How Flowering Plants Sense and Interact with Their
26.4 Porifera: The Sponges 571
26.5 Cnidaria: Jellyfish and Other Radially Symmetric
Environments 640
Animals 572 29.1 Overview of Plant Behavioral Responses 641
26.6 Lophotrochozoa: The Flatworms, Rotifers, Bryozoans, 29.2 Plant Hormones 644
Brachiopods, Mollusks, and Annelids 574 Evolutionary Connections: Plant Gibberellin Responses Evolved
Quantitative Analysis: How Many Flukes? 576 in a Stepwise Manner 646
26.7 Ecdysozoa: The Nematodes and Arthropods 582 29.3 Plant Responses to Light 648
26.8 Deuterostomia: The Echinoderms and Chordates 590

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 xxvi Detailed Table of Contents

29.4 Plant Responses to Gravity and Touch 651 32.3 General Principles of Homeostasis 708
29.5 Plant Responses to Attack 653 32.4 Homeostatic Regulation of Body Temperature 711
32.5 Homeostasis of Internal Fluids 716
Feature Investigation: Cade and Colleagues Discovered Why
CHAPTER 30 Athletes’ Performances Wane on Hot Days 718

How Flowering Plants Obtain and Transport Water, Mineral

Nutrients, and Organic Compounds 658 CHAPTER 33
30.1 Plant Nutritional Requirements 659 Neuroscience I: The Structure, Function, and Evolution of
30.2 The Roles of Soil in Plant Nutrition 662 Nervous Systems 722
30.3 Transport at the Cellular Level 666
Quantitative Analysis: The Water Potential Equation Can Be Used 33.1 Cellular Components of Nervous Systems 723
to Predict Cellular Water Status 670 33.2 Electrical Properties of Neurons and the Resting
Evolutionary Connections: Plants Have Evolved Cellular Membrane Potential 726
Adaptations to Drought Stress 670 Quantitative Analysis: An Ion’s Equilibrium Potential Depends on
30.4 Plant Transport at the Tissue Level 671 Its Concentration Gradient 728
30.5 Long-Distance Transport in Plants 673 33.3 Generation and Transmission of Electrical Signals Along
Neurons 729
33.4 Communication at Synapses 734
CHAPTER 31
33.5 Evolution and Development of Nervous Systems 737
Evolutionary Connections: Animals Evolved Increasingly
How Flowering Plants Reproduce and Develop 682 Complex Nervous Systems 737
31.1 An Overview of Flowering Plant Reproduction 683 33.6 Structure and Function of the Nervous Systems of
31.2 Flower Production, Structure, and Development 687 Vertebrates 740
31.3 Male and Female Gametophytes and Double Feature Investigation: Gaser and Schlaug Discovered That the
Fertilization 690 Sizes of Certain Brain Structures Differ Between
Musicians and Nonmusicians 745
31.4 Embryo, Seed, Fruit, and Seedling Development 692
33.7 Impact on Public Health 747
31.5 Asexual Reproduction in Flowering Plants 696
Evolutionary Connections: Gene Expression Changes Explain the
Evolution of Plantlets in Kalanchoë 697 CHAPTER 34

UNIT VII Animals Neuroscience II: How Sensory Systems Allow Animals to
Interact with the Environment 752
34.1 Introduction to Sensation 753
34.2 Mechanoreception 754
34.3 Thermoreception and Nociception 759
34.4 Photoreception 760
Evolutionary Connections: Color Vision Is an Ancient Adaptation
in Animals 763
34.5 Chemoreception 767
Feature Investigation: Buck and Axel Discovered a Family
of Olfactory Receptor Proteins That Bind Specific Odor
anakondasp/Shutterstock Molecules 768
34.6 Impact on Public Health 770

CHAPTER 32
General Features of Animal Bodies, and Homeostasis as a
Key Principle of Animal Biology 701
32.1 Organization of Animal Bodies 702
Evolutionary Connections: Organ Development and Function Are
Controlled by Hox Genes 707
32.2 Relationship Between Structure and Function 707
CHAPTER 35
How Muscles and Skeletons Are Adaptations for
Movement, Support, and Protection 774
35.1 Types of Animal Skeletons 775
35.2 Skeletal Muscle Structure and the Mechanism of
Force Generation 777

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 Detailed Table of Contents xxvii

Evolutionary Connections: Myosins Are an Ancient and Diverse

Family of Proteins 780 CHAPTER 38
35.3 Types of Skeletal Muscle Fibers and Their
Functions 784 How Endocrine Systems Influence the Activities of
35.4 Impact on Public Health 786 All Other Organ Systems 857
38.1 Types of Hormones and Their Mechanisms of Action 858
CHAPTER 36 38.2 Links Between the Endocrine and Nervous Systems 861
38.3 Hormonal Control of Metabolism and Energy Balance 863
Circulatory and Respiratory Systems: Transporting Solutes Feature Investigation: Banting, Best, Collip, and MacLeod Were
and Exchanging Gases 789 the First to Isolate Active Insulin 867
38.4 Hormonal Control of Mineral Balance 870
36.1 Types of Circulatory Systems 790 Evolutionary Connections: Hormones and Receptors Evolved as
Evolutionary Connections: A Four-Chambered Heart Evolved Tightly Integrated Molecular Systems 872
from Simple Contractile Tubes 792 38.5 Hormonal Control of Growth and Development 874
36.2 The Composition of Blood 793 38.6 Hormonal Control of Reproduction 876
36.3 The Four-Chambered Vertebrate Heart and Its 38.7 Impact on Public Health 876
Function 795
36.4 Blood Vessels 799
36.5 Relationship Among Blood Pressure, Blood Flow, and CHAPTER 39
Resistance 802
Quantitative Analysis: Cardiac Output and Resistance Determine The Production of Offspring: Reproduction and
Blood Pressure 803 Development 879
36.6 Physical Properties of Gases 805
39.1 Overview of Sexual and Asexual Reproduction 880
36.7 Types of Respiratory Systems 806
Feature Investigation: Paland and Lynch Provided Evidence That
36.8 Structure and Function of the Mammalian Respiratory
Sexual Reproduction May Promote the Elimination of Harmful
System 809 Mutations in Populations 881
36.9 Mechanisms of Gas Transport in Blood 812 39.2 Gametogenesis and Fertilization 883
Quantitative Analysis: The Ability of Hemoglobin to Bind Oxygen
39.3 Human Reproductive Structure and Function 887
Is Affected by Factors Such as Temperature, CO2, and pH 813
39.4 Pregnancy and Birth in Mammals 892
36.10 Control of Ventilation 815
39.5 General Events of Embryonic Development 895
36.11 Impact on Public Health 816
39.6 Impact on Public Health 901

CHAPTER 37
CHAPTER 40
Digestive and Excretory Systems:
Immune Systems: How Animals Defend Against Pathogens
Maintaining Nutrient, Water, and Energy Balance
and Other Dangers 905
and Removing Waste 822
40.1 Types of Pathogens 906
37.1 Overview of Animal Nutrition 823 40.2 Innate Immunity 907
37.2 General Principles of Digestion and Absorption of Evolutionary Connections: Innate Immune Responses
Nutrients 826 Require Proteins That Recognize Features Common to Many
37.3 Vertebrate Digestive Systems 827 Pathogens 909
Evolutionary Connections: Evolution and Genetics Explain Feature Investigation: Lemaitre and Colleagues Identified an
Lactose Intolerance 833 Immune Function for Toll Protein in Drosophila 910
37.4 Nutrient Use and Storage 837 40.3 Adaptive Immunity in Vertebrates 912
37.5 Regulation of the Absorptive and Postabsorptive States in 40.4 Impact on Public Health 923
Vertebrates 839
37.6 Excretory Systems in Different Animal Groups 842
37.7 Structure and Function of the Mammalian Kidneys 845
CHAPTER 41
37.8 Impact on Public Health 851
Feature Investigation: Marshall, Warren, and Coworkers An Example of a System-Wide Response to a Challenge
Demonstrated a Link Between Bacterial Infection and Ulcers 852 to Homeostasis 927
41.1 Effects of Hemorrhage on Blood Pressure and Organ
Function 928

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 xxviii Detailed Table of Contents

41.2 The Rapid Phase of the Homeostatic Response to

Hemorrhage 930 CHAPTER 44
Evolutionary Connections: Baroreceptors May Have Evolved to
Minimize Increases in Blood Pressure in Vertebrates 932 Communities and Ecosystems: Ecological Organization
41.3 The Secondary Phase of the Homeostatic Response to at Large Scales 980
Hemorrhage 934
44.1 Patterns of Species Richness and Species Diversity 981
41.4 Impact on Public Health 938
Quantitative Analysis: Calculating Species Diversity 983
44.2 Species Richness and Community Stability 985
UNIT VIII Ecology 44.3 Succession: Community Change 986
44.4 Island Biogeography 989
Feature Investigation: Simberloff and Wilson’s Experiments
Tested the Predictions of the Equilibrium Model of Island
Biogeography 991
44.5 Food Webs and Energy Flow 993
44.6 Biomass Production in Ecosystems 997

CHAPTER 45

Biomes: How Climate Affects the Distribution of

Kirill KukhmarITAR-TASS News Agency/Alamy Stock Photo Species on Earth 1003
45.1 Climate and Its Relationship to Biological
Communities 1004
CHAPTER 42 45.2 Major Biomes 1010
Evolutionary Connections: Plate Tectonics and Biogeography
Behavioral Ecology: The Struggle to Find Food and Mates Help to Explain Species Distribution 1014

and to Pass on Genes 943

CHAPTER 46
42.1 The Influence of Genetics
and Learning on Behavior 944
Feature Investigation: Tinbergen’s Experiments Showed That
The Age of Humans 1017
Digger Wasps Learn the Positions of Landmarks to Find Their 46.1 Human Population Growth 1018
Nests 945 46.2 Global Warming and Climate Change 1021
42.2 Communication 948 46.3 Pollution and Human Influences on Biogeochemical
42.3 Living in Groups and Game Theory 950 Cycles 1024
Quantitative Analysis: Game Theory Establishes Whether Feature Investigation: Stiling and Drake’s Experiments with
Individuals Fight or Flee 952 Elevated CO2 Showed an Increase in Plant Growth but a Decrease in
42.4 Altruism 953 Herbivory 1025
42.5 Mating Systems 956 46.4 Pollution and Biomagnification 1030
46.5 Habitat Destruction 1032
CHAPTER 43 46.6 Overexploitation 1035
46.7 Invasive Species 1038
Population Growth and Species Interactions 960
43.1 Measuring Population Size and Density 961 CHAPTER 47
Quantitative Analysis: Mark-Recapture Can Be Used to Estimate
Population Size 962 Biodiversity and Conservation Biology 1043
43.2 Demography 963 47.1 Genetic, Species, and Ecosystem Diversity 1044
43.3 How Populations Grow 966 47.2 Value of Biodiversity to Human Welfare 1045
43.4 Species Interactions 968 47.3 Biodiversity and Ecosystem Function 1047
Evolutionary Connections: Organisms Have Evolved Many
47.4 Conservation Strategies 1049
Defenses Against Natural Enemies 971
43.5 The Spread of Pathogens 975 Appendix A Periodic Table of the Elements A-1
Appendix B Answers to In-Chapter and End-of-Chapter Questions B-1
Index I-1

bro23659_fm_i-xxviii.indd 28 30/09/22 8:58 PM


An Introduction to Biology
Geoff Sperring/Shutterstock
Biology is the study of life. The diverse
forms of life found on Earth provide
biologists with an amazing array of
organisms to study. In many cases, the
investigation of living things leads to
discoveries with far-reaching benefits.
Certain ancient civilizations, such as
the Greeks, Romans, and Egyptians,
discovered that the bark of the white
willow tree (Salix alba) could be used
to fight fever. Chemists determined that
willow bark contains a substance called
salicylic acid, which led to the develop-
ment of the related compound acetyl-
salicylic acid, more commonly known
as aspirin (Figure 1.1). Today, aspirin is
taken for fever and pain relief.
As a more recent example, re-
searchers determined that the venom
from certain poisonous snakes con-
tains a chemical that lowers blood
pressure in humans. By analyzing
that chemical, scientists have devel-
oped drugs called ACE inhibitors for
the treatment of high blood pressure
(Figure 1.2).
These are just a couple of the
many discoveries that make biology
bro23659_ch01_001-019.indd 1
1
A tuskless female elephant with her offspring.
As discussed in Section 1.2, ivory poaching has
increased the frequency of tuskless elephants in
Africa.
Chapter Outline
1.1 Principles of Biology and the
Levels of Biological Organization
1.2 Biological Evolution
1.3 Classification of Living Things
1.4 Biology as a Scientific Discipline
Assessing Your Knowledge and Skills
an intriguing discipline. The study of
life not only reveals the fascinating
characteristics of living species but
also leads to the development of med-
icines that benefit the lives of people.
To make new discoveries, biolo-
gists view life from many different
perspectives. What is the composi-
tion of living things? How is life orga-
nized? How do organisms reproduce?
Sometimes the questions posed by
biologists are fundamental and even
philosophical in nature. How did liv-
ing organisms originate? Can we live
forever? What is the physical basis for
memory? Can we save endangered
species?
Future biologists will continue to
make important advances. Biologists
are scientific explorers looking for
answers to some of life’s most endur-
ing mysteries. Unraveling these mys-
teries presents exciting challenges
to the best and brightest minds.
The rewards of a career in biology
include the excitement of forging into
uncharted territory, the thrill of mak-
ing discoveries that can improve the
23/09/22 10:04 AM
 2 CHAPTER 1
health and lives of people, and the goal of trying to preserve
the environment and protect endangered species. For these
and many other compelling reasons, students seeking chal-
lenging and rewarding careers may wish to choose biology
as a lifelong pursuit.
In this chapter, we will begin our survey of biology by
examining the basic principles that underlie the characteristics
O C
OH
O ACE inhibitor (Lotensin)
C
Aspirin
O CH3
Figure 1.1 The white willow (Salix alba) and aspirin. Modern aspirin,
acetylsalicylic acid, was developed after analysis of a chemical
found in the bark of the white willow tree.
Blickwinkel/Hartl/Alamy Stock Photo
1.1 Principles of Biology and the Levels
of Biological Organization
Learning Outcomes
1. Describe the principles of biology.
2. Explain how life can be viewed at different levels of
biological complexity.
Because biology is the study of life, a good way to begin a biology
textbook is to distinguish living organisms from nonliving objects. At
first, the distinction might seem obvious. A person is alive, but a rock
is not. However, the distinction between living and nonliving may
seem less obvious when we consider microscopic entities. Is a bac-
terium alive? What about a virus or a chromosome? In this section,
we will examine the principles that underlie the characteristics of all
forms of life and explore other broad principles in biology. We will
then consider the levels of organization that biologists study, ranging
from atoms and small molecules to very large geographic areas.
The Study of Life Has Revealed a Set
of Unifying Principles
In the course of studying a vast number of species, biologists have
learned that a set of principles applies to all fields of biology. Twelve
broad principles are described in Figure 1.3. The first eight principles
are often used as criteria for defining the basic features of life. You
bro23659_ch01_001-019.indd 2
of all living organisms. We will then take a deeper look at the
process of evolution and how it explains the unity and diver-
sity that we observe among living and extinct species. Finally,
we will explore the general approaches that scientists follow
when making new discoveries.
N
O O OCH2CH3
CH2COOH
Figure 1.2 The Brazilian arrowhead viper and an inhibitor of high
blood pressure. Originally found in the venom of the Brazilian
arrowhead viper (Bothrops jararaca/jararacussa), angiotensin-
converting enzyme (ACE) inhibitors, including benazepril
(Lotensin, chemical structure shown here), are commonly used to
treat high blood pressure.
Francois Gohier/Science Source
will see these 12 principles at many points as you progress through
this textbook. In particular, we will draw your attention to them at the
beginning of each unit, and we will refer to them within particular
figures in Chapters 2 through 47. It should be noted that the princi-
ples of biology are also governed by the laws of chemistry and phys-
ics, which are discussed in Chapters 2, 3, and 6.
Principle 1: Cells are the simplest units of life. The term
organism can be applied to all living things. Organisms maintain an
internal order that is separated from the environment (Figure 1.3a).
The simplest unit of such organization is the cell, which we will exam-
ine in Unit II. One of the foundations of biology is the cell theory,
which states that (1) all living things are composed of one or more
cells, (2) cells are the smallest units of life, and (3) new cells come
from pre-existing cells by cell division. Some organisms are unicellu-
lar and are composed of one cell, whereas others, such as plants and
animals, are multicellular and contain many cells. In plants and ani-
mals, each cell has an internal order, and the cells within the organism
have specific arrangements and functions.
Principle 2: Living organisms use energy. The maintenance of
organization requires energy. Therefore, all living organisms acquire
energy from the environment and use that energy to maintain their
internal order. Cells carry out a variety of chemical reactions that are
responsible for the breakdown of energy-yielding nutrients. Such reac-
tions often release energy in a process called cellular respiration.
The energy may also be used to synthesize the components that
make up individual cells and living organisms. Chemical reactions
23/09/22 10:04 AM
 (a) (Principle 1) Cells: (g) ( Principle 7) Structure and Function:
Cells are the simplest units of The structural features of
life. Organisms maintain an living organisms determine
internal order. The simplest their functions. In the example
unit of organization is the cell. seen here, webbed feet (on
Yeast cells are shown here. ducks) function as paddles for
13.7 μm swimming. Nonwebbed feet
(on chickens) function better for
walking on the ground.
(b) ( Principle 2) Energy and Matter:
Living organisms use energy (h) ( Principle 8) Emergent Properties:
and are composed of matter. New properties of life emerge
Organisms need energy to from complex interactions. Our
maintain internal order. These ability to see is an emergent
algae harness light energy via property due to interactions
photosynthesis. Energy is used among many types of cells in
in chemical reactions collectively the eye and neurons that send
known as metabolism. signals to the brain.

(c) (Principle 3) Environmental Interactions: (i) ( Principle 9) Experimentation:

Living organisms interact with Biology is an experimental
their environment. Organisms science. The discoveries
respond to environmental of biology are made via
changes. This plant is growing experimentation, which leads to
toward the light. theories and biological principles.

(d) ( Principle 4) Homeostasis: Male gametes (j) ( Principle 10) Quantitative Analysis:
Living organisms maintain T t Biology is a quantitative science.

Female gametes
homeostasis. Organisms regulate their Biologists analyze data in
cells and bodies, maintaining relatively T TT Tt a quantitative way and use
stable internal conditions, a process mathematical approaches to
called homeostasis. For example, make predictions. For example, a
t Tt tt
this bird maintains its internal body Punnett square is used to predict
temperature on a cold day. the outcome of genetic crosses.
(e) ( Principle 5) Genetic Material: (k) ( Principle 11) Modeling:
The genetic material provides a Biologists use models and
blueprint that allows organisms to grow, simulations to test experimental
develop, and reproduce. To sustain life predictions and convey their
over many generations, organisms must ideas. For example, molecular
reproduce. Due to the transmission biologists propose models
of genetic material, offspring tend to that describe the structures of
have traits like their parents. proteins.
(f) ( Principle 6) Evolution: (l) ( Principle 12) Biology and Society:
Populations of organisms evolve Biology affects our society. Many
from one generation to the next and discoveries in biology have had
are related by an evolutionary history. major effects on our society. For
Populations of organisms change example, biologists developed
over the course of many generations. Bt-corn, which is resistant to
Evolution results in traits that promote insect pests and is widely
survival and reproductive success. planted by farmers.
Evolution results in new species that
are related to each other.

Figure 1.3 Twelve principles of biology. The first eight principles are often used as criteria for defining the basic features of life. Note: The
12 principles described here were modeled after the core concepts and core competencies described in Vision and Change in Undergraduate
Biology, a report that was published in 2009 and organized by the American Association for the Advancement of Science. Vision and Change
proposed several core concepts and core competencies (see http://visionandchange.org). We have slightly modified and expanded them
into 12 principles to make them more accessible to beginning biology students. The five Vision and Change core concepts are related to
our principles in the following manner: (1) evolution (principle 6); (2) structure and function (principle 7); (3) information flow, exchange, and
storage (principle 5); (4) pathways and transformations of energy and matter (principles 2, 3, and 4); and (5) systems (principles 1 and 8). The
last four principles are modeled after four core competencies (that is, core skills) described in Vision and Change: ability to apply the process
of science (principle 9); ability to use quantitative reasoning (principle 10); ability to use models and simulations (principle 11); and ability to
understand the relationship between science and society (principle 12).
(a) Steve Gschmeissner/Science Photo Library/Getty Images; (b) Robert La Salle/Aqua-Photo/Alamy Stock Photo; (c) Martin Shields/Alamy Stock Photo;
(d) Cliff Keeler/Alamy Stock Photo; (e) Frank Krahmer/Getty Images; (f) Luiz Kagiyama/Shutterstock; (g) G.K. & Vikki Hart/Getty Images; (h) Maria Teijeiro/Getty
Images; (i) Corbis/SuperStock; (k) ©Daniel Gage, University of Connecticut; (l) Bill Barksdale/age fotostock

BioConnections:  Look ahead to Figure 4.15. Which of these principles is this figure emphasizing?

bro23659_ch01_001-019.indd 3 23/09/22 10:06 AM

 4 CHAPTER 1
involved with the breakdown and synthesis of cellular molecules are
collectively known as metabolism. Plants, algae, and certain bac-
teria directly harness light energy to produce their own nutrients in
the process of photosynthesis (Figure 1.3b). They are the primary
producers of food on Earth. In contrast, some organisms, such as ani-
mals and fungi, are consumers—they must use other organisms or the
products of other organisms as food to obtain energy.
Principle 3: Living organisms interact with their environment.
To survive, living organisms must interact with their environment,
which includes other organisms they may encounter. All organisms
must respond to environmental changes. For example, in the winter,
many species of mammals develop a thicker coat of fur that protects
them from the cold temperatures. Plants respond to changes in the
angle of the Sun. If you place a plant next to a lamp, it will grow
toward the light (Figure 1.3c).
Principle 4: Living organisms maintain homeostasis. Although
life is a dynamic process, living cells and organisms regulate their cells
and bodies to maintain relatively stable internal conditions, a process
called homeostasis (from the Greek, meaning “to stay the same”).
The degree to which homeostasis is achieved varies among different
organisms. For example, most mammals and birds maintain a relatively
stable body temperature in spite of changing environmental tempera-
tures (Figure 1.3d), whereas reptiles and amphibians tolerate a wider
fluctuation in body temperature. By comparison, all organisms contin-
ually regulate their cellular metabolism so that nutrient molecules are
used at an appropriate rate and new cellular components are synthe-
sized when they are needed.
Principle 5: The genetic material provides a blueprint that
allows organisms to grow, develop, and reproduce. All
living organisms grow and develop specific traits. To sustain life
from one generation to the next, organisms must also reproduce,
or generate offspring (Figure 1.3e). A key feature of reproduction is
that offspring tend to have characteristics that greatly resemble those
of their parent(s). How is this possible? All living organisms contain
genetic material composed of deoxyribonucleic acid (DNA), which
provides a blueprint for the organization, development, and func-
tion of living things. During reproduction, a copy of this blueprint is
transmitted from parent to offspring. DNA is heritable, which means
that offspring inherit DNA from their parents.
As discussed in Unit III, genes, which are segments of DNA,
govern the characteristics, or traits, of organisms. Most genes are tran-
scribed into a type of ribonucleic acid (RNA) molecule called mes-
senger RNA (mRNA), which is then translated into a polypeptide
with a specific amino acid sequence. A protein is composed of one
or more polypeptides. The structures and functions of proteins are
largely responsible for the traits of living organisms.
Principle 6: Populations of organisms evolve from one gener-
ation to the next and are related by an evolutionary history.
The first five characteristics of life, which we have just considered,
apply to individual organisms over the short run. Over the long run,
another universal characteristic of life is biological evolution, or sim-
ply evolution, which is a heritable change in a population of organisms
from generation to generation. As a result of evolution, populations
become better adapted to the environment in which they live. For
bro23659_ch01_001-019.indd 4
example, the long snout of an anteater is an adaptation that enhances its
ability to obtain food, namely ants, from hard-to-reach places (Figure
1.3f). Over the course of many generations, the fossil record indicates
that the long snout occurred via biological evolution in which modern
anteaters evolved from populations of organisms with shorter snouts.
Evidence from a variety of sources, including the fossil record
and DNA sequences, also indicates that all organisms on Earth share
a common ancestry. For example, the different species of mammals
that currently exist shared a common ancestor in the past, which was
also a mammal. We will discuss evolutionary relationships further in
Section 1.2. In many chapters of this textbook, you will find a subsec-
tion called Evolutionary Connections, which focuses on the evolu-
tionary aspects of the chapter’s material.
Principle 7: The structural features of living organisms deter-
mine their function. In addition to the preceding six characteristics
of life, biologists have identified other principles that are important in all
fields of biology. The principle that structure determines function per-
tains to very tiny biological molecules as well as very large biological
structures. For example, at the microscopic level, a cellular protein called
actin naturally assembles into structures that are long filaments. The
function of these filaments is to provide support and shape to cells. At the
macroscopic level, let’s consider the feet of different birds (Figure 1.3g).
Aquatic birds have webbed feet that function as paddles for swimming.
By comparison, the feet of nonaquatic birds are not webbed and are bet-
ter adapted for grasping food, perching on branches, and running along
the ground. In this case, the structure of a bird’s feet, webbed versus non-
webbed, is a critical feature that affects their function.
Principle 8: New properties of life emerge from complex inter-
actions. In biology, when individual components in an organism
interact with each other or with the external environment to create
novel structures and functions, the resulting characteristics are called
emergent properties. For example, the human eye is composed of
many different types of cells that are organized to sense incoming light
and transmit signals to the brain (Figure 1.3h). Our ability to see is an
emergent property of this complex arrangement of different cell types.
Principle 9: Biology is an experimental science. We now turn
to principles that involve skills that are used in the study of life. Biology
is an inquiry process. In other words, biologists are curious about the
characteristics of living organisms and ask questions about those charac-
teristics. For example, a cell biologist may wonder why a cell produces a
specific protein when it is confronted with high temperature. An ecolo-
gist may ask herself why a particular bird eats insects in the summer and
seeds in the winter. To answer such questions, biologists typically gather
additional information and ultimately form a hypothesis, which is a pro-
posed explanation for a natural phenomenon. The next stage is to design
one or more experiments to test the validity of the hypothesis (Figure
1.3i). Like evolution, experimentation is such a key aspect of biology that
most chapters of this textbook include a Feature Investigation—an actual
research study that showcases the experimental approach.
Principle 10: Biology is a quantitative science. Biologists use
quantitative reasoning in a variety of ways, as you will see throughout
this textbook. For example, the results of experiments are often ana-
lyzed using statistics. Also, biologists may use mathematical models
23/09/22 10:06 AM

to make predictions. In Chapter 15, you will see how a Punnett square
is used to predict the outcome of crosses (Figure 1.3j). In Chapter 43,
equations are used to predict different patterns of population growth
over the course of several generations. Many chapters have a sub-
section that emphasizes quantitative reasoning, an important skill for
careers in science and medicine. In these subsections, the quantitative
nature of a given topic is described, and then students are asked to
solve a problem related to that topic.
Principle 11: Biologists use models and simulations to
test experimental predictions and convey their ideas. A
scientific model, or simply a model, is a conceptual, mathematical,
or physical depiction of a real-world phenomenon. Models may con-
vey different aspects of life. For example, Figure 1.3k shows a model
for the molecular structure of a protein. Why are models useful?
Models are often used as a working hypothesis that helps research-
ers visualize or explain biological phenomena. Such models form the
basis for conducting further experiments. Models are evaluated for
their consistency with experimental data, which enables researchers
to accept, reject, or refine them. Another useful aspect of models is
that they allow biologists to convey their ideas in a relatively simple
way. Most of the figures in this textbook are models that have been
derived from the interpretation of experimental data.
Principle 12: Biology affects our society. The influence of biol-
ogy is not confined to textbooks and classrooms. The work of biol-
ogists has far-reaching effects in our society. For example, biologists
have discovered drugs that are used to treat many different human dis-
eases. Likewise, biologists have created technologies that have many
uses. Examples include the use of microorganisms to make medical
products, such as human insulin, and the genetic engineering of crops
to make them resistant to particular types of insect pests (Figure 1.3l).
Living Organisms Are Studied
at Different Levels of Organization
The organization of living organisms can be analyzed at different lev-
els of biological complexity, starting with the smallest level of organ-
ization and progressing to levels that are physically much larger and
more complex. Figure 1.4 depicts a scientist’s view of the levels of
biological organization.
1. Atoms. An atom is the smallest unit of an element that has the
chemical properties of the element. All matter is composed of
atoms.
2. Molecules and macromolecules. As discussed in Unit I, atoms bond
with each other to form molecules. Many smaller molecules can
bond together to form a large polymer called a macromolecule.
Carbohydrates, proteins, and nucleic acids (DNA and RNA) are
important macromolecules found in living organisms.
3. Cells. Molecules and macromolecules associate with each other
to form larger structures such as cells. A cell is surrounded
by a membrane and contains a variety of molecules and
macromolecules. As noted earlier, a cell is the simplest unit of life.
4. Tissues. In multicellular organisms such as plants and animals,
many cells of the same type associate with each other to form
tissues. An example is muscle tissue.
bro23659_ch01_001-019.indd 5
AN INTRODUCTION TO BIOLOGY 5
5. Organs. In complex multicellular organisms, an organ is
composed of two or more types of tissue and carries out a
particular function. For example, the heart is composed of several
types of tissue, including muscle, nervous, and connective tissue.
6. Organ systems. In multicellular species, organs are typically
a part of a larger, interacting system. In animals, such as birds
and mammals, the heart is a part of the circulatory system that
pumps blood throughout the body.
7. Organism. All living things can be called organisms.
Biologists classify organisms as belonging to a particular
species, which is a related group of organisms that share a
distinctive form and set of attributes in nature. The members of
the same species are closely related genetically. In Units VI and
VII, we will examine plants and animals at the levels of cells,
tissues, organs, organ systems, and complete organisms.
8. Population. A group of organisms of the same species that
occupy the same environment is called a population.
9. Community. A biological community is an assemblage of
populations of different species that live in the same environment.
The types of species found in a community are determined by the
environment and by the interactions of the species with each other.
10. Ecosystem. Researchers may extend their work beyond
living organisms and also study the physical environment.
Ecologists analyze ecosystems, which are formed by the
interactions between a community of organisms and its physical
environment. Unit VIII considers biological organization from
populations to ecosystems.
11. Biosphere. The biosphere includes all of the places on the
Earth where living organisms exist. Life is found in the air, in
bodies of water, on the land, and in the soil.
1.1 Reviewing the Concepts
• Biology is the study of life. Discoveries in biology help us understand
how life exists, and they have many practical applications, such as
the development of drugs to treat human diseases (Figures 1.1, 1.2).
• Eight principles underlie the characteristics that are common to
all forms of life (Figure 1.3). All living things (1) are composed of
cells as their simplest unit; (2) use energy; (3) interact with their
environment; (4) maintain homeostasis; and (5) have genetic
material for the development of traits and for reproduction. Also,
(6) populations of organisms evolve from one generation to the
next and are connected by an evolutionary history (Figure 1.3).
Additional important principles of biology that apply to living
organisms are that (7) structure determines function and (8) new
properties emerge from complex interactions.
• Biology also involves certain skills, including (9) biology is an
experimental science; (10) biology is a quantitative science; (11)
models and simulations are used to test experimental predic-
tions and convey ideas; and (12) biology influences our society.
• Living organisms can be viewed at different levels of biological
organization: atoms, molecules and macromolecules, cells, tis-
sues, organs, organ systems, organisms, populations, communi-
ties, ecosystems, and the biosphere (Figure 1.4).
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 6 CHAPTER 1

1 Atoms 2 Molecules and

macromolecules

3 Cells

4 Tissues
5 Organs
6 Organ
systems

7 Organism

11 Biosphere

8 Population

10 Ecosystem
9 Community

Figure 1.4 The levels of biological organization.

Concept Check:  At which level of biological organization would you place a herd of buffalo?

d. Populations of organisms evolve from one generation to the next.

1.1 Testing Your Knowledge e. All of the above are correct.
1. The wing of a bird, the wing of an insect, and the wing of a bat 2. Which of the following is the most complex level of biological
have similar shapes. Which principle of biology does this obser- organization?
vation pertain to? a. organism
a. Living organisms use energy. b. tissue
b. Living organisms maintain homeostasis. c. community
c. Structure determines function. d. population

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1.2 Biological Evolution
Learning Outcomes
1. Explain the two basic mechanisms by which evolutionary
change occurs: vertical descent with mutation and
horizontal gene transfer.
2. Describe how evolution accounts for unity and diversity in
biology.
Unity and diversity are two words often used to describe the living
world. As we have seen, all modern forms of life display a common
set of characteristics that distinguish them from nonliving objects. In
this section, we will explore how this unity of common traits is rooted
in the phenomenon of biological evolution. Life on Earth is united by
an evolutionary past in which modern organisms have evolved from
populations of pre-existing organisms. This unity is a core principle
of biology.
However, evolutionary unity does not mean that organisms are
exactly alike. The Earth has many different types of environments,
ranging from tropical rain forests to salty oceans, from hot, dry des-
erts to cold mountaintops. Diverse forms of life have evolved in ways
that help them prosper in the different environments the Earth has to
offer. In this and the following section, we will begin to examine the
unity and diversity that exist within the biological world.
Modern Forms of Life Are Connected
by an Evolutionary History
Life began on Earth as primitive cells about 3.5–4 billion years ago
(bya). Since that time, populations of living organisms have undergone
evolutionary changes that ultimately gave rise to the species we see
today. Understanding the evolutionary history of species can provide
key insights into an organism’s structure and function because evolu-
tionary change involves modifications of characteristics in pre-existing
populations. Over long periods of time, populations may change so that
structures with a particular function become modified to serve a new
function. For example, the wing of a bat is used for flying, and the flip-
per of a dolphin is used for swimming. Evidence from the fossil record
indicates that both structures were modified from a front limb that was
used for walking in a pre-existing ancestor (Figure 1.5).
Evolutionary change occurs by two mechanisms: vertical descent
with mutation and horizontal gene transfer. Let’s take a brief look at
each of these mechanisms.
Vertical Descent with Mutation Is the Primary
Way That Evolution Occurs
The traditional way to study evolution is to examine a progression of
changes in a series of related ancestral species. Such a series is called
a lineage. Biologists depict such evolutionary change in a diagram
like the one shown in Figure 1.6, which shows a portion of the line-
age that gave rise to modern horses. In this mechanism of evolution,
called vertical evolution, new species evolve from pre-existing ones
by the accumulation of mutations, which are heritable changes in
the genetic material of organisms.
bro23659_ch01_001-019.indd 7
AN INTRODUCTION TO BIOLOGY 7
Ancestral
limb
Modification over time
Dolphin
Bat wing flipper
Figure 1.5 An example of a modification that has occurred as a result of
biological evolution. The wing of a bat and the flipper of a dolphin are
modifications of a limb that was used for walking in a pre-existing
ancestor.
Concept Check:  Among mammals, give two examples of
how the tail has been modified and has different purposes.
But why would some mutations accumulate in a population and
eventually change the characteristics of an entire species? One rea-
son is that a mutation may alter the traits of organisms in a way that
increases their chances of survival and reproduction. When a muta-
tion causes such a beneficial change, the frequency of the mutation
may increase in a population from one generation to the next, a pro-
cess called natural selection. This topic is discussed in Units IV
and V. Evolution also involves the accumulation of neutral changes
that do not benefit or harm a species, and it sometimes involves rare
changes that may be harmful.
With regard to the horses shown in Figure 1.6, the fossil record
has revealed adaptive changes in various traits such as size and
tooth morphology. The first horses were the size of dogs, whereas
modern horses typically weigh more than a half ton. The teeth of
Hyracotherium were relatively small compared with those of mod-
ern horses. Over the course of millions of years, horses’ teeth have
increased in size, and a complex pattern of ridges has developed on
the molars.
How do evolutionary biologists explain these changes in horse
characteristics? They can be attributed to natural selection in which
changing global climates favored the survival and reproduction of
horses with certain types of traits. Over North America, where much
of horse evolution occurred, large areas changed from dense forests
to grasslands. Horses with genetic variation that made them larger
were more likely to escape predators and be able to travel greater
distances in search of food. The changes seen in horses’ teeth are con-
sistent with a dietary shift from eating tender leaves to eating grasses
and other types of vegetation that are more abrasive and require more
chewing.
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 8 CHAPTER 1

0
Hippidium and
Equus other genera
5
Nannippus
Styohipparion
Hipparion Neohipparion Pliohippus
10
Sinohippus Megahippus
Calippus
Archaeohippus

Merychippus
20 Anchitherium
Hypohippus
Millions of years ago (mya)

Parahippus

Miohippus

40 Mesohippus

Paleotherium
Epihippus
Propalaeotherium

Pachynolophus Orohippus

55
Hyracotherium

Figure 1.6 An example of vertical evolution: the horse lineage. This diagram shows the horse lineage. The highlighted branch gave rise to the
modern horse genus (Equus), which evolved from ancestors that were much smaller. The vertical evolution shown here occurred due to the
accumulation of mutations that altered traits of the species.

Concept Check:  What is the relationship between vertical evolution and natural selection?

problem. This can occur by the transfer of an antibiotic-resistance

Horizontal Gene Transfer Can Also Promote
Evolutionary Change
The most common way for genes to be transferred is in a vertical
manner. This can involve the transfer of genetic material from a
mother cell to daughter cells, or it can occur via gametes—sperm and
egg—that unite to form a new organism. However, as discussed in
Chapter 21, genes are sometimes transferred between organisms by
horizontal gene transfer (HGT)—a process in which an organism
incorporates genetic material from another organism without being
the offspring of that organism.
In some cases, HGT can occur between members of differ-
ent species. For example, you may have heard in the news media
that resistance to antibiotics among bacteria is a persistent medical
gene from one bacterial species to another via HGT (look ahead to
Figure 17.12).
Traditionally, biologists have described evolution using dia-
grams such as that in Figure 1.6, which depict the vertical evolution
of species over a long timescale. In this view, all living organisms
evolved from a common ancestor, resulting in a “tree of life” that
could describe the evolution that gave rise to all modern species. Now
that we understand the great importance of HGT in the evolution
of life on Earth, biologists have re-evaluated the way evolution has
occurred over time. Rather than a tree of life, a more appropriate way
to view the unity of living organisms is a “web of life” (as discussed
in Chapter 21; look ahead to Figure 21.14), which accounts for both
vertical descent with mutation and HGT.

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EVOLUTIONARY CONNECTIONS
The Frequency of the Tuskless Elephant Appears to Be
Increasing in Elephant Populations Due to Poaching
Evolution explains the unity and diversity among different spe-
cies. Throughout this textbook, subsections entitled Evolutionary
Connections will relate a specific topic to evolution. As described
earlier in Figure 1.3a–f, living organisms have a unifying set of
features because all species evolved from a common set of ancestors.
Evolution via natural selection results in adaptations that favor repro-
ductive success, which explains the diversity we see among different
species. An adaptation is a characteristic in a species that is the
result of natural selection. Some adaptations may affect the ability of
individuals to survive to reproductive age in their native environment.
For example, the warm coat of polar bears allows them to withstand
the cold arctic temperatures. Other adaptations, such as internal ges-
tation in mammals, may directly affect the ability of individuals to
successfully reproduce. In our Evolutionary Connections subsections,
we will often explore the characteristics of species to appreciate how
evolution has resulted in various types of adaptations.
In addition to natural selection, the activities of humans can also
alter the frequency of traits in populations of animals, plants, and
microorganisms. As discussed in Chapter 19, selective breeding
has altered the traits of domesticated species, including dogs and
agricultural crops (look ahead to Figure 19.7). Human activities have
also affected the frequencies of traits in wild species. For example,
the widespread use of antibiotics has caused an increase in the fre-
quency of antibiotic-resistant strains of bacteria. Another striking
example is the increase in frequency of the tuskless elephant in Africa
(Figure 1.7). As its name suggests, the tuskless elephant lacks the tusks
that are usually present in wild elephants. In their native environment,
a strong pair of tusks is an advantage for elephants, allowing them
to dig for water, to strip bark for food, and for males to compete for
mates. On the other hand, tusks can be a liability due to poaching.
Figure 1.7 A comparison of a tuskless (left) and a tusked (right) elephant.
Volker Schlichting/123RF
bro23659_ch01_001-019.indd 9
AN INTRODUCTION TO BIOLOGY 9
Elephants are often killed for their tusks, due to the value of ivory,
though such a practice is illegal. However, the tuskless elephants are
undesirable to poachers.
In a well-protected population, the frequency of tuskless ele-
phants is typically less than 2%, but that percentage can dramatically
increase due to poaching. For example, civil war and poaching in
Mozambique have led to a greater proportion of tuskless elephants.
During the conflict from 1977 to 1992, fighters on both sides slaugh-
tered elephants for ivory to finance war efforts. In the region that
is now Gorongosa National Park, about 90% of the elephants were
killed. Among the survivors, about 50% of the females were tuskless,
whereas before the civil war, less than 20% lacked tusks. In 2021,
mathematical modeling by American evolutionary biologist Shane
Campbell-Staton and colleagues indicated that this shift is the result
of hunting pressure: The selective killing of tusked elephants has led
to the birth of more tuskless offspring.
All tuskless elephants are females. How do we explain this
observation? The gene that determines the tuskless trait is carried
on the X chromosome. It is found in two alleles, tuskless, which
we will designate as T, and normal tusks, designated t. The T allele
is dominant. Female mammals carry two X chromosomes (XX),
whereas males carry one X chromosome and one Y chromosome
(XY). If a female elephant is Tt, she will be tuskless, because the
tuskless allele (T) is dominant. Males, however, inherit only one
X chromosome so they cannot be Tt. If a male inherits the t allele,
he will have tusks. However, the T allele in males is lethal during
embryonic development. Therefore, adult male elephants cannot be
tuskless.
Campbell-Staton and colleagues have also attempted to identify
the gene(s) that is/are involved in the tuskless trait. A particularly
interesting candidate is a gene called AMELX, which is carried on the
X chromosome in elephants and humans. In humans, a deletion in the
X chromosome that carries the AMELX gene is lethal in males and
causes amelogenesis imperfecta in females. Such females, who carry
an X chromosome with the deletion and the other X chromosome
without the deletion, display several facial abnormalities, including
the absence of teeth called maxillary lateral incisors. In elephants, the
maxillary lateral incisors become tusks.
1.2 Reviewing the Concepts
• Changes in species often occur as a result of modification of pre-
existing structures (Figure 1.5).
• During vertical evolution, mutations in a lineage alter the char-
acteristics of species. Individuals with greater reproductive suc-
cess are more likely to contribute certain mutations to future
generations, a process known as natural selection. Over the long
run, this process alters species and may produce new species
(Figure 1.6).
• Horizontal gene transfer is the transfer of genes from one organ-
ism to another organism that is not its offspring. It may involve
the transfer of genes between different species. Along with
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 10 CHAPTER 1
vertical descent with mutation, it is an important process in bio-
logical evolution, producing a web of life.
• Humans have influenced the frequency of traits in domesticated
and wild species, including the frequency of the tuskless trait in
elephant populations (Figure 1.7)
1.2 Testing Your Knowledge
1. Which of the following is an example of horizontal gene
transfer?
a. the transmission of genes from a mother cell to a daughter
cell during cell division
b. the transmission of a mutant gene from a father to his
daughter
c. the transfer of an antibiotic-resistance gene from one
bacterial species to another
d. all of the above
e. both a and b
1.3 Classification of Living Things
Learning Outcome
1. Outline how organisms are classified (taxonomy).
As biologists discover new species, they try to place them into groups
based on their evolutionary history. This is a difficult task because
researchers estimate that the Earth has between 5 and 50 million dif-
ferent species! The rationale for categorization is usually based on
vertical evolution. The field of biology that is concerned with the
grouping and classification of species is termed taxonomy.
Why is taxonomy useful? First, taxonomy allows us to appreci-
ate the amazing diversity of life on Earth. Also, because taxonomy is
based on evolution, it provides a view of the evolutionary relation-
ships among living species and the relationships between living and
extinct species.
The Classification of Living Organisms Allows Biologists
to Appreciate the Unity and Diversity of Life
Let’s first consider taxonomy on a broad scale. From an evolution-
ary perspective, all forms of life can be placed into three large cat-
egories, or domains, called Bacteria, Archaea, and Eukarya
(Figure 1.8). Bacteria and archaea are microorganisms that are also
termed prokaryotic because their cell structure is relatively simple
(look ahead to Figure 4.8). At the molecular level, bacterial and
archaeal cells show significant differences in their compositions. For
example, the lipids in their cell membranes are biochemically differ-
ent. By comparison, organisms in the domain Eukarya are termed
eukaryotic; they have cells with internal compartments that serve
various functions. A defining distinction between prokaryotic and
eukaryotic cells is that eukaryotic cells have a nucleus in which
the genetic material is surrounded by a membrane. The organisms
in domain Eukarya are divided into seven broad categories called
supergroups.
bro23659_ch01_001-019.indd 10
Each Species Is Placed into a Taxonomic Hierarchy
Taxonomy involves multiple levels in which particular species are
placed into progressively smaller and smaller groups whose mem-
bers are more closely related to each other evolutionarily. Such an
approach emphasizes the unity and diversity of different species.
As an example, let’s consider clownfish, which are found in the
Indian and Pacific Oceans and are popular among saltwater aquar-
ium enthusiasts (Figure 1.9). Several species of clownfish have been
identified.
One species of clownfish, which is orange with white stripes, has
several common names, including ocellaris clownfish. The broadest
grouping for this clownfish is the domain, Eukarya, followed by pro-
gressively smaller divisions, from supergroup (Opisthokonta) to king-
dom (Animalia) and eventually to species. In the animal kingdom,
clownfish are part of a phylum, Chordata, the chordates, which is sub-
divided into classes. Clownfish are in a class called Actinopterygii,
which includes all ray-finned fishes. The common ancestor that gave
rise to ray-finned fishes arose about 420 million years ago (mya).
Actinopterygii is subdivided into several smaller orders. The clown-
fish are in the order Perciformes (bony fish). The order is, in turn,
divided into families; the clownfish belong to the family of marine
fish called Pomacentridae, which are often brightly colored. Families
are divided into genera (singular, genus). The genus Amphiprion is
composed of 28 different species; these are various types of clown-
fish. Therefore, the genus contains species that are very similar to
each other in form and have evolved from a common (extinct) ances-
tor that lived relatively recently on an evolutionary timescale.
Biologists use a two-part description, called binomial
nomenclature, to provide each species with a unique scientific
name. The scientific name of the ocellaris clownfish is Amphiprion
ocellaris. The first word is the genus, and the second word is the spe-
cific epithet, or species descriptor. By convention, the genus name is
capitalized, whereas the specific epithet is not. Both names are itali-
cized. Scientific names are usually Latinized, which means they are
made similar in appearance to Latin words. The origins of scientific
names are typically Latin or Greek, but they can come from a variety
of sources, including a person’s name.
1.3 Reviewing the Concepts
• Taxonomy is the grouping of species according to their evolu-
tionary relatedness to other species. Going from broad to nar-
row groups, each species is placed into a domain, a supergroup,
a kingdom, a phylum, a class, an order, a family, and a genus
(Figures 1.8, 1.9).
1.3 Testing Your Knowledge
1. Which of the following is the broadest group?
a. phylum
b. kingdom
c. class
d. species
e. genus
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 6.2 μm 3.2 μm

(a) Domain Bacteria: Mostly unicellular prokaryotes that inhabit (b) Domain Archaea: Unicellular prokaryotes that often live in extreme
many diverse environments on Earth environments, such as hot springs

375 μm

Protists: Mostly unicellular and some multicellular organisms Plants: Multicellular organisms that can carry out
that are now subdivided into seven broad groups based photosynthesis
on their evolutionary relationships

Fungi: Unicellular and multicellular organisms that have a cell Animals: Multicellular organisms that usually have a nervous
wall but cannot carry out photosynthesis; fungi usually survive system and are capable of locomotion; they must eat other
on decaying organic material organisms or the products of other organisms to live

(c) Domain Eukarya: Unicellular and multicellular organisms having cells with internal compartments that serve various functions

Figure 1.8 The three domains of life. (a) Bacteria and (b) Archaea are domains consisting of species with prokaryotic cells. The third domain,
(c) Eukarya, comprises species that are eukaryotes.
(a) BSIP/age fotostock; (b) Eye of Science/Science Source; (c Protists) Dr. Norbert Lange/Shutterstock; (c Plants) Alexander Dunkel/Sieboldianus/E+/Getty Images;
(c Fungi) Ondrej83/Shutterstock; (c Animals) Ingram Publishing/age fotostock

BioConnections:  Look ahead to Figure 21.1. Are fungi more closely related to plants or animals?

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 12 CHAPTER 1

Taxonomic Ocellaris Approximate time Approximate Examples

group clownfish when the common number of
is found in ancestor for this modern species
group arose in this group

Domain Eukarya 2,000 mya > 5,000,000

Supergroup Opisthokonta > 1,000 mya > 1,000,000

Kingdom Animalia 600 mya > 1,000,000

Phylum Chordata 525 mya 65,000

Class Actinopterygii 420 mya 30,000

Order Perciformes 80 mya 7,000

Family Pomacentride ~ 40 mya 360

Genus Amphiprion ~ 9 mya 28

Species ocellaris < 3 mya 1

Figure 1.9 Taxonomic classification of the clownfish.

Concept Check:  Why is it useful to place organisms into taxonomic groupings?

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 AN INTRODUCTION TO BIOLOGY 13

1.4 Biology as a Scientific Discipline

Learning Outcomes
1. Explain how researchers study biology at different levels,
ranging from molecules to ecosystems.
2. SCISKILLS ⊲ Distinguish between discovery-based science
and hypothesis testing.
3. SCISKILLS ⊲ Describe the steps of the scientific method,
also called hypothesis testing.
4. SCISKILLS ⊲ Explain the usefulness of models in biology.

What is science? Surprisingly, the definition of science is not easy to Ecologists study species in Anatomists and physiologists
state. Most people have an idea of what science is, but actually articu- their native environments. study how the structures of
lating that idea proves difficult. In biology, we can define science as organisms are related to their
the observation, identification, experimental investigation, and theo- functions.
retical explanation of natural phenomena. (a) Ecology—population/community/ (b) Anatomy and physiology—tissue/
Science is conducted in different ways and at different levels. ecosystem levels organ/organism levels
Some biologists study the molecules that compose life, and oth-
ers try to understand how organisms survive in their natural envi-
ronments. Experimentally, they often focus their efforts on model
organisms—organisms studied by many different researchers so
that they can compare their results and determine scientific prin-
ciples that apply more broadly to other species, including humans.
Examples of model organisms include Escherichia coli (a bacterium),
Saccharomyces cerevisiae (a yeast), Drosophila melanogaster (a fruit
fly), Caenorhabditis elegans (a nematode worm), Mus musculus (a
mouse), and Arabidopsis thaliana (a flowering plant). Model organ-
isms offer experimental advantages over other species. For example,
E. coli is a very simple organism that can be easily grown in the lab-
oratory. By limiting their work to a few model organisms, researchers
can gain a deeper understanding of these species, and their results Cell biologists often use Molecular biologists and
may apply broadly to other species, including humans. microscopes to learn how biochemists study the molecules
In this section, we will begin by examining how biologists gener- cells function. and macromolecules that make
up cells.
ally follow an experimental process, called the scientific method,
to test their ideas. We will explore how scientific knowledge makes (c) Cell biology—cellular levels (d) Molecular biology—atomic/
predictions that can be experimentally tested. However, not all dis- molecular levels
coveries are the result of researchers following the scientific method.
Some discoveries are made simply by gathering new information. As
illustrated earlier in Figures 1.1 and 1.2, the characterization of many
living organisms has led to the development of important medicines.
In this section, we will also consider how researchers often set out on
fact-finding missions aimed at uncovering new information that may
eventually lead to important discoveries in biology.
Systems biologists may study
Biologists Investigate Life at Different Levels groups of molecules. The
microarray shown in the inset
of Organization determines the expression of
In Figure 1.4, we examined the various levels of biological organ- many genes simultaneously.
ization. The study of these different levels depends not only on the
scientific interests of biologists but also on the tools available to (e) Systems biology—all levels, shown here at the molecular level
them.
∙∙ The study of organisms in their natural environments is a
Figure 1.10 Biological investigation at different levels of organization.
(a) Thierry Berrod/Mona Lisa Productions/Science Source; (b) Diane Nelson;
branch of biology called ecology, which considers populations, (c) Erik Isakson/Blend Images; (d) Aslysun/Shutterstock; (e) Andrew Brookes/
communities, and ecosystems (Figure 1.10a). Getty Images; (e inset) Alfred Pasieka/Science Source

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 14 CHAPTER 1
∙∙ Researchers who examine the structures and functions of plants
and animals are working in disciplines called anatomy and
physiology, respectively (Figure 1.10b).
∙∙ With major advances in microscopy in the 20th century,
cell biology, which is the study of cells and their interactions,
became an important branch of biology and remains so today
(Figure 1.10c).
∙∙ In the 1970s, genetic tools became available for studying single
genes and the proteins they code. This genetic technology
enabled researchers to study individual molecules, such as
proteins, in living cells and thereby gave rise to the field
of molecular biology. Together with biochemists and
biophysicists, molecular biologists focus their efforts on the
structure and function of the molecules of life (Figure 1.10d).
Such researchers want to understand how biology works
at the molecular and even atomic levels. This approach is
called reductionism—reducing complex systems to simpler
components as a way to understand how the systems work.
∙∙ More recently, scientists have invented tools that allow them to
study groups of genes and groups of proteins. Systems biology
is aimed at understanding how emergent properties arise from
complex interactions. At the molecular or cellular level, systems
biology may involve the investigation of groups of proteins
with a common purpose (Figure 1.10e). For example, a systems
biologist may conduct experiments that try to characterize an
entire cellular process, which is driven by dozens of different
proteins.
A Hypothesis Is a Proposed Idea, Whereas a Theory Is a
Broad Explanation Backed by Extensive Evidence
Let’s now consider the process of science. In biology, a hypothesis
is a proposed explanation for a natural phenomenon. It is a propo-
sition based on previous observations or experimental studies. For
example, with knowledge of seasonal changes, you might hypothe-
size that maple trees drop their leaves in the autumn because of the
shortened amount of daylight. An alternative hypothesis might be
that the trees drop their leaves because of lower temperatures. In
biology, a hypothesis requires more work by researchers to evaluate
its validity.
A useful hypothesis must make predictions—expected out-
comes that can be shown to be correct or incorrect. It should also
be testable, which means that the hypothesis can be shown to be
consistent or inconsistent with data that are obtained via experimenta-
tion. If a hypothesis is incorrect, it should be falsifiable—it can be
shown to be incorrect by additional observations or experimentation.
Alternatively, a hypothesis may be correct, so further experiments will
not disprove it. In such cases, we say that the researchers have failed
to reject the hypothesis. Even so, in science, a hypothesis is never
really proven but rather always remains provisional. Researchers
accept the possibility that perhaps they have not yet conceived of the
correct hypothesis. After many experiments, biologists may conclude
that a hypothesis is consistent with known data, but they should never
say the hypothesis is proven. In such cases, researchers may say that
their data support their hypothesis.
bro23659_ch01_001-019.indd 14
By comparison, a theory, as the term is used in biology, is a
broad explanation of some aspect of the natural world that is sub-
stantiated by a large body of evidence. Biological theories incorpo-
rate observations, hypothesis testing, and the laws of other disciplines
such as chemistry and physics. Theories are powerful because they
allow us to make many predictions about the properties of living
organisms.
As an example, let’s consider the theory that DNA is the genetic
material and that it is organized into units called genes. An over-
whelming body of evidence has substantiated this theory. Thousands
of living species have been analyzed at the molecular level. All of them
have been found to use DNA as their genetic material and to express
genes that produce the proteins that lead to their characteristics. This
theory makes many valid predictions. For example, certain types of
mutations in genes are expected to affect the traits of organisms. This
prediction has been confirmed experimentally. Similarly, this theory
predicts that genetic material is copied and transmitted from parents
to offspring. Through comparisons of the DNA of parents and off-
spring, this prediction has also been confirmed. Furthermore, the
theory explains the observation that offspring resemble their parents.
Overall, two key attributes of a scientific theory are
1. consistency with a vast amount of known data, and
2. the ability to make many correct predictions.
The meaning of the term theory is sometimes muddled because
the word is used differently depending on the situation. In everyday
language, a theory is often viewed as little more than a guess. For
example, a person might say, “My theory is that Professor Simpson
missed class on Wednesday because he thought Professor Idossa
was supposed to teach class that day.” However, in biology, a theory
is much more than a mere guess. A theory is an established set of
ideas that explains a vast amount of data and offers valid predictions
that can be tested. Theories are viewed as knowledge, which is the
awareness and understanding of information.
Discovery-Based Science and Hypothesis Testing Are
Scientific Approaches That Help Us Understand Biology
The path that leads to an important discovery is rarely a straight
line. Rather, scientists ask questions, make observations, ask modi-
fied questions, and may eventually conduct experiments to test their
hypotheses. The first attempts at experimentation may fail, and new
experimental approaches may be needed. To suggest that scientists
follow a rigid scientific method is an oversimplification of the process
of science. Scientific advances often occur as scientists dig deeper
and deeper into a topic that interests them. Curiosity is the key phe-
nomenon that sparks scientific inquiry. How is biology actually con-
ducted? As discussed next, researchers typically follow two general
types of approaches: discovery-based science and hypothesis testing.
Discovery-Based Science The collection and analysis of data
without having a preconceived hypothesis is called discovery-based
science, or simply discovery science. Why is discovery-based sci-
ence carried out? The information gained from discovery-based
science may lead to the formation of new hypotheses and, in the
long run, may have practical applications that benefit people.
Researchers, for example, have identified and begun to investigate
23/09/22 10:07 AM
 AN INTRODUCTION TO BIOLOGY 15

previously unknown genes within humans without already know-
ing the function of those genes. The goal is to gather additional
clues that may eventually allow them to propose a hypothesis that
explains a gene’s function. Discovery-based science often leads to
hypothesis testing.
Hypothesis Testing In biological science, the scientific method—
also known as hypothesis testing—is a process that is usually fol-
lowed to formulate and test the validity of a hypothesis. This strategy
may be described as a five-step process:
1. Observations are made regarding natural phenomena.
2. These observations lead to a hypothesis that tries to explain the
phenomena. A useful hypothesis is one that is testable because it
makes specific predictions.
3. Experimentation is conducted to determine if the predictions are
correct.
4. The data from the experiment are analyzed.
5. The hypothesis is considered to be consistent with the data, or it
is rejected.
The scientific method is intended to be an objective way to
gather knowledge. As an example, let’s return to the question of why
maple trees drop their leaves in autumn. By observing the length of
daylight throughout the year and comparing those data with the time
of the year when leaves fall, one hypothesis might be that leaves fall
in response to shorter periods of daylight (Figure 1.11). This hypoth-
esis makes a prediction—exposure of maple trees to shorter amounts
of daylight will cause their leaves to fall. To test this prediction,
researchers would design and conduct an experiment.
How is hypothesis testing conducted? Although hypothesis test-
ing may follow many paths, certain experimental features are com-
mon to this approach. First, data are often collected in two parallel
ways. One set of experiments is done on the control group, whereas
another set is conducted on the experimental group. In an ideal
experiment, the control and experimental groups differ by only one
factor. For example, an experiment can be conducted in which two
groups of trees are observed, and the only difference between their
environments is the length of time they receive light each day.
To conduct such an experiment, researchers would grow small
trees in a greenhouse, where they could keep other factors such as

1 OBSERVATIONS The leaves on maple trees fall in autumn when the days get colder and shorter.

2 HYPOTHESIS The shorter amount of daylight causes the leaves to fall.

3 EXPERIMENTATION
Small maple trees are grown in
two greenhouses where the only
variable is the length of light.

Control group: Experimental group:

Amount of daily light remains Amount of daily light becomes
constant for 180 days. progressively shorter for 180 days.

4 THE DATA 5 CONCLUSION The hypothesis cannot be rejected.

Number of leaves dropped

200
A statistical analysis
per tree after 180 days

can determine if
the control and the
100 experimental data
are significantly
different. In this
case, they are.

Control Experimental
group group

Figure 1.11 The steps of the scientific method, also known as hypothesis testing. In this example, the goal is to test the hypothesis that maple trees
drop their leaves in the autumn due to the shorter amount of daylight.

Concept Check:  What is the purpose of a control group in hypothesis testing?

bro23659_ch01_001-019.indd 15 23/09/22 10:08 AM

 16 CHAPTER 1
temperature, water, and nutrients the same between the control and
experimental groups while providing the two groups with differ-
ing amounts of light via artificial lighting. In the control group, the
number of hours of light provided would be kept constant each day,
whereas in the experimental group, the amount of light provided
each day would become progressively shorter to mimic seasonal light
changes. The researchers would then record the number of leaves
dropped by the two groups of trees over a certain period of time.
The result of experimentation is a set of data from which a biolo-
gist tries to draw conclusions. Biology is a quantitative science. When
experimentation involves control and experimental groups, a com-
mon form of analysis is to determine if the data collected from the
two groups are truly different. Biologists apply statistical analyses to
their data to determine if outcomes from the control and experimental
groups are likely to differ because of the single variable that differs
between the two groups. A statistics primer is provided at the web-
site for this textbook, and several problems throughout the textbook
will ask you to apply them to a biological problem. When differences
between the control and experimental data are statistically significant,
they are not likely to have occurred as a matter of random chance.
In our example in Figure 1.11, the trees in the control group
dropped far fewer leaves than did those in the experimental group.
Statistical analysis can determine if the data collected from the two
greenhouses are significantly different from each other. If the two
sets of data are found not to be significantly different, the hypothesis
will be rejected. Alternatively, if the differences between the two sets
of data are significant, as shown in Figure 1.11, biologists can con-
clude that the hypothesis is consistent with the data and, therefore,
cannot be rejected. A hallmark of science is that valid experiments
are repeatable, which means that similar results are obtained when
an experiment is conducted on multiple occasions. For our example
in Figure 1.11, the data would be valid only if the experiment were
repeatable.
As described next, discovery-based science and hypothesis test-
ing are often used together to learn more about a particular scientific
topic. As an example, let’s look at how both approaches led to suc-
cesses in the study of the disease called cystic fibrosis.
The Study of Cystic Fibrosis Provides Examples of
Discovery-Based Science and Hypothesis Testing
Let’s consider how biologists made discoveries related to the disease
cystic fibrosis (CF), which affects about 1 in every 3,500 Americans.
Persons with CF produce abnormally thick and sticky mucus that
obstructs the lungs and leads to life-threatening lung infections. The
thick mucus also blocks ducts in the pancreas, which prevents the
digestive enzymes this organ produces from reaching the intestine.
Without these enzymes, the intestine cannot fully absorb amino acids
and fats, which can cause malnutrition. Persons with this disease may
also experience liver damage because the thick mucus can obstruct
the liver. On average, people with CF in the United States currently
live into their mid 40s. Fortunately, as more advances have been made
in treatment, their life expectancy has steadily increased.
Because of this disease’s medical significance, many scientists
are conducting studies aimed at gaining greater information about the
underlying cause of CF. The hope is that knowing more about the
bro23659_ch01_001-019.indd 16
disease may lead to improved treatment options and perhaps even a
cure. As described next, discovery-based science and hypothesis test-
ing have been critical to gaining a better understanding of CF.
The CFTR Gene and Discovery-Based Science In 1935,
American physician Dorothy Andersen determined that cystic
fibrosis is a genetic disorder. Persons with CF have inherited two
faulty CFTR genes, one from each parent. (We now know this gene
codes a protein named the cystic fibrosis transmembrane regulator,
abbreviated CFTR.) In the 1980s, researchers used discovery-based
science to identify this gene. Their search for the CFTR gene did not
require any preconceived hypothesis regarding the function of the
gene. Rather, they used genetic strategies similar to those described
in Chapter 18. Research groups headed by Lap-Chee Tsui, Francis
Collins, and John Riordan identified the CFTR gene in 1989.
The discovery of the CFTR gene made it possible to devise diag-
nostic testing methods to determine if a person carries a faulty ver-
sion of the gene. In addition, the characterization of the CFTR gene
provided important clues about its function. Researchers observed
striking similarities between the CFTR gene and other genes that
were already known to code proteins called transport proteins, which
function in the transport of substances across membranes. Based on
this observation, as well as other kinds of data, the scientists hypothe-
sized that the function of the normal CFTR gene is to code a transport
protein. In this way, the identification of the CFTR gene led them to
conduct experiments aimed at testing a hypothesis about its function.
The CFTR Gene and Hypothesis Testing Researchers inter-
ested in the CFTR gene also considered studies showing that
patients with CF have an abnormal regulation of salt balance across
their plasma membranes. They hypothesized that the normal CFTR
gene codes a transport protein that functions in the transport of
chloride ions (Cl−) across the membranes of cells (Figure 1.12). This
hypothesis led to experimentation that tested normal cells and cells
from CF patients for their ability to transport Cl−. The CF cells
were found to be defective in chloride transport. In 1990, scientists
successfully transferred the normal CFTR gene into cells from CF
patients in the laboratory. The introduction of the normal gene cor-
rected the cells’ defect in chloride transport.
Overall, the results showed that the CFTR gene codes a protein
that transports Cl− across the plasma membrane. A mutation in this
gene causes it to code a defective transporter, leading to a salt imbal-
ance that affects water levels outside the cell, which explains the thick
and sticky mucus in CF patients. In this example, hypothesis testing
provided a way to evaluate a hypothesis about how a disease is caused
by a genetic change.
Observation and Experimentation
Form the Core of Biology
Biology is largely about the process of discovery. Therefore, a recur-
ring theme of this textbook is how scientists design experiments,
analyze data, and draw conclusions. Although each chapter contains
many examples of data collection and experiments, a consistent ele-
ment is a Feature Investigation—an actual study by current or past
researchers. Some of these involve discovery-based science, in which
biologists collect and interpret data in an attempt to make discoveries
23/09/22 10:08 AM
Another random document with
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The Project Gutenberg eBook of
Les nuits champêtres
 This ebook is for the use of anyone anywhere in the United
States and most other parts of the world at no cost and with
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laws of the country where you are located before using this
eBook.

Title: Les nuits champêtres

Author: J.-Charles Laveaux

Release date: February 16, 2024 [eBook #72973]

Language: French

Original publication: Lausanne: J. P. Heubach & c, 1783

Credits: Enrico Segre

*** START OF THE PROJECT GUTENBERG EBOOK LES NUITS

CHAMPÊTRES ***
 L E S

N U I T S
C H A M P Ê T R E S.
 L E S
N U I T S
C H A M P Ê T R E S,
Par M. DE LA VEAUX.

Restat ut his ergo me ipse regam solerque

elementis.

H O R A T.

A L A U S A N N E,

Chez J. P. HEUBACH & Comp.

M. D C C. L X X X I V.
 A S O N

ALTESSE SÉRÉNISSIME
MONSEIGNEUR

L E L A N D G R AV E
R É G N A N T
DE HESSE-CASSEL.

M O N S E I G N E U R,

Le suffrage du Public est la récompense la plus flatteuse des

Ecrivains. J'ai tâché de le mériter, en composant ce petit Essai; je
suis sûr de l'obtenir, si je me suis rendu digne de celui de V O T R E
A L T E S S E S É R É N I S S I M E. En dédiant ce petit Ouvrage au
Protecteur éclairé des Sciences & des Arts, je me acquitte pour ma
part de l'hommage & de la recennoissance que lui doivent les Gens
de Lettres de toutes les Nations. Il est aisé de protéger les Lettres, il
ne l'est pas de le faire avec discernement. Ceux qui connoissent les
personnes de mérite qui composent l'Académie de Hesse-Cassel,
conviendront avec moi que les Gens de Lettres qui ont Le bonheur
de mériter l'attention gracieuse de V O T R E A L T E S S E
S É R É N I S S I M E, doivent compter sur les suffrages de le
postérité.

Je suis avec la soumission la plus respectueuse,

 M O N S E I G N E U R,

DE VOTRE ALTESSE SÉRÉNISSIME,

Berlin, ce 1 Avril 1783.

Le très-humble & très-

obéissant serviteur
DE LA VEAUX.
 P R É FA C E .

I L n'est point d'ame sensible qui n'ait goûté, dans

la solitude, ces instans délicieux où l'homme,
écartant les prestiges du mensonge, rentre dans son
propre cœur, pour y chercher les étincelles de la
vérité. Quel plaisir, après avoir été ballotté pendant
quelque temps sur la mer de ce monde, de le retirer
sur le sommet d'un rocher paisible, pour y considérer
en sûreté les tempêtes & les naufrages qui s'y
succédent! Heureux celui qui peut alors oublier un
instant les vains préjugés dont son ame est remplie!
les miseres de l'humanité disparoissent à ses yeux,
l'auguste Vérité remplit son cœur dune joie pure. Ce
n'est que dans ces instans & dans ceux qui
précedent la mort, que l'homme peut apprendre à
connoître ce qu'il est sur la terre & ce que la terre est
pour lui.
 Rassasié du monde que j'ai vu sous différens
aspects, la solitude est devenue pour moi un besoin.
Je m'occupe quelquefois à écrire les sentimens & les
réflexions qu'elle m'inspire; c'est un choix de ces
réflexions que je donne ici au Public. Je n'ai prétendu
faire ni des Poëmes, ni des Traités philosophiques;
j'ai voulu peindre les diverses impressions que mon
ame éprouvoit dans ces momens.

S'il est vrai, comme l'a dit Jean-Jacques

Rousseau, qu'on écrit bien, quand on est convaincu
de la vérité de ce qu'on écrit, les Nuits Champêtres
doivent avoir quelque mérite. Elles ne contiennent
pas un sentiment que je n'aye puisé dans mon cœur,
pas une vérité dont je ne sois convaincu. La plupart
des petites histoires que j'y raconte ne sont point le
fruit de mon imagination; le fond en est vrai: je n'ai
fait que changer la scene & les acteurs. Le caractere
de Dorval qu'on trouve dans la sixieme Nuit, n'est
point imaginé, & le trait de cet homme bienfaisant
que je rapporte dans la neuvieme, est vrai jusques
dans les moindres détails.
 Je suis bien éloigné de croire que cet Ouvrage ne
trouvera point de critiques, dans un siecle où l'on ne
lit guere que pour critiquer. Quelques Gens de
Lettres ont lu la premiere Nuit; les uns l'ont louée
avec trop de politesse, d'autres l'ont condamnée elle
& les sœurs qu'elle devoit avoir. Quelques Gens du
monde ont lu l'Ouvrage entier, & m'ont assuré, d'un
air vrai, l'avoir lu avec plaisir. Quoi qu'il en soit, je ne
serois point insensible au succès qu'il pourroit avoir;
ce seroit un grand plaisir pour moi de procurer
quelques instans agréables à mes Lecteurs. Mais s'il
est condamné à n'être connu que par des critiques
ameres, je m'en consolerai aisément; je n'ai jamais
regardé, comme un fort grand malheur, de ne pas
savoir tourner ses pensées au goût de son siecle; &
d'ailleurs, je me suis fait des principes de bonheur
qui sont indépendans de la plume des Journalistes.
En un mot, s'il est décidé que l'Ouvrage soit
mauvais, je serai le premier à le jetter au feu, & je
tâcherai de m'occuper plus utilement une autre fois.
 TA B L E .

PREMIERE NUIT. La Campagne, p. 1

SECONDE NUIT. Dieu, 33
TROISIEME NUIT. L'Homme, 53
QUATRIEME NUIT. La Science, 87
CINQUIEME NUIT. La Société, 109
SIXIEME NUIT. Le Bienfaisance, 150
SEPTIEME NUIT. L'Amitié, 169
HUITIEME NUIT. L'Amour, 186
NEUVIEME NUIT. Le Bonheur, 205
DIXIEME NUIT. La Mort, 224
 L E S N U I T S
CHAMPÊTRES.

P R E M I E R E N U I T.
L a C a m pa g n e .

J E vous quitte enfin, tristes asiles de l'orgueil & de l'esclavage,

villes superbes où le vice rampe si souvent sous la vertu humiliée, où
l'or tient lieu de tout, où les sentimens précieux de la nature sont
étouffés sous le brillant attirail de la folie! Je te revois, Campagne
chérie, doux asile du bonheur & de l'innocence! Mon ame, dégagée
de ses chaînes, respire l'air délicieux de la liberté; mes idées, mes
sentimens s'étendent comme le vaste horizon dont les bornes
échappent à ma vue: un charme secret s'empare de tous mes sens;
je vais être heureux. Rendu à ma chere cabane, je vais remplir la
destination de la nature. Un travail modéré suffit à mes besoins; mon
jardin, mon champ & mon troupeau offrent en abondance les vrais
biens à ma main laborieuse. A l'abri de vos traits ensanglantés,
barbare Calomnie! perfide Médisance! je verrai de loin les victimes
innocentes qui tombent sous vos coups; je pleurerai sur ces
infortunés & je bénirai le sort qui me soustrait à vos fureurs.

Toutes les fois que le soleil disparoissant de notre horizon, ira

répandre ses bienfaits sur un autre hémisphere, je cesserai mon
travail, pour venir errer nonchalamment le long des bords de ce
ruisseau, ou je me coucherai négligemment auprès de sa source
sacrée. Là, le spectacle ravissant de la nature élévera mon ame &
charmera mon cœur. J'écarterai les préjugés funestes dont les
nuages épais obscurcissent la vérité. Seul avec la nature, j'écouterai
sa voix, j'ouvrirai mon ame à ses inspirations divines, j'étudierai les
devoirs qu'elle me prescrit, je jouirai de tous les plaisirs qu'elle
m'offre. Je déchirerai les enveloppes étrangeres sous lesquelles
l'homme se cache à mes yeux, je le verrai tel qu'il est.

Déjà je sens l'influence secrete de la nature, elle semble répandre

autour de moi une atmosphere d'innocence & de volupté. Un feu
secret s'allume dans mon cœur; il s'augmente, il coule dans toutes
mes veines: état délicieux! Tout m'enchante, m'intéresse, me ravit;
tout me remplit de l'idée de la liberté & du bonheur. Mes fers ont
disparu, la volupté brûle mon cœur, & mes yeux répandent les
larmes délicieuses du sentiment.

La nuit répand sur l'Univers un calme profond qui se communique

à mon ame; les Zéphyrs m'apportent doucement le baume
voluptueux des fleurs; les arbres, en courbant au-dessus de ma tête
leurs branches touffues, augmentent l'obscurité de ma retraite; & la
lune dont la lumiere tremblotante se joue à travers le feuillage, vient
argenter les flots du ruisseau qui fuit à mes côtés. Le bruit des eaux
qui se brisent en tombant sur ces rochers escarpés, se mêle au
vaste silence de la nature, sans paroître l'interrompre, ou plutôt il
semble l'augmenter encore.
 Tous les êtres jouissent ici du repos & du bonheur. Mille insectes
divers se jouent dans l'herbe humide & fraîche, les poissons folâtrent
sans crainte au milieu des eaux; & les oiseaux, sur leurs nids,
savourent voluptueusement les plaisirs de l'amour & du repos:
récompense délicieuse des travaux que la tendresse paternelle leur
a fait essuyer pendant la chaleur du jour.

Innocens animaux, vous goûtez en paix les douceurs de la

liberté! la nuit semble vous envelopper de ses ombres bienfaisantes
pour vous soustraire à la cruauté de l'homme: profitez de ces doux
instans; bientôt le jour va paroître, & votre tyran avec lui.

Aimable rossignol, fais entendre les accens de ta voix; charme

les ennuis de ta compagne chérie, occupée à répandre dans son nid
la chaleur qui va faire éclorre les doux fruits de vos amours
mutuelles; que tes chants fassent retentir tous les échos d'alentour:
ils doivent charmer toute la nature, c'est la tendresse qui les inspire!
Et vous, sensibles tourterelles, livrez-vous sans réserve aux
douceurs de l'amour & de la fidélité; que vos tendres plaintes soient
les expressions de vos plaisirs! Entrelacez vos becs amoureux; que
le frémissement voluptueux de vos ailes agite doucement le
feuillage; mais fuyez dès que l'aurore éclairera vos retraites: l'homme
paroîtra; peut-être que sa main cruelle vous ôtera la vie. Il sait vous
arracher à vos plaisirs innocens; mais, hélas! il ne sait pas les goûter.

J'entends remuer les branches de la forêt, un animal frappe la

terre & s'avance de mon côté, il sort du milieu des broussailles, il
paroît au clair de la lune; c'est un cerf qui vient se désaltérer dans
l'onde pure du ruisseau. Que la forme de son corps est noble & belle!
on diroit que la nature a pris plaisir à répandre sur lui l'élégance & la
beauté. Hélas! à quoi servent ces dons? Demain, si-tôt que l'aurore
annoncera le retour de la lumiere, une meute de chiens cruels,
excités par des hommes plus cruels encore, le forceront à sortir de
sa retraite, & le poursuivront jusqu'à ce que ses genoux chancelans
tremblent sous le poids de son corps accablé. Barbares! quel mal
vous ont fait ces animaux timides, pour prendre plaisir à les
tourmenter? Vous êtes des hommes, & vous vous faites un plaisir de
la douleur d'un être foible qui fuit devant vous? Le cerf pleure sa
défaite & sa mort; le lievre blessé tourne sur vous un regard
languissant, qui vous reproche votre cruauté; la perdrix expirante
semble implorer votre pitié pour sa couvée fugitive. Mais comment
vous laisseriez-vous attendrir par ces animaux innocens? les maux
que vous leur faites souffrir ne sont qu'une foible image des
tourmens dont vous accablez vos semblables.

Que manquera-t-il à mon bonheur, si cette retraite peut me

soustraire à votre cruauté? Monde faux & trompeur, je ne regretterai
point les illusions par lesquelles tu éblouis les insensés qui te
consacrent leur cœur! je les verrai sans envie jouir des prétendus
biens dont tu les combles. J'ai vu tes favoris s'empresser après des
bulles brillantes & légeres que l'air promene à son gré, & qui se
dissipent à l'instant qu'on croit les saisir; je les ai vus se tourmenter
pendant toute leur vie pour des chimeres qui doivent faire leur
malheur, & périr enfin sans avoir goûté le repos. J'ai vu dans tes
cercles brillans l'or adoré sous mille noms pompeux. J'ai vu le Dieu
de la Frivolité dicter insolemment ses arrêts ridicules, & les répandre
sur toute la terre. J'ai vu ceux que tu appelles sages se jouer avec
les hochets de la folie, & danser au son de ses grelots. L'homme
sensible cherche en vain sur cette terre un cœur où il puisse reposer
son cœur, il n'en trouve point. Le souffle empoisonné de l'avarice, de
la perfidie & du mensonge a flétri toutes les vertus. L'homme de bien
est seul sur la terre. Il ressemble à ces plantes transportées dans
des climats étrangers, qui se desséchent & penchent leur tige
languissante, faute d'avoir une nourriture qui leur convienne.

Le sommeil bienfaisant regne dans la cabane du laboureur & du

berger. Il n'est point interrompu par les cris aigus d'une conscience
criminelle, ni par les monstres affreux de la haine, de la perfidie, de la
vengeance; ni par les vains projets de l'avarice & de l'ambition.
L'innocence & la paix regnent avec lui. Le chant des oiseaux va
bientôt réveiller ces mortels fortunés. Il me semble les voir regarder
avec joie l'aurore naissante. Leurs forces renouvellées circulent avec
impétuosité dans leurs membres reposés, & y portent par-tout le
besoin du travail. Bientôt ils se répandent dans la campagne, &
reprennent en souriant leurs utiles travaux. Peuvent-ils le refuser aux
transports de la reconnoissance, lorsqu'ils voient, lorsqu'ils entendent
toute la nature célébrer le retour de la lumiere; lorsqu'ils voient le
soleil darder sur leurs moissons & sur leurs fruits les rayons
bienfaisans dont la chaleur précieuse va travailler en silence à
l'œuvre merveilleuse de la maturité, ou faire éclorre les germes
féconds que la terre renferme dans son sein?

Soit que le laboureur recueille l'herbe fleurie de ses prés, soit qu'il
trace lentement un pénible sillon, ou qu'il coupe les épis courbés
pour en former de lourdes gerbes, soit qu'il soulage les branches
affaissées de ses arbres fruitiers, soit enfin qu'il remplisse ses
celliers de la dépouille vermeille de la vigne; la joie, l'espérance ou le
plaisir charment toujours ses travaux.

Dormez tranquillement, heureux habitans de la Campagne,

jouissez des plus doux présens des cieux, & n'enviez point le sort du
riche dont l'éclat vous éblouit. Pendant que la nature répand sur vos
membres fatigués les bienfaits du repos, pendant qu'elle prépare
autour de vous les plaisirs qui doivent charmer votre réveil; le crime,
au milieu des villes, aiguise ses poignards, prépare ses noirs
poisons, & marque ses victimes. Le remords affreux vole de palais
en palais, il seme par-tout épouvante & l'effroi. Il tire avec fracas les
rideaux pompeux du riche coupable, & fait siffler autour de lui ses
horribles serpens. L'un couché sur des coussins qui semblent
préparés par les mains de la volupté, pousse au ciel des cris aigus
que lui arrache la douleur cruelle. Il souffre des maux que le travail &
la frugalité ont écartés de vos chaumieres. Ennemi de la nature, il a
voulu lui arracher les plaisirs destinés à ses favoris; il a voulu jouir du
bonheur & du repos sans l'avoir mérité par son travail & son
innocence: il en est puni; la nature outragée se venge de sa violence
& de ses mépris. Un autre, plongé dans le désespoir, est prêt à
s'arracher lui-même une vie que les suites honteuses du vice lui ont
rendue odieuse. Ici la perfidie & le mensonge trament leurs intrigues
secretes; là le jeu, la débauche & les profusions de toute espece
renversent des fortunes & plongent les familles dans le désespoir. Je
vois ces malheureux lever les mains au ciel, & ramper dans la
bassesse & la misere. Qu'ils seroient heureux, si leurs bras étoient
accoutumés au travail, si leurs cœurs ne connoissoient d'autres
besoins que les vôtres!
 Et quels plaisirs pourroient donc être comparés à ceux dont vous
jouissez! S'il en étoit quelques-uns, ce seroient ceux de l'homme utile
qui, au milieu de ces villes, travaille à diminuer les maux de
l'humanité. Mais, hélas! qu'il paye cher le plaisir de faire du bien à
ses semblables! le fanatisme lui prépare des persécutions & des
fers. Ce monstre affreux se traîne sur les restes des bûchers que sa
fureur alluma & que la raison éteignit; ses membres livides fouillent
parmi les cendres & les ossemens; il cherche quelque étincelle qui
puisse rallumer ces flammes odieuses. Que dis-je? hélas!
Campagnes innocentes, vous n'êtes pas à l'abri de ses fureurs. Il
poursuit ses déplorables victimes jusques dans le sanctuaire sacré
de la nature, & vos ruisseaux ont été teints plus d'une fois du sang
qu'il a versé. La mort, la mort même, n'est pas un asile assuré contre
sa férocité. Il s'acharne sur des cadavres palpitans, il les emporte en
les secouant avec fureur, il craint que la terre ne les dérobe à sa
rage; & si la raison tremblante ose leur rendre en secret les derniers
devoirs, il se jette sur la fosse en poussant des hurlemens affreux,
ronge la terre qui les cache, & la couvre d'écume & de sang.

L'oubli, oubli seul de la nature a causé tous ces maux. Ce n'est

pas celui qui consacre tous les instans de sa vie à des travaux utiles
qui alluma le premier les flambeaux de la haine, de la superstition &
du fanatisme; c'est celui qui; renonçant aux avantages d'une vie
laborieuse & innocente, chercha dans la crédulité & dans la foiblesse
de ses semblables des ressources contre les besoins qui naissent en
foule de l'oisiveté & des vices.

Heureuse innocence, précieuse médiocrité, vous étiez destinées

à faire le bonheur de l'homme! il est malheureux dès qu'il vous
abandonne, & ce n'est qu'à la Campagne qu'il peut vous retrouver
dans toute votre pureté.

Quel bruit frappe mes oreilles? Un char pompeux s'avance, il

s'ouvre; c'est un riche que le dégoût de la ville, ou plutôt le dégoût de
sa propre existence, conduit à la Campagne. Il vient y chercher le
bonheur & le repos, il n'y trouvera ni l'un ni l'autre. Ce n'est qu'à ses
amis que la nature accorde ses faveurs; & il apporte avec lui tout
l'attirail des vices & des préjugés de la ville. Des hommes ses