Biology : concepts and investigations

Fourth Edition Mariëlle Hoefnagels

Visit to download the full and correct content document:
https://ebookmass.com/product/biology-concepts-and-investigations-fourth-edition-ma
rielle-hoefnagels/
BIOLOGY
This page intentionally left blank
BIOLOGY
Concepts and Investigations

FOURTH EDITION

Mariëlle Hoefnagels
The University of Oklahoma

Media Contributions by
Matthew S. Taylor
BIOLOGY: CONCEPTS AND INVESTIGATIONS, FOURTH EDITION

Published by McGraw-Hill Education, 2 Penn Plaza, New York, NY 10121. Copyright © 2018 by McGraw-Hill
Education. All rights reserved. Printed in the United States of America. Previous editions © 2015, 2012, and 2009.
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 Education, 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.

1 2 3 4 5 6 7 8 9 0 DOW 20, 19, 18, 17

ISBN 978-0-07-802420-7
MHID 0-07-802420-X

Chief Product Officer, SVP Products & Markets: G. Scott Virkler

Vice President, General Manager, Products & Markets: Marty Lange
Vice President, Content Design & Delivery: Betsy Whalen
Director, Biology: Lynn Breithaupt
Brand Manager: Michelle Vogler
Director of Product Development: Rose Koos
Product Developer: Anne Winch
Marketing Manager: Britney Ross
Digital Product Analyst: Christine Carlson
Director, Content Design & Delivery: Linda Avenarius
Program Manager: Angela R. FitzPatrick
Content Project Manager (print): Peggy Selle
Content Project Manager (assessment): Emily Windelborn
Buyer: Laura Fuller
Designer: Tara McDermott
Content Licensing Specialists: Lori Hancock/Deanna Dausener
Cover Image: (front) ©mustamin/Getty Images RF
Compositor: MPS Limited
Typeface: 10/12 STIX MathJax Main
Printer: R. R. Donnelley

All credits appearing on page or at the end of the book are considered to be an extension of the copyright page.
Leaf: ©image broker/Alamy RF
Flames: ©Don Farrall/Getty Images RF

Library of Congress Cataloging-in-Publication Data

Names: Hoefnagels, Mariëlle.

Title: Biology : concepts and investigations.
Description: Fourth edition / Mariëlle Hoefnagels, The University of Oklahoma;
media contributions, Matthew S. Taylor. | New York, NY : McGraw-Hill Education, [2018] | Includes index.
Identifiers: LCCN 2016047958 | ISBN 9780078024207 (alk. paper)
Subjects: LCSH: Biology–Textbooks.
Classification: LCC QH307.2 .H64 2018 | DDC 570–dc23 LC record available at https://lccn.loc.gov/2016047958

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 Education, and McGraw-Hill Education does not guarantee
the accuracy of the information presented at these sites.

www.mhhe.com
Brief Contents
UNIT 1 Science, Chemistry, and Cells UNIT 5 Plant Life
1 The Scientific Study of Life 2 22 Plant Form and Function 460
2 The Chemistry of Life 18 23 Plant Nutrition and Transport 482
3 Cells 44 24 Reproduction and Development of Flowering
4 The Energy of Life 70 Plants 496
5 Photosynthesis 88
6 Respiration and Fermentation 104
UNIT 6 Animal Life
UNIT 2 DNA, Inheritance, 25 Animal Tissues and Organ Systems 518
26 The Nervous System 534
and Biotechnology 27 The Senses 558
7 DNA Structure and Gene Function 120 28 The Endocrine System 574
8 DNA Replication, Binary Fission, and Mitosis 146 29 The Skeletal and Muscular Systems 590
9 Sexual Reproduction and Meiosis 166 30 The Circulatory System 608
10 Patterns of Inheritance 186 31 The Respiratory System 628
11 DNA Technology 216 32 Digestion and Nutrition 644
33 Regulation of Temperature and Body Fluids 664
34 The Immune System 680
UNIT 3 The Evolution of Life 35 Animal Reproduction and Development 700
12 The Forces of Evolutionary Change 236
13 Evidence of Evolution 260
14 Speciation and Extinction 280
UNIT 7 The Ecology of Life
15 The Origin and History of Life 304 36 Animal Behavior 728
37 Populations 748
38 Communities and Ecosystems 766
UNIT 4 The Diversity of Life 39 Biomes 788
16 Viruses 330 40 Preserving Biodiversity 808
17 Bacteria and Archaea 344
18 Protists 360
19 Plants 378
20 Fungi 398
21 Animals 416

v
 About the Author Mariëlle Hoefnagels is an associate professor in the
Department of Biology and the Department of Microbiology and

Plant Biology at the University of Oklahoma, where she teaches

courses in introductory biology, mycology, and science writing.

She has received the University of Oklahoma General Education

Teaching Award and the Longmire Prize (the Teaching Scholars

Award from the College of Arts and Sciences). She has also

been awarded honorary memberships in several student honor

societies.

Dr. Hoefnagels received her B.S. in environmental science

from the University of California at Riverside, her M.S. in soil

science from North Carolina State University, and her Ph.D. in

plant pathology from Oregon State University. Her dissertation

work focused on the use of bacterial biological control agents

to reduce the spread of fungal pathogens on seeds. In addition

©Davenport Photos
to authoring Biology: Concepts and Investigations and Biology:

The Essentials, her recent publications have focused on creating

investigative teaching laboratories and methods for teaching

experimental design in beginning and advanced biology classes.

She frequently gives presentations on study skills and related

topics to student groups.

vi
Preface
Vision and Change in Undergraduate Biology Education: A Call
to Action encourages instructors to improve student engagement
and learning in introductory biology courses. The central idea
of the original Vision and Change report—and of the confer-
ences and reports that followed—is that we need to turn away
from teaching methods that reward students who memorize and
regurgitate superficial knowledge. Instead, we need to emphasize
deeper learning that requires students to understand and apply
course content. This idea is precisely what I have tried to achieve
since I started teaching at the University of Oklahoma in 1997,
and it has been a guiding principle in the creation of my books
and digital material as well.
This edition retains what users have always loved about
this book: the art program, readable narrative, handy study
tips, ­Investigating Life essays, and tutorial animations. We
also supply a variety of supplements that make teaching easier,
including eye-catching PowerPoint® lectures with integrated
clicker questions that assess conceptual understanding. As you
examine this new edition, however, I hope you will see an even
stronger emphasis on connections and the “big picture.” Our
most prominent new feature, Survey the Landscape, shows
how each chapter’s content fits into the unit’s overall empha-
sis. Students often struggle to connect new topics to what they
have learned previously; Survey the Landscape is designed to
help them keep an eye on the big picture. These new figures,
which appear in each chapter opener, can be integrated with
the Pull It Together figure in every chapter’s summary to help
students see the “forest” and the “trees.”
Many other changes to this book reflect the growing num-
bers of instructors and students who are embracing digital text-
books. Much of our work for the fourth edition was behind-
the-scenes adjustments that make the narrative and art more
digital-friendly. Moreover, SmartBook® user data from thou-
sands of students using the third edition helped us to identify
passages that needed clarification. The user data also guided
us as we created a carefully selected array of digital Learning
­Resources to accompany many probes in SmartBook. In addi-
tion, many chapters have bonus features for ebook users, includ-
ing new digital-only miniglossaries and figures.
I agree with the Vision and Change report’s call for instruc-
tors to embrace active learning techniques, but I also believe that
one set of tools and techniques does not work in every classroom.
For that reason, my team and I are proud to create a package that
gives you the flexibility to teach introductory biology in a way
that works best for you. The following sections illustrate the fea-
tures and resources for this edition that can help you meet your
teaching goals.
I hope that you and your students enjoy this text and that it
helps cultivate an understanding of, and deep appreciation for,
biology.
Mariëlle Hoefnagels
The University of Oklahoma
vii
 Author’s Guide To Using This Textbook
This guide lists the main features of each chapter and describes some of the ways that I use them
in my own classes.

The Learning Outline introduces the chapter’s

©Davenport Photos

main headings and helps students keep the big

Is It Easier Being Green?
Food is expensive. It would be much cheaper and easier if we
could feed ourselves using photosynthesis. Imagine the benefit of
being photosynthetic: You could make your own food, free of
charge, simply by sitting outside in the sun.
Of course, your body would have to have some new adapta-
picture in mind.
Each heading is a complete sentence that summarizes the
most important idea of the section.
The gradual change in leaf colors as a chapter unfolds
L E AR NI N G OU T L I N E indicates where the student is in the chapter’s big picture.
5.1 Life Depends on Photosynthesis
5.2 Sunlight Is the Energy Source for Photosynthesis Students can also flip to the end of the chapter before
5.3 Photosynthesis Occurs in Two Stages

5.4 The Light Reactions Begin Photosynthesis

tions for photosynthesis to work. Your skin would have to be
green, for starters. You might even have skin flaps that capture
extra sunlight. You wouldn’t eat, so you would need another
way to acquire essential minerals; perhaps your feet would
grow rootlike extensions that would absorb water and nutrients
from soil.
Maybe photosynthetic cows, pigs, and chickens—or pets
such as dogs and cats—would be a better idea. Feed-free animals
5.5 The Carbon Reactions Produce Carbohydrates
5.6 C3, C4, and CAM Plants Use Different Carbon Fixation
Pathways
5.7 Investigating Life: Solar-Powered Salamanders
M ULT IP L E CHOICE QUESTIONS
starting to read; the chapter summary and Pull It Together
concept map can serve as a review or provide a preview of
what’s to come.
CHAPTER 5 Photosynthesis 103
7. In 1941, biologists exposed photosynthesizing cells to water
containing a heavy oxygen isotope, designated 18O. The “labeled”

Concept maps help students see the big

would be a commercial and environmental triumph, costing less isotope appears in the O2 gas released in photosynthesis, showing that
1. Where does the energy come from to drive photosynthesis? the oxygen came from the water. Where would the 18O have ended up
to own and generating less waste than the animals we raise now. a. A chloroplast c. The sun if the researchers had used 18O-labeled CO2 instead of H2O?
picture.
Fortunately or unfortunately, scientists will probably never
b. ATP d. Glucose
be able to create photosynthetic people, chickens, or pooches. 8. Over the past decades, the CO2 concentration in the atmosphere has
Mammals and birds move, breathe, pump blood, and maintain 2. Animals and other rely on that carry out increased. (a) Predict the effect of increasing carbon dioxide
high body temperatures. All of this activity would likely require photosynthesis. concentrations on photorespiration. (b) Scientists suggest that
energy beyond what photosynthesis alone could supply.
Some animals, however, have adopted the “green” lifestyle
a. autotrophs; autotrophs c. heterotrophs; autotrophs
b. heterotrophs; heterotrophs d. autotrophs; heterotrophs New Survey the Landscape concept maps at the start of
increasing CO2 concentrations are leading to higher average global
temperatures. If temperatures are increasing, does this change your
by harboring live-in photosynthetic partners (see section 5.7).
The closest to a true plant–animal hybrid is probably the sea slug
3. Photosynthesis is an example of an
because .
chemical reaction each chapter illustrate how the pieces of the entire unit
answer to part (a)?
9. How does photosynthesis help compensate for increasing
Elysia chlorotica, a solar-powered mollusk with chloroplasts
(photosynthetic organelles) in the cells lining its digestive tract.
a. exergonic; energy is released by the reaction center pigment
b. endergonic; light energy is used to build chemical bonds
fit together. These
atmospheric CO ?new figures
Where does
2 the CO go?integrate
2 with the existing
Does cutting down
SURVEY THE LANDSCAPE forests likely increase or decrease the rate of CO accumulation in
The chloroplasts come from algae in the slug’s diet. As the ani-
mal grazes, it punctures the algal cells and discards everything
c. exergonic;
Science, Chemistry,light energy is captured by pigment molecules
and Cells
d. endergonic; the reactions occur inside a cell
Pull It Together
the atmosphere?concept maps in the chapter summary. 2

but the chloroplasts, which migrate into the animal’s cells. Light 10. How is the CAM pathway adaptive in a desert habitat?
4. The evolution
scientific of photosynthesis resulted in
passes through the slug’s skin and strikes the food-producing
chloroplasts. Once its “solar panels” are in place, the animal may
Life

consists carry
is the
a. anstudy
increase
of
Biology
in the amount of O2 in the atmosphere. After spending
11. Explain howclass 4timeis discussing
C photosynthesis the key
based on a spatial arrangement
structures, whereas CAM photosynthesis is temporally based.
of points in
out theb. the initial appearance of heterotrophs.
constructing
12. Explainconcept
why each of themaps, I have my about students draw
of units
not eat again for months! called functions of
c. global warming. following misconceptions
Perhaps the most famous animals to “farm” photosynthetic ATP makes
d. an increase in the Respiration
amount of CO2 in the atmosphere. photosynthesis is false: (a) Only plants are autotrophs. (b) Plants do
partners are corals. Inside the coral are single-celled protists
called dinoflagellates, which use the sun’s energy to feed the
Cells
5. Only high-energy
uses
light can penetrate the ocean and reach concept maps of their
not need cellular own.
respiration because they carry out photosynthesis.
coral. In exchange, the animals provide a home for the protists.
consist of Carbohydrates
photosynthetic organisms in coral reefs. What color of light would (c) Chlorophyll is the only plant pigment.
makes
Sometimes, however, the partners break up. Corals under stress Molecules
you predictProteins
include
these organisms use?
Photosynthesis
sometimes expel their dinoflagellates, or the protists may leave a. Red c. Blue
on their own. The reef then turns white. The coral animals even- consist of
b. encodes
Yellowsuch as d. Orange PULL IT TOGETHER
tually die, endangering the entire reef ecosystem. Pollution, dis- 6. DNAThe primary function of the light reactions is to , whereas
Atoms Enzymes
ease, shading, excessively warm water, and ultraviolet radiation the primary function of the carbon reactions is to .
all trigger coral bleaching. Biologists predict that global climate
a. convert the sun's energy into chemical energy; store chemical energy
change will only make this problem worse. Photosynthesis
Photosynthetic cellsb.useuse light
carbon energy
dioxide, to produce
water, and sunlight ATP;
to use chemical energy to produce
Corals and sea slugs are not the only animals whose lives build carbohydrates. These ATPmolecules, which make up much of a
depend on photosynthesis. Yours does, too, as you will learn in plant's body, can also be used to store energy and generate ATP. occurs in
c. store light; use light energy to produce carbon two stages
the next two chapters. For more details, study the Pull It Together feature in the chapter summary.
d. transfer heat captured from light to electrons; use electrons to
generate organic molecules
89
7. Photorespiration becomes more likely when produce
Light Carbon
a. CO2 concentrations are high in leaf cells. reactions reactions
b. stomata remain closed in C3 plants. is energy
c. glucose concentrations are low in leaf cells. release as source for
d. ATP binds to rubisco. is electron waste is electron produce
source for product source for
8. A plant that opens its stomata only at night is a ATP
a. C2 plant. c. C4 plant. is carbon
b. C3 plant. d. CAM plant. H2O O2 source for
NADPH
Answers to these questions are in appendix A. is energy CO2
source for
WRIT E IT OUT Sugar
Light
1. Imagine that multiple simultaneous volcanic eruptions send black ash
into Earth’s atmosphere, making photosynthesis impossible anywhere
Figure 5.15 Pull It Together: Photosynthesis.
on Earth for many years. What would be the consequence to plants?
To animals? To microbes?
2. Other stars in the galaxy emit light at different wavelengths than the Refer to figure 5.15 and the chapter content to answer the following questions.
sun. If photosynthesis evolved on a planet around one of these stars, 1. Review the Survey the Landscape figure in the chapter introduction,
how might it be different from and similar to photosynthesis on Earth? and then add enzymes, cells, molecules, and respiration to the Pull It
3. Define these terms and arrange them from smallest to largest: thylakoid Together concept map.
membrane; photosystem; chloroplast; granum; reaction center. 2. How would you incorporate the Calvin cycle, rubisco, C3 plants, C4
4. Would a plant grow better in a room painted blue or in a room painted plants, and CAM plants into this concept map?
green? Explain your answer. 3. One possible connecting phrase in the concept map is “Chlorophyll
5. Determine whether each of the following molecules is involved in the reflects wavelengths of light.” Fill in the blank and explain
light reactions, the carbon reactions, or both and explain how: O2, your answer.
CO2, carbohydrates, chlorophyll a, photons, NADPH, ATP, H2O. 4. Build another small concept map showing the relationships among the
6. Of the many groups of photosynthetic bacteria, only cyanobacteria use terms chloroplast, stroma, grana, thylakoid, photosystem, and chlorophyll.
chlorophyll a. How does this observation support the hypothesis that 5. Besides respiration, what happens to the sugar produced in
cyanobacteria gave rise to the chloroplasts of today’s plants and algae? photosynthesis?

viii
 AUTHOR’S GUIDE ix

Coral Competition. These two brain coral colonies are competing for space off the coast of Roatan, Honduras. Photosynthetic algae, which live inside each coral
animal’s tissues, provide one source of their food.
©Andrew J. Martinez/Science Source

Learn How to Learn study tips help students LEARN HOW TO LEARN
See What’s Coming

develop their study skills. Start by reviewing the Survey the Landscape figure at the start of each chapter to see how the material fits with the rest of the unit. Then check out the
Learning Outline. Each heading is a complete sentence that summarizes the most important idea of the section. Read through these statements before
you start each chapter. You can also flip to the end of the chapter before you start to read; the chapter summary and Pull It Together concept map can

Each chapter has one Learn How to Learn study tip, and a
provide a preview of what’s to come.

complete list is in Appendix F.

I present a Study Minute in class each week, with
examples of how to use these study tips.
CHAPTER 5 Photosynthesis 101

Investigating Life describes a real experiment INVESTIGATING LIFE

Average Net
Radioactivity
Average
Hourly Source of

focusing on an evolutionary topic related to

Difference (Light Change in Carbon
5.7 Solar-Powered Salamanders Sample
Whole egg
Minus Dark) Carbon Increase

This chapter’s opening essay described two examples of solar-

each chapter’s content. powered animals that live in the ocean: sea slugs and corals. But
marine invertebrates are not the only animals with this unusual
lifestyle. The eggs of spotted salamanders have live-in algae of
Embryo

12,041 dpm* 294.5 ng

Carbon fixation
by algae in egg
their own.
Each case concludes with critical thinking questions Spotted salamanders are amphibians that live throughout the
forests of North America. On rainy evenings in the spring, these Embryo alone

that can be used as an in-class group activity. The animals mate in temporary ponds, where the females lay masses
of fertilized eggs (figure 5.12). Each egg contains a tiny embryo
627 dpm* 15.4 ng
Transfer of
sugars from
algae to embryo

studies touch on concepts found in other units; you can and is surrounded by a thick jelly layer. Then, something unusual
happens: Microscopic green algae somehow find each egg and
enter the jelly layer. The algae reproduce and carry out photosyn-
*dpm = disintegrations per minute, a measure of radioactivity

encourage students to draw a concept map illustrating the thesis in the protective confines of their new homes.
Biologists have known since 1986 that the green algae boost
Figure 5.13 Thanks for the Snack. Using a radioactive isotope of
carbon, researchers measured the amount of carbon transferred from

relationships between ideas. You might also use the case the O2 concentration inside the salamander eggs, a real benefit to egg-dwelling green algae to salamander embryos.
embryos that cannot yet breathe on their own. But do algae also

as a basis for discussion of the nature of science.

feed the embryos a steady diet of sugars? Erin Graham, Robert
Sanders, and two other researchers at Temple University wanted transfer some of the radioactive sugar to the embryos.
to learn more. i diffusion, section 4.5A
The team gathered algae-infected eggs from the wild and in- To differentiate between these two possibilities, the team in-

Connect interactive and test bank questions focus on cubated them for nearly 2 hours in a solution containing CO2 that
was “tagged” with a radioactive isotope of carbon. They knew
cubated some eggs in the light, allowing both diffusion and pho-
tosynthesis to occur. A second set of eggs was incubated in total
darkness. Photosynthesis is not possible in the dark, but diffusion
the Investigating Life cases. Questions assess students’
that only algae—not salamander eggs or embryos—would be
able to use CO2 directly in photosynthesis. After the incubation continues. Subtracting the amount of radioactive carbon in dark-
period, they rinsed off the excess solution and measured the treated embryos from the amount in light-treated embryos should
understanding of the science behind the Investigating amount of radioactive carbon in each egg and embryo. i radio-
active isotopes, section 2.1C
therefore reveal the effect of photosynthesis.
After the experiment was complete, radioactivity measure-

Life case and their ability to integrate those concepts with The researchers reasoned that any radioactive carbon in a
salamander embryo could come from one of two sources. The
carbon might simply diffuse in from the solution, without any
information from other units. help from the algae. Alternatively, the algae might use the
tagged carbon to produce sugars in photosynthesis, then
102 UNIT ONE Science, Chemistry, and Cells
ments revealed that eggs and embryos incubated in the light in-
corporated more radioactive carbon than did their dark-treated
counterparts, a sure sign that the algae were sharing their carbon
with their tiny hosts (figure 5.13). This sugar supplement can
help a developing embryo survive.
It is not surprising that animals would take on photosynthet-
ic partners, but what’s in it for the algae? They probably benefit
from the partnership as well. A developing embryo releases CO2
in respiration (see chapter 6). Perhaps this extra shot of CO2
makes photosynthesis more efficient for the algae, completing
the exchange of materials between two allies from different king-
doms of life.
Source: Graham, Erin R., Scott A. Fay, Adam Davey, and Robert W. Sanders. 2013.
Intracapsular algae provide fixed carbon to developing embryos of the salamander
Ambystoma maculatum. Journal of Experimental Biology, vol. 216, pages 452–459.

CH A P T E R S UM M ARY Leaf Cell

5.7 MASTERING CONCEPTS
5.1 Life Depends on Photosynthesis Chloroplast 1. On average, what percentage of the 294.5 ng of carbon that the
∙ Autotrophs produce their own organic compounds from inorganic algae fix each hour is transferred to the embryo? Refer to
starting materials such as CO2 and water. Heterotrophs rely on organic figure 5.13.
molecules produced by other organisms.
Figure 5.12 Spawning. A spotted salamander lays eggs in a pool 2. Identify a standardized variable, an independent variable, and a
A. Photosynthesis Builds Carbohydrates Out of Carbon Dioxide and Water
of water. dependent variable in the experiment.
∙ Photosynthesis converts kinetic energy in light to potential energy in the Granum ©George Grall/National Geographic Creative
Light reactions
covalent bonds of carbohydrates, according to the following chemical (in thylakoid
equation: membranes) Stroma
light energy Light
6CO2 + 6H2O C6H12O6 + 6O2 Light Chlorophyll
energy energy
B. Plants Use Carbohydrates in Many Ways + H NADP++ H+ NADPH
∙ Plants use glucose and other sugars to grow, generate ATP, nourish
nonphotosynthetic plant parts, and produce cellulose and many other
biochemicals. Most store excess carbohydrates as starch or sucrose. H2O H+
2e–
C. The Evolution of Photosynthesis Changed Planet Earth
1/2 O + 2H+ H+
∙ Before photosynthesis evolved, organisms were heterotrophs. The first 2 H+
autotrophs made new food sources available. 3 CO2
∙ Over billions of years, oxygen produced in photosynthesis changed H+
Earth’s climate and the history of life. Rubisco
ADP + P enzyme
5.2 Sunlight Is the Energy Source for Photosynthesis ATP
Carbon
A. What Is Light? reactions ATP
∙ Visible light is a small part of the electromagnetic spectrum. ADP + P
(in stroma)
∙ Photons move in waves. The shorter the wavelength, the more kinetic NADPH
energy per photon. Visible light occurs in a spectrum of colors NADP+ + H+
representing different wavelengths. ATP ADP + P
B. Photosynthetic Pigments Capture Light Energy
∙ Chlorophyll a is the primary photosynthetic pigment in plants. Accessory 1 P
pigments absorb wavelengths of light that chlorophyll a cannot absorb. PGAL
C. Chloroplasts Are the Sites of Photosynthesis
∙ Plants exchange gases with the environment through pores called stomata.
∙ Leaf mesophyll cells contain abundant chloroplasts.
Figure 5.14 Light
and Carbon Reactions.
Glucose, starch,
sucrose The Chapter Summary highlights key points
∙ A chloroplast contains a gelatinous fluid called the stroma. The fluid
surrounds the grana, which are stacks of pancake-shaped thylakoid
membranes. Photosynthetic pigments are embedded in the thylakoid
membranes, which enclose the thylakoid space.
5.5 The Carbon Reactions Produce Carbohydrates
and terminology from the chapter.
∙ A photosystem consists of proteins, antenna pigments, and a reaction ∙ The carbon reactions use energy from ATP and electrons from NADPH
center. in carbon fixation reactions that add CO2 to organic compounds.
∙ In the Calvin cycle, rubisco catalyzes the reaction of CO2 with ribulose
5.3 Photosynthesis Occurs in Two Stages bisphosphate (RuBP) to yield two molecules of PGA. These are
∙ The light reactions of photosynthesis produce ATP and NADPH; these converted to PGAL, the immediate product of photosynthesis. PGAL
molecules provide energy and electrons for the sugar-producing carbon later becomes glucose and other carbohydrates.
reactions (figure 5.14).
5.6 C3, C4, and CAM Plants Use Different Carbon
5.4 The Light Reactions Begin Photosynthesis Fixation Pathways
A. Light Striking Photosystem II Provides the Energy to Produce ATP ∙ The Calvin cycle is also called the C3 pathway. Most plant species are
∙ Photosystem II captures light energy and sends electrons from reactive C3 plants, which use only this pathway to fix carbon.
chlorophyll a along the electron transport chain. ∙ Photorespiration wastes carbon and energy when rubisco reacts with O2
∙ Electrons from chlorophyll are replaced with electrons from water. O2 is instead of CO2.
the waste product. ∙ The C4 pathway reduces photorespiration by separating two carbon
∙ The energy released in the electron transport chain drives the active fixation reactions into different cells. In mesophyll cells, CO2 is fixed as
transport of protons (H+) into the thylakoid space. The protons diffuse a four-carbon molecule, which moves to a bundle-sheath cell and
out through channels in ATP synthase. This movement powers the liberates CO2 to be fixed again in the Calvin cycle.
phosphorylation of ADP to ATP. ∙ In the CAM pathway, desert plants such as cacti open their stomata and
∙ The coupling of the proton gradient and ATP formation is called take in CO2 at night, storing the fixed carbon in vacuoles. During the
chemiosmotic phosphorylation. day, they split off CO2 and fix it in chloroplasts in the same cells.
B. Electrons from Photosystem I Reduce NADP+ to NADPH
∙ Light striking photosystem I re-energizes the electrons, which pass to an 5.7 Investigating Life: Solar-Powered Salamanders
enzyme that uses them to reduce NADP+. The product of this reaction is ∙ The eggs of spotted salamanders contain cells of green algae, which
NADPH. provide O2 and carbon to the animal’s embryo.
x AUTHOR’S GUIDE

CHAPTER 5 Photosynthesis 95
Write It Out and Mastering Concepts
5.3 Photosynthesis Light
H2O CO2 questions are useful for student review or as
Occurs in Two Stages
Inside a chloroplast, photosynthesis occurs in two stages: the
Chloroplast
short in-class writing assignments.
light reactions and the carbon reactions. Figure 5.7 summarizes ATP

I compile them into a list of Guided Reading Questions

the entire process, and sections 5.4 and 5.5 describe each part in
greater detail. Light NADPH Carbon
The light reactions convert solar energy to chemical energy. reactions
NADP+ reactions
(You can think of the light reactions as the “photo-” part of pho-
tosynthesis.) In the chloroplast’s thylakoid membranes, pigment
ADP
that help students focus on material I cover in class. I
molecules in two linked photosystems capture kinetic energy
from photons and store it as potential energy in the chemical O2 Sugar
also use them as discussion questions in Action Centers,
bonds of two molecules: ATP and NADPH.
Recall from chapter 4 that ATP is a nucleotide that stores po-
tential energy in the covalent bonds between its phosphate groups. Figure 5.7 Overview of Photosynthesis. In the light reactions, pigment where students can come for additional help with course
ATP forms when a phosphate group is added to ADP (see figure 4.9).
The other energy-rich product of the light reactions, NADPH, is a
molecules capture light energy and transfer it to molecules of ATP and
NADPH. The carbon reactions use this energy to build sugar molecules out of
carbon dioxide.
material.
coenzyme that carries pairs of energized electrons. In photosynthe-
sis, these electrons come from one of the two reaction center chloro-
phyll molecules. Once the light reactions are under way, chlorophyll, Because the carbon reactions do not directly require light,
in turn, replaces its “lost” electrons by splitting water molecules, they are sometimes called the “dark reactions” of photosynthesis.
yielding O2 as a waste product. i coenzymes, section 4.4B This term is misleading, however, because the carbon reactions
These two resources (energy and “loaded” electron carriers) occur mostly during the day, when the light reactions are produc-
set the stage for the second part of photosynthesis: the carbon ing ATP and NADPH. A more accurate alternative would be the
reactions. In the carbon reactions, the chloroplast uses ATP, the “light-independent reactions.”
high-energy electrons in NADPH, and CO2 to produce sugar
molecules. (These reactions are the “-synthesis” part of photo- 5.3 MASTERING CONCEPTS
synthesis.) The ATP and NADPH come from the light reactions, 1. What happens in each of the two main stages of photosynthesis?
and the CO2 comes from the atmosphere. Once inside the leaf,
CO2 diffuses into a mesophyll cell and across the chloroplast
2. Explain the role of each of the products of the light reactions and CHAPTER 5 Photosynthesis 93
the carbon reactions.
membrane into the stroma, where the carbon reactions occur.

Burning Question Leaf

Chlorophyll a Sunlight
Why do leaves change colors in the fall?
Chlorophyll b
Most leaves are80 Carotenoids
green throughout a plant’s growing season, al- These carefully timed Reflected
events help the plant conserve resources.
Relative absorption (percent)

though there are exceptions; some ornamental plants, for example,
have yellow or purple foliage. The familiar green color comes
from chlorophyll a, the most abundant pigment in photo-
synthetic plant parts. But the leaf also has other photo-
60 Carotenoids contribute brilliant
synthetic pigments.
yellow, orange, and red hues. Purple pigments, such
as anthocyanins, are not photosynthetically active,
but they do protect leaves from damage by ultraviolet
radiation.
Carotenoids40 are less abundant than chlorophyll, so
they usually remain invisible to the naked eye during the
growing season. As winter approaches, however, deciduous
plants prepare to shed their leaves. Anthocyanins accumulate while
chlorophyll degrades, and the now “unmasked” accessory pigments
20
reveal their colors for a short time as a spectacular autumn display.
These pigments soon disappear as well, and the dead leaves turn brown
After all, about 75% of a leaf’s light
proteins occur in its chloroplasts.
Rather than simply letting the first frost kill the leaves, the plant
dismantles these proteins before the leaves die. The plant
stores the valuable nitrogen and other nutrients from these
molecules in living tissues that will survive the winter.
a. Spring brings a flush of fresh, green leaves. The Mesophyll
energy to produce the foliage comes from glucose the
plant produced during the last growing season and cells
stored as starch. The new leaves make food throughout
the spring and summer, so the tree can grow—both above
the ground and below—and produce fruits and seeds. As the
©Corbis RF days grow shorter and cooler in autumn, the cycle will con-
tinue, and the colorful pigments will again participate in one of na-
ture’s great disappearing acts.

Submit your burning question to

Burning
Stoma Questions cover topics that students
CO
and fall to the ground. Marielle.Hoefnagels@mheducation.com
a. wonder about. O +HO 2
2 2
0
400 500 600 700
I ask my students to write down a Burning Question on
Wavelength of light (nanometers)
b.
Mesophyll cell the first day of class. I answer all of them during the
Figure 5.4 Everything but Green. (a) Overall, a leaf reflects green and semester, whenever a relevant topic comes up in class.
yellow wavelengths of light and absorbs the other wavelengths. (b) Each type
Figure
of pigmentIt Outsomequestions
absorbs reinforce
wavelengths of light chapter
and reflects others. Nucleus
concepts and typically have numeric
lobster shells, and the flesh of salmon all owe their distinctive
Central
vacuole
answers (supporting student
colors to carotenoid pigments, which math
the animals must obtain Mitochondrion Chloroplasts

skills).
from their diets.)
Only absorbed light is useful in photosynthesis. Accessory
pigments absorb wavelengths that chlorophyll a cannot, so they
Students can work on these in small groups, in
extend the range of light wavelengths that a cell can harness.
TEM
b. 15 μm (false color)
class,
This or
is ain Action
little like theCenter.
membersMost
of the could easily
same team on a be
quiz
used as each
show, clicker questions
contributing as well.
answers from a different area of
expertise.
Chloroplast DNA Outer
membrane
Figure It Out Inner
membrane
If you could expose plants to just one wavelength of light at a time, would a
wavelength of 300 nm, 450 nm, or 600 nm produce the highest photosynthetic rate?
Answer: 450 nm

Stroma
Granum Ribosomes
c.
C. Chloroplasts Are the Sites
Thylakoid
of Photosynthesis membrane with
In plants, leaves are the main organs of photosynthesis. Their embedded
Granum pigments
broad, flat surfaces expose abundant surface area to sunlight. But
light is just one requirement for photosynthesis. Water is also
essential; roots absorb this vital ingredient, which moves up
Thylakoid
stems and into the leaves. And plants also exchange CO2 and O2 space
with the atmosphere. How do these gases get into and out of
leaves?
The answer is that CO2 and O2 enter and exit a plant
through stomata (singular: stoma), tiny openings in the epi-
d.
dermis of a leaf or stem (figure 5.5a). Stomata allow for gas
exchange, but water evaporates through the same openings. Figure 5.5 Leaf and Chloroplast Anatomy. (a) The tissue inside a leaf
When the plant loses too much water, pairs of specialized is called mesophyll. (b) Each mesophyll cell contains many chloroplasts. (c) A
“guard cells” surrounding each stoma collapse against one an- chloroplast contains light-harvesting pigments, embedded in (d) the stacks of
other, closing the pores. Stomata therefore help balance the thylakoid membranes that make up each granum.
competing needs of gas exchange and water conservation. Photos: (leaves): ©Steve Raymer/National Geographic Stock; (mesophyll): Electron micrograph by
i leaf epidermis, section 22.2B Wm. P. Wergin, courtesy of Eldon H. Newcomb, University of Wisconsin-Madison
Animated Tutorials Explain Complicated Topics
Animated tutorials guide students through
complicated topics, using illustrations and
examples from the book.
We created these tutorials to walk students through the most
difficult material, step by step. Each tutorial places the topic
in context, explains one or more concepts and related figures
taken directly from the Hoefnagels text, and returns to the
big picture at the end. You can assign the tutorials with
accompanying critical thinking questions from the interactive
question banks, or you can use the tutorials embedded in
PowerPoint® slides in your presentations.
Your students can review the tutorials through SmartBook.
Topics are listed below.
Organization of Life Genetic Variation: The Basis of Natural Organ System Interactions
Scientific Method and Interpreting a Selection Example of Negative Feedback
Graph Understanding the Hardy–Weinberg Action Potential
Chemical Bonding Equation The Synapse
Dehydration Synthesis and Hydrolysis Evidence for Evolution Overview of the Senses
Protein Structure Evidence for Human Evolution Sense of Vision
Anatomy of a Cell Membrane Radiometric Dating Sense of Hearing
ATP Reading an Evolutionary Tree Cell Responses to Hormones
Enzymes Origin of Life Role of ATP in Muscle Contraction
Reaction Energetics Endosymbiont Theory The Heartbeat
Osmosis Viral Replication Respiratory Surfaces
Cell Structure Lytic and Lysogenic Cycles Digestion and Food Molecules
Overview of Photosynthesis Replication of HIV Nephron Function
Light Reactions Prokaryote Diversity Adaptive Immunity
The Calvin Cycle Protist Diversity Allergies
Overview of Respiration Plant Diversity Oogenesis
Mitochondrial Electron Transport Chain Moss Reproductive Cycle Human Male Reproductive System
Fermentation Fern Reproductive Cycle Human Female Reproductive System
Protein Synthesis Conifer Reproductive Cycle Ovarian and Menstrual Cycles
Overview of DNA Replication Sexual Reproduction in Angiosperms Proximate and Ultimate Behaviors
Stages of Mitosis Basidiomycete Reproductive Cycle Population Growth Models
Stages of Meiosis Diversity of Fungi Biomagnification
Comparison of Mitosis and Meiosis Animal Diversity Water Cycle
Crossing Over Overview of Plant Tissues Nitrogen Cycle
Nondisjunction Phloem Sap Transport Phosphorus Cycle
Homologous Chromosomes Water Movement Through the Xylem Carbon Cycle
Constructing and Interpreting a Punnett Alternation of Generations Earth’s Climate and Biomes
Square Fruit Development CO2 and Earth’s Average Temperature
DNA Profiling Auxin and Phototropism Threats to Biodiversity
Mechanisms of Evolution Overview of Animal Tissues

xi
 Required=Results
©Getty Images/iStockphoto

McGraw-Hill Connect®
Learn Without Limits
Connect is a teaching and learning platform that
is proven to deliver better results for students and
instructors.
Connect empowers students by continually adapting
to deliver precisely what they need, when they need
it, and how they need it, so your class time is more
engaging and effective.

73% of instructors who use

Connect require it; instructor Using Connect improves retention
satisfaction increases by 28% rates by 19.8%, passing rates by
when Connect is required. 12.7%, and exam scores by 9.1%.

Analytics
Connect Insight®
Connect Insight is Connect’s new one-of-a-kind
visual analytics dashboard—now available for both
instructors and students—that provides at-a-
glance information regarding student performance,
which is immediately actionable. By presenting assignment,
assessment, and topical performance results together with
a time metric that is easily visible for aggregate or individual
results, Connect Insight gives the user the ability to take a
just-in-time approach to teaching and learning, which was Students can view
never before available. Connect Insight presents data that their results for any
empowers students and helps instructors improve class
performance in a way that is efficient and effective.
Connect course.

Mobile
Connect’s new, intuitive mobile interface gives students and
instructors flexible and convenient, anytime–anywhere access to all
components of the Connect platform.
Adaptive
THE ADAPTIVE
READING EXPERIENCE
DESIGNED TO TRANSFORM
THE WAY STUDENTS READ

More students earn A’s and

B’s when they use McGraw-Hill
Education Adaptive products.

SmartBook®
Proven to help students improve grades and
study more efficiently, SmartBook contains the
same content within the print book, but actively
tailors that content to the needs of the individual.
SmartBook’s adaptive technology provides precise,
personalized instruction on what the student
should do next, guiding the student to master
and remember key concepts, targeting gaps in
knowledge and offering customized feedback,
and driving the student toward comprehension
and retention of the subject matter. Available on
tablets, SmartBook puts learning at the student’s
fingertips—anywhere, anytime.

Over 8 billion questions have

been answered, making McGraw-
Hill Education products more
intelligent, reliable, and precise.

www.mheducation.com
Acknowledgments
It takes an army of people to make a textbook, and while I don’t
work with everyone directly, I greatly appreciate the contributions
of each person who makes it possible.
Matt Taylor has worked alongside me at every stage, from
first draft to finished product; in addition, he has seamlessly
­integrated the book’s approach into our digital assets. His careful
work and insights are invaluable.
I appreciate the help of my colleagues at The University of
Oklahoma, including Doug Gaffin, Ben Holt, Michael Markham,
and J. P. Masly. Helpful colleagues from other institutions i­ nclude
Marjorie Weber.
I am grateful to former OU student Emily North, who spent
many hours scrutinizing art and page layouts. In addition, con-
versations with students in my classes spark many good ideas.
My team at McGraw-Hill is wonderful. Thank you to Man-
aging Director Lynn Breithaupt and Executive Brand Manager
Michelle Vogler, who help us create the best book and digital
resources possible. Product Developer Anne Winch retains her
amazing ability to juggle an ever-increasing slate of tasks, all
Changes by Chapter
UNIT 1 Science, Chemistry, and Cells
∙∙Chapter 1 (The Scientific Study of Life): Updated fig-
ure 1.3 to use memories as examples of emergent properties;
improved figure 1.12 to more clearly define independent and
dependent variables; within table 1.2, clarified definitions of
independent variable, dependent variable, and standardized
variable. Added several learning tools to SmartBook®: a fig-
ure depicting a simple evolutionary tree in section 1.2; a figure
showing the taxonomic hierarchy in section 1.2; a miniglossary
of scientific knowledge in section 1.3; a figure describing how
to interpret graphs in the chapter summary.
∙∙Chapter 2 (The Chemistry of Life): In table 2.1, clarified that
the mass of an electron is not exactly zero; added figure 2.3,
which compares a neutral hydrogen atom with H+; improved
description of electron orbitals and energy shells; improved
explanation of polar covalent bonds in water molecules in nar-
rative and in figure 2.10; clarified definition of hydrophobic;
updated illustration of cellulose in figure 2.18 and illustration
xiv
while remaining both responsive and funny. Chris L ­ oewenberg,
Britney Ross, and Jenna Paleski contribute energy and great
ideas to the marketing side. Emily Tietz continues to provide
outstanding service in photo selections. I also appreciate ­Angie
Fitzpatrick, April Southwood, and Peggy Selle for their im-
­
pressive skills at the interface between us and the production
team. Also among the talented folks at McGraw-Hill are E ­ mily
­Windelborn, Tara ­McDermott, Lori Hancock, Lorraine Buczek,
and Jane Peden, who has made life easier in countless ways.
Thanks to all of you for all you do.
My family and friends continue to encourage me. Thank you
to my parents and sister for their pride and support. I also thank
my friends Kelly Damphousse, Ben and Angie Holt, Michael
Markham and Kristi Isacksen, Karen and Bruce Renfroe, Ingo
and Andrea Schlupp, Clarke and Robin Stroud, Matt Taylor and
Elise Knowlton, and Mark Walvoord. Cats Sidecar, Smudge, and
­Snorkels were worthy companions in my office as well. Finally, my
husband, Doug Gaffin, is always there for me, helping in countless
large and small ways. I could not do this work without him.
of triglycerides in figure 2.25; added figure 2.26, which shows
the difference between solid and liquid lipids; wrote new Inves-
tigating Life about chemical defenses in tawny crazy ants.
∙∙Chapter 3 (Cells): Based on SmartBook user data, clarified
that high surface area to volume improves cell transport effi-
ciency; improved Apply It Now box to teach students that most
of their cells are not their own; replaced Burning Question
with a new box on artificial cells; clarified distinction between
­cytoplasm and cytosol in narrative and several figures; in fig-
ure 3.10, improved accuracy of double bond in unsaturated
fatty acid; adjusted figures 3.15 and 3.18 for accuracy; clarified
the function of plasmodesmata. Added a new learning tool to
­SmartBook: a table comparing mitochondria and chloroplasts
in section 3.4.
∙∙Chapter 4 (The Energy of Life): In chapter opening essay,
added how human evolutionary history affects our food cravings;
improved illustration of potential and kinetic energy in figure 4.1;
based on SmartBook user data, clarified definition of heat; updated
figure 4.10 to clearly show uses of ATP; improved explanations of

negative and positive feedback; in figure 4.18, corrected position
of cell membranes in plant cells in hypertonic surroundings; wrote
new Apply It Now box on boosting metabolism; added aquaporins
to Burning Question on headaches, to improve the connection to
the chapter; wrote new Investigating Life on membrane proteins
in electric fish; improved clarity of summary figure 4.24. Added a
new learning tool to SmartBook: in the chapter summary, a mini-
glossary defining metabolism terms.
∙∙Chapter 5 (Photosynthesis): Based on SmartBook user data,
changed “oxidized” to “consumed” in section 5.1C to emphasize
that the first organisms obtained C from their surroundings; in
narrative and figure 5.8, clarified that the photosynthetic elec-
tron transport chain includes the entire pathway from photosys-
tem II to the formation of NADPH; improved figure 5.9 to show
the fates of the carbohydrates produced in the carbon reactions;
clarified that rubisco is not involved in the C4 pathway; based
on SmartBook user data, explained why C4 plants have greater
water use efficiency than C3 plants; clarified the distinction
between C4 and CAM plants; wrote new Investigating Life on
carbon translocation from algae to salamander embryos. Added
new learning tools to SmartBook: a miniglossary of leaf anatomy
in section 5.2; a miniglossary of light reactions in section 5.4.
∙∙Chapter 6 (Respiration and Fermentation): Changed chapter
title to “Respiration and Fermentation” to better match the title
of chapter 5; improved consistency of use of the terms hydrogen
ion gradient, H+ gradient, and proton gradient; updated Burning
Question on diet pills. Made several changes based on Smart-
Book user data: clarified the paragraph debunking the myth
that lactic acid causes muscle soreness after intense exercise;
modified figure 6.9 to show how glycerol and some amino acids
enter metabolism and to show that nitrogen is stripped from
amino acids and eliminated as waste; improved explanation of
why aerobic respiration must have evolved after O2-generating
photosynthesis. Added several learning tools to SmartBook: a
table in section 6.3 showing where the reactions of respiration
occur; a miniglossary of mitochondrion anatomy in section 6.3;
a miniglossary of aerobic respiration in section 6.5.
UNIT 2 DNA, Inheritance, and
Biotechnology
∙∙Chapter 7 (DNA Structure and Gene Function): Clarified in
section 7.3 that DNA must be “unpacked” for the cell to use its
genetic information; based on SmartBook user data, explained
that terminator sequence is part of DNA, not RNA; improved
definition of promoter in narrative and position of promoter in
figure 7.10; extended “cookbook” analogy to the participants
in translation; improved description of the lac operon; added
information about epigenetics; made an explicit connection
between transcription factors and signal transduction; clarified
bold-faced terms related to mutations; reworked figure 7.24
(Investigating Life) to show an evolutionary tree of FOXP2 pro-
tein changes; added summary figure 7.26 to show three types of
RNA. Added a new learning tool to SmartBook: a miniglossary
of protein synthesis in the chapter summary.
CHANGES BY CHAPTER xv
∙∙Chapter 8 (DNA Replication, Binary Fission, and Mitosis):
Made many small clarifications to the narrative describing chro-
mosome structure and the events of cell division; improved fig-
ure 8.8 to include a yarn analogy of DNA compaction; based
on SmartBook user data, clarified that a compacted chromo-
some is unavailable for transcription and improved the defini-
tion of semiconservative replication; added information about
epigenetics in relation to cancer; improved the list of ways that
cancer cells differ from normal cells; updated figure showing
ways to reduce cancer risk. Added new learning tools to Smart-
Book: miniglossary of cell division in section 8.1; table compar-
ing binary fission and mitotic cell division in chapter summary.
∙∙Chapter 9 (Sexual Reproduction and Meiosis): In chapter
opening essay, added possible implications of fetal screening
on human evolution; increased relevance of box on mules by
explaining why mules are desirable; clarified the use of the
word align in talking about the events of meiosis and the origin
of genetic variation; revised box on multiple births to focus on
their rising incidence; improved connection between problems
during meiosis and abnormalities in chromosome number and
structure; added aneuploid cell as a contrast to polyploid cell.
Added several learning tools to SmartBook: a miniglossary of
variability in meiosis in section 9.5; a table comparing asexual
and sexual reproduction in section 9.6; a miniglossary of chro-
mosome abnormalities in section 9.7.
∙∙Chapter 10 (Patterns of Inheritance): Reworked allele des-
ignations for all figures and narrative relevant to yellow and
green peas; improved connection between proteins and traits;
replaced figure 10.14 to show how genotypic ratios differ in
crosses between linked and unlinked genes; based on Smart-
Book user data, clarified explanation of product rule; related
epigenetics to environmental effects on gene expression; based
on SmartBook user data, explained why males cannot be
symptomless carriers of X-linked traits; revised Investigating
Life to incorporate information about next-generation Bt cot-
ton; improved lightbulb analogy of dominance relationships
in figure 10.34. Added several new learning tools to Smart-
Book: a miniglossary of tracking inheritance in section 10.3;
a miniglossary of gene linkage in section 10.5; a miniglossary
of dominance relationships in section 10.6; a miniglossary of
modes of inheritance in section 10.8.
∙∙Chapter 11 (DNA Technology): Added CRISPR as an exam-
ple of a new DNA technology to chapter opening essay; based
on SmartBook user data, clarified that scientists use DNA to
reveal species relationships; updated figure 11.5 to show mod-
ern DNA sequencing; added figure 11.6, which shows similar-
ity between a gene of humans and a homologous gene in other
species; improved application of DNA profiling in figure 11.9;
revised description of somatic cell nuclear transfer; based on
SmartBook user data, clarified that pseudogenes are noncod-
ing DNA and that gene therapy provides supplemental DNA
(not replacement DNA); wrote new Investigative Life on gene
transfer between GMOs and their wild relatives. Added a new
learning tool to SmartBook: a table listing some additional uses
of DNA analysis in section 11.2.
xvi CHANGES BY CHAPTER
UNIT 3 The Evolution of Life
∙∙Chapter 12 (The Forces of Evolutionary Change): Updated
chapter opening essay to mention CDIFF and CRE, two of the
three most dangerous antibiotic-resistant bacteria; improved
figure 12.8 to connect natural selection with mutations in DNA;
reworked Burning Question to explain why there is no such
thing as a “pinnacle of evolution”; improved narrative and figure
explaining Hardy–Weinberg equilibrium; clarified descriptions
of genetic drift and nonrandom mating; based on SmartBook
user data, clarified the effect of gene flow on genetic diversity;
reworked Pull It Together (figure 12.24) to explain how each
mechanism of evolution affects allele frequencies. Added new
learning tools to SmartBook: a miniglossary of populations and
evolution in section 12.1; a miniglossary of evolutionary mech-
anisms in section 12.7.
∙∙Chapter 13 (Evidence of Evolution): In figure 13.2, replaced
Tertiary Period with Paleogene and Neogene; added mole eyes
as an example of a vestigial structure in figure 13.11; expanded
the list of vestigial structures in the narrative; based on Smart-
Book user data, improved figure 13.17 to better show how
mutations in enhancers affect gene expression; in Investigating
Life, clarified evidence that Najash was terrestrial. Added sev-
eral learning tools to SmartBook: a miniglossary of fossil aging
terms in section 13.2; a miniglossary of comparative anatomy
terms in section 13.4; a figure showing all five lines of evidence
for evolution in the chapter summary.
∙∙Chapter 14 (Speciation and Extinction): Reorganized sec-
tion on gradualism and punctuated equilibrium for clarity;
added figure 14.17, which distinguishes ancestral and derived
features; improved figure 14.18 to more clearly explain the
anatomy of a phylogenetic tree; revised Burning Question to
explain how each condition boosts the evolution rate; wrote
new Investigating Life on ecological interactions that boost
speciation rates. Added several learning tools to SmartBook:
a miniglossary of macroevolution in section 14.1; a miniglos-
sary of reproductive barriers in section 14.2; a miniglossary of
speciation patterns in section 14.3.
∙∙Chapter 15 (The Origin and History of Life): Improved
illustrations of primary and secondary endosymbiosis in fig-
ure 15.9; rearranged section on human evolution for clarity;
based on SmartBook user data, clarified distinction between
“early” and “recent” Homo; replaced Investigating Life with
an essay about the human and chimpanzee genome sequencing
projects. Added a new learning tool to SmartBook: a table sum-
marizing biodiversity changes over time in section 15.3.
UNIT 4 The Diversity of Life
∙∙Chapter 16 (Viruses): Revised chapter opening essay to
include the most recent Ebola outbreak; reworked several head-
ings to improve clarity; reorganized paragraphs on viral enve-
lope for clarity; rewrote the passage on latent animal viruses;
based on SmartBook user data, explained which cells are
infected by herpes simplex virus type I (cold sores); improved
figure 16.5, which shows HIV replication; updated Investigat-
ing Life to include newer data. Added a new learning tool to
SmartBook: a miniglossary of viral infections in section 16.3.
∙∙Chapter 17 (Bacteria and Archaea): Based on SmartBook
user data, improved figure 17.5 and clarified description of
Gram-positive and Gram-negative cells; differentiated between
exotoxins and endotoxins; wrote new Investigating Life on
antibiotic-resistant bacteria in pig farms. Added several learn-
ing tools to section 17.2 of SmartBook: a miniglossary of pro-
karyote anatomy; a miniglossary of prokaryote classification; a
miniglossary of gene transfer.
∙∙Chapter 18 (Protists): Based on SmartBook user data, clari-
fied differences between feeding and reproductive stages for
plasmodial slime molds; corrected figure showing overall tree
of life. Added new learning tools to SmartBook: a miniglossary
of types of algae in section 18.2; a table summarizing the life
cycles of plasmodial and cellular slime molds in section 18.3.
∙∙Chapter 19 (Plants): Improved organization of introduction
to section 19.1; clarified description of alternation of genera-
tions; reworked description of double fertilization; improved
description of ovules; added paragraph about gluten sensitivity.
Added a new learning tool to SmartBook: a miniglossary of
plant reproduction in section 19.1.
∙∙Chapter 20 (Fungi): Based on SmartBook user data, clarified
description of figure 20.2; revised Apply It Now: Fungi and
Human Health to focus on infection prevention; improved dis-
tinction between endophytes and mycorrhizal fungi. Added new
learning tools to SmartBook: a miniglossary of fungal anatomy in
section 20.1; a miniglossary of fungal interactions in section 20.7.
∙∙Chapter 21 (Animals): Wrote new chapter opening essay to
emphasize the uses of animal products in everyday objects;
reorganized section 21.1 for clarity; improved definitions of
ectoderm and endoderm; based on SmartBook user data, clar-
ified that both indirect and direct development may occur in
mollusks; also based on SmartBook user data, rearranged sec-
tion on echinoderm defenses to put related content together;
clarified description of amniotic eggs and the term amniote;
reworked and simplified the Investigating Life section. Added
new learning tools to SmartBook: a miniglossary of animal
clades in section 21.1; a miniglossary of arthropod diversity
in section 21.8.
UNIT 5 Plant Life
∙∙Chapter 22 (Plant Form and Function): Added art of ground
tissue cell types in figure 22.4; clarified the description of cells
in phloem tissue; defined primary and secondary growth ear-
lier in the chapter; improved illustrations of stem and root cross
sections in figures 22.9 and 22.13; based on SmartBook user
data, clarified distinction between monocots and eudicots in
leaf cross sections; wrote new Apply It Now box on the topic of
fire-resistant trees and shrubs; based on SmartBook user data,
reworked figure 22.17 to better illustrate the definition of bark.
Added a new learning tool to SmartBook: a miniglossary of
plant anatomy in section 22.3.

∙∙Chapter 23 (Plant Nutrition and Transport): Revised Apply
It Now box on fertilizers; reworked some titles so the words
xylem and phloem are more prominent in the chapter’s main
headers; clarified explanation of sink in description of pressure
flow theory. Added a new learning tool to SmartBook: a mini-
glossary of plant transport in the chapter summary.
∙∙Chapter 24 (Reproduction and Development of ­ Flowering
Plants): Clarified relationship between carpel and ovary;
improved illustration of mature monocot and eudicot seeds in
figure 24.9; based on SmartBook user data, adjusted labeling
on figure depicting corn and bean seed germination; wrote new
Apply It Now box on plants that attack caterpillars; improved
figures 24.22 and 24.23, which illustrate photoperiod’s role in
flowering. Added new learning tools to SmartBook: a miniglos-
sary of angiosperm reproduction in section 24.2; a miniglossary
of plant tropisms in the chapter summary.
UNIT 6 Animal Life
∙∙Chapter 25 (Animal Tissues and Organ Systems): Wrote
new chapter opener on physiological changes that happen as a
person runs a marathon; revised box on organ donation to focus
on artificial organs; revised many glossary terms for consis-
tency; developed new figure 25.14, which summarizes organ
system interactions. Added a new learning tool to SmartBook: a
miniglossary of animal anatomy and physiology in section 25.1.
∙∙Chapter 26 (The Nervous System): Added narrative and glos-
sary definitions for membrane potential; clarified the signifi-
cance of the “all-or-none” nature of an action potential; reworked
the explanation of graded potentials and action potentials; dis-
tinguished between action potential and neural impulse; added
figure 26.7, which illustrates how a neural impulse is similar to
a line of firecrackers exploding; improved explanation of how
the sympathetic nervous system can have both instantaneous
effects and prolonged effects (via adrenal hormones); wrote
new Burning Question on whether neurons communicate at
the speed of light; replaced Investigating Life with an essay on
a grasshopper mouse’s reaction to a scorpion sting. Based on
SmartBook user data, added new learning tools to SmartBook:
a table of action potential events in section 26.3; a miniglossary
of nervous system communication in section 26.3.
∙∙Chapter 27 (The Senses): Wrote new Investigating Life on
taste detection in whales. Based on SmartBook user data, added
new learning tools to SmartBook: a miniglossary of the visual
information pathway in section 27.4; a miniglossary of the
auditory information pathway in section 27.5.
∙∙Chapter 28 (The Endocrine System): Based on SmartBook
user data, clarified in the introduction to section 28.2 that hor-
mone receptors may be on the cell surface, in the cytoplasm,
or in the nucleus; improved the explanation of the overall role
of the hypothalamus and pituitary; reworked section on adre-
nal hormones and their regulation. Added new learning tools to
SmartBook: a miniglossary of hormones and responses in sec-
tion 28.2; a chapter summary table comparing the origins and
functions of many hormones.
CHANGES BY CHAPTER xvii
∙∙Chapter 29 (The Skeletal and Muscular Systems): Added
micrograph to illustration of skeletal muscle organization;
improved figure 29.23, which summarizes the relationship
between muscles and bones. Added several new learning tools
to SmartBook: a miniglossary of the skeletal system in sec-
tion 29.3; a table summarizing the steps of muscle contraction
in section 29.4; a miniglossary of the muscular system in the
chapter summary.
∙∙Chapter 30 (The Circulatory System): Reworked introduc-
tion to section 30.1 for clarity; improved passage describing red
blood cells; clarified description of blood clotting; added infor-
mation on the possible effects of overexercising. Added new
learning tools to SmartBook: a miniglossary of the heartbeat
in section 30.4; a miniglossary of blood vessels in section 30.5.
∙∙Chapter 31 (The Respiratory System): Wrote new chapter
opener on competitive breath-holding; based on SmartBook
user data, clarified the features that all respiratory surfaces
have in common; revised the description of red blood cells’ role
in carrying O2 and CO2; improved section on the functions of
CO2 and blood pH in regulating the breathing rate; updated fig-
ure 31.17 to compare and contrast external and internal respira-
tion. Added a new learning tool to SmartBook: a miniglossary
of breathing in the chapter summary.
∙∙Chapter 32 (Digestion and Nutrition): Reworked fig-
ure 32.9 to emphasize the types of teeth; added information
on essential amino acids and essential fatty acids; wrote new
Burning Question on maximizing the nutrient content of food;
added new table on the calorie content of various beverages;
replaced Investigating Life essay with one about the evolu-
tionary cost of a sweet tooth; reworked Pull It Together (fig-
ure 32.22) to better cover the chapter’s content. Added a new
learning tool to SmartBook: a miniglossary of digestive fluids
in section 32.3.
∙∙Chapter 33 (Regulation of Temperature and Body Fluids):
Clarified the process of urination in the human urinary system.
Added a new learning tool to SmartBook: a miniglossary of
temperature homeostasis in section 33.1.
∙∙Chapter 34 (The Immune System): Reworked figure 34.4,
which provides an overview of innate defenses; clarified the
roles of white blood cells in innate defenses; improved the pas-
sage about cytotoxic T cells; made many small improvements
to the narrative about adaptive immunity; based on SmartBook
user data, clarified the role of MHC proteins; updated informa-
tion about SCID. Added a new learning tool to SmartBook: a
miniglossary of adaptive immunity in section 34.3.
∙∙Chapter 35 (Animal Reproduction and Development):
Wrote new chapter opener about intersex conditions; com-
bined contraception and sexually transmitted diseases in a
new section on reproductive health; clarified the events of
fertilization and prenatal development; improved explanation
of the placenta’s structure and function; wrote a new box on
male pregnancy; made a new summary figure showing the
stages of human development. Added a new learning tool to
SmartBook: a miniglossary on embryonic support structures
in section 35.5.
xviii CHANGES BY CHAPTER
UNIT 7 The Ecology of Life
∙∙Chapter 36 (Animal Behavior): Based on SmartBook user
data, improved figure 36.1 to differentiate between proximate
and ultimate causes of behavior; clarified the definition of
search image; combined multiple figures to make figure 36.11,
which shows many types of defenses against predation. Added
a new learning tool to SmartBook: a miniglossary of innate and
learned behaviors in section 36.2.
∙∙Chapter 37 (Populations): Revised population numbers
and graphs in figures 37.6 and 37.8 for accuracy and clarity;
updated information about China’s former one-child policy.
Added a new learning tool to SmartBook: a miniglossary of
population growth in section 37.3.
∙∙Chapter 38 (Communities and Ecosystems): Wrote new
chapter opening essay about sustainable meat production
and home gardening; improved explanation of coevolution in
crossbills; updated information about mercury in tuna in fig-
ure 38.17. Added new learning tool to SmartBook: a miniglos-
sary of diversity and succession in section 38.2.
∙∙Chapter 39 (Biomes): Based on SmartBook user data, revised
figure 39.4 to show a convection cell from the perspective of
Earth’s surface; improved explanation and illustration of El
Niño in the Apply It Now box; updated Investigating Life
text and figure 39.26 for clarity. Added new learning tools to
SmartBook: a miniglossary of lake zones in section 39.4; a
miniglossary of ocean zones in section 39.5.
∙∙Chapter 40 (Preserving Biodiversity): Updated data on
endangered species; added inset map to figure 40.8 showing
the location of the Great Pacific Garbage Patch; revised sec-
tion on global climate change to include information from the
recent Paris conference; mentioned the sources of methane and
N2O in the atmosphere; corrected the amount of CO2 released
by human activities; updated photo and information about the
ozone hole in figure 40.12; improved figure 40.15 to show how
the Arctic sea ice minimum has changed over time; updated
Pull It Together (figure 40.27) to better illustrate connections
between the topics. Added a new learning tool to SmartBook: a
miniglossary of pollution in section 40.3.
Contents
UNIT 1 Science, Chemistry, and Cells
|
1 The Scientific
Study of Life 2
1.1 What Is Life? 4
A. Life Is Organized 4
B. Life Requires ©Franco Banfi/WaterF/age fotostock
Energy 6
C. Life Maintains Internal Constancy 6
D. Life Reproduces, Grows, and Develops 7
E. Life Evolves 7
1.2 The Tree of Life Includes Three Main Branches 8
1.3 Scientists Study the Natural World 10
A. The Scientific Method Has Multiple Interrelated Parts 10
B. An Experimental Design Is a Careful Plan 11
C. Theories Are Comprehensive Explanations 12
D. Scientific Inquiry Has Limitations 13
3 | Cells 44
E. Biology Continues to Advance 14
1.4 Investigating Life: The Orchid and the Moth 15
2 | The Chemistry of Life 18
2.1 Atoms Make Up All Matter 20
A. Elements Are Fundamental Types of Matter 20
B. Atoms Are Particles of Elements 20
C. Isotopes Have Different Numbers of Neutrons 21
2.2 Chemical Bonds Link Atoms 22
A. Electrons Determine Bonding 22
B. In an Ionic Bond, One Atom Transfers Electrons to
Another Atom 23
C. In a Covalent Bond, Atoms Share Electrons 24
D. Partial Charges on Polar Molecules Create Hydrogen
Bonds 25
2.3 Water Is Essential to Life 26
A. Water Is Cohesive and Adhesive 26
B. Many Substances Dissolve in Water 26
C. Water Regulates Temperature 27
D. Water Expands As It Freezes 27
E. Water Participates in Life’s Chemical Reactions 28
2.4 Cells Have an Optimum pH 28
A. The pH Scale Expresses Acidity or Alkalinity 29
B. Buffers Regulate pH 29
2.5 Cells Contain Four Major Types of Organic Molecules 30
A. Large Organic Molecules Are Composed of Smaller
Subunits 30
B. Carbohydrates Include Simple Sugars and
Polysaccharides 31
C. Proteins Are Complex and Highly Versatile 32
D. Nucleic Acids Store and Transmit Genetic Information 34
E. Lipids Are Hydrophobic and Energy-Rich 36
2.6 Investigating Life: Chemical Warfare on a Tiny
Battlefield 41
3.1 Cells Are the Units of Life 46
A. Simple Lenses Revealed the First Glimpses of Cells 46
B. The Cell Theory Emerges 46
C. Microscopes Magnify Cell Structures 47
D. All Cells Have Features in Common 49
3.2 Different Cell Types Characterize Life’s Three
Domains 50
A. Domain Bacteria Contains Earth’s Most Abundant
Organisms 50
B. Domain Archaea Includes Prokaryotes with Unique
Biochemistry 51
C. Domain Eukarya Contains Organisms with Complex
Cells 52
3.3 A Membrane Separates Each Cell from Its
Surroundings 54
3.4 Eukaryotic Organelles Divide Labor 56
A. The Nucleus, Endoplasmic Reticulum, and Golgi
Interact to Secrete Substances 57
B. Lysosomes, Vacuoles, and Peroxisomes Are Cellular
Digestion Centers 59
xix
xx CONTENTS
C. Mitochondria Extract Energy from Nutrients 60
D. Photosynthesis Occurs in Chloroplasts 60
3.5 The Cytoskeleton Supports Eukaryotic Cells 62
A. Proteins Form the Cytoskeleton 62
B. Cilia and Flagella Help Cells Move 63
3.6 Cells Stick Together and Communicate with One
Another 64
A. Animal Cell Junctions Occur in Several Forms 64
B. Cell Walls Are Strong, Flexible, and Porous 64
3.7 Investigating Life: The Tiniest Compass 67
4 | The Energy of Life 70
4.1 All Cells Capture and Use Energy 72
A. Energy Allows Cells to Do Life’s Work 72
B. The Laws of Thermodynamics Describe Energy
Transfer 73
4.2 Networks of Chemical Reactions Sustain Life 74
A. Chemical Reactions Absorb or Release Energy 74
B. Linked Oxidation and Reduction Reactions Form
Electron Transport Chains 75
4.3 ATP Is Cellular Energy Currency 76
A. Coupled Reactions Release and Store Energy in ATP 76
B. ATP Represents Short-Term Energy Storage 77
4.4 Enzymes Speed Biochemical Reactions 78
A. Enzymes Bring Reactants Together 78
B. Enzymes Have Partners 79
C. Cells Control Reaction Rates 79
4.5 Membrane Transport May Release Energy or Cost
Energy 80
A. Passive Transport Does Not Require Energy Input 80
B. Active Transport Requires Energy Input 83
C. Endocytosis and Exocytosis Use Vesicles to Transport
Substances 83
4.6 Investigating Life: Energy Efficiency in an Electric
Fish 85
5 | Photosynthesis 88
5.1 Life Depends on Photosynthesis 90
A. Photosynthesis Builds Carbohydrates Out of Carbon
Dioxide and Water 90
B. Plants Use Carbohydrates in Many Ways 90
C. The Evolution of Photosynthesis Changed Planet Earth 91
5.2 Sunlight Is the Energy Source for Photosynthesis 92
A. What Is Light? 92
B. Photosynthetic Pigments Capture Light Energy 92
C. Chloroplasts Are the Sites of Photosynthesis 93
5.3 Photosynthesis Occurs in Two Stages 95
5.4 The Light Reactions Begin Photosynthesis 96
A. Light Striking Photosystem II Provides the Energy to
Produce ATP 96
B. Electrons from Photosystem I Reduce NADP+ to
NADPH 97
5.5 The Carbon Reactions Produce Carbohydrates 98
5.6 C3, C4, and CAM Plants Use Different Carbon Fixation
Pathways 99
5.7 Investigating Life: Solar-Powered Salamanders 101
6 | Respiration and Fermentation 104
6.1 Cells Use Energy in Food to Make ATP 106
6.2 Cellular Respiration Includes Three Main Processes 107
6.3 In Eukaryotic Cells, Mitochondria Produce Most
ATP 108
6.4 Glycolysis Breaks Down Glucose to Pyruvate 109
6.5 Aerobic Respiration Yields Abundant ATP 110
A. Pyruvate Is Oxidized to Acetyl CoA 110
B. The Krebs Cycle Produces ATP and Electron
Carriers 110
C. The Electron Transport Chain Drives ATP
Formation 110
6.6 How Many ATPs Can One Glucose Molecule
Yield? 112
6.7 Other Food Molecules Enter the Energy-Extracting
Pathways 113
6.8 Some Energy Pathways Do Not Require Oxygen 114
A. Anaerobic Respiration Uses an Electron Acceptor Other
Than O2 114
B. Fermenters Acquire ATP Only from Glycolysis 115
6.9 Photosynthesis and Respiration Are Ancient
Pathways 116
6.10 Investigating Life: Hot Plants Offer Heat Reward 117
UNIT 2 DNA, Inheritance, and
Biotechnology
7
| DNA Structure
and Gene
Function 120
7.1 Experiments Identified
the Genetic Material 122
©Dr. Gopal Murti/Science Source
A. Bacteria Can Transfer Genetic Information 122
B. Hershey and Chase Confirmed the Genetic Role of
DNA 123
7.2 DNA Is a Double Helix of Nucleotides 124
7.3 DNA Contains the “Recipes” for a Cell’s Proteins 126
A. Protein Synthesis Requires Transcription and
Translation 126
B. RNA Is an Intermediary Between DNA and a
Protein 127

7.4 Transcription Uses a DNA Template to Build RNA 128
A. Transcription Occurs in Three Steps 128
B. mRNA Is Altered in the Nucleus of Eukaryotic Cells 129
7.5 Translation Builds the Protein 130
A. The Genetic Code Links mRNA to Protein 130
B. Translation Requires mRNA, tRNA, and Ribosomes 131
C. Translation Occurs in Three Steps 132
D. Proteins Must Fold Correctly After Translation 133
7.6 Cells Regulate Gene Expression 134
A. Operons Are Groups of Bacterial Genes That Share
One Promoter 134
B. Eukaryotic Organisms Use Many Regulatory
Mechanisms 135
7.7 Mutations Change DNA Sequences 138
A. Mutations Range from Silent to Devastating 138
B. What Causes Mutations? 139
C. Mutations May Pass to Future Generations 140
D. Mutations Are Important 140
7.8 Investigating Life: Clues to the Origin of Language 141
|
8 DNA Replication, Binary Fission,
and Mitosis 146
8.1 Cells Divide and Cells Die 148
A. Sexual Life Cycles Include Mitosis, Meiosis, and
Fertilization 148
B. Cell Death Is Part of Life 148
8.2 DNA Replication Precedes Cell Division 150
10 | Patterns of Inheritance 186
8.3 Prokaryotes Divide by Binary Fission 152
8.4 Chromosomes Condense Before Cell Division 152
8.5 Mitotic Division Generates Exact Cell Copies 153
A. DNA Is Copied During Interphase 154
B. Chromosomes Divide During Mitosis 155
C. The Cytoplasm Splits in Cytokinesis 156
8.6 Cancer Arises When Cells Divide Out of Control 156
A. Chemical Signals Regulate Cell Division 156
B. Cancer Cells Break Through Cell Cycle Controls 157
C. Cancer Cells Differ from Normal Cells in Many
Ways 157
D. Cancer Treatments Remove or Kill Abnormal Cells 159
E. Genes and Environment Both Can Increase Cancer
Risk 160
8.7 Apoptosis Is Programmed Cell Death 162
8.8 Investigating Life: Cutting Off a Tumor’s Supply Lines
in the War on Cancer 163
9 | Sexual Reproduction and Meiosis 166
9.1 Why Sex? 168
9.2 Diploid Cells Contain Two Homologous Sets of
Chromosomes 169
CONTENTS xxi
9.3 Meiosis Is Essential in Sexual Reproduction 170
A. Gametes Are Haploid Sex Cells 170
B. Specialized Germ Cells Undergo Meiosis 170
C. Meiosis Halves the Chromosome Number and
Scrambles Alleles 171
9.4 In Meiosis, DNA Replicates Once, but the Nucleus
Divides Twice 172
A. In Meiosis I, Homologous Chromosomes Pair Up
and Separate 172
B. Meiosis II Yields Four Haploid Nuclei 173
9.5 Meiosis Generates Enormous Variability 174
A. Crossing Over Shuffles Alleles 174
B. Homologous Pairs Are Oriented Randomly During
Metaphase I 174
C. Fertilization Multiplies the Diversity 175
9.6 Mitosis and Meiosis Have Different Functions:
A Summary 176
9.7 Errors Sometimes Occur in Meiosis 177
A. Cells May Inherit Too Many or Too Few
Chromosomes 177
B. Changes in Chromosome Structure May Be
Harmful 178
9.8 Haploid Nuclei Are Packaged into Gametes 180
A. In Humans, Gametes Form in Testes and
Ovaries 180
B. In Plants, Gametophytes Produce Gametes 181
9.9 Investigating Life: Evolving Germs Select for Sex in
Worms 182
10.1 Chromosomes Are Packets of Genetic Information:
A Review 188
10.2 Mendel’s Experiments Uncovered Basic Laws
of Inheritance 189
A. Why Peas? 189
B. Dominant Alleles Appear to Mask Recessive
Alleles 189
C. For Each Gene, a Cell’s Two Alleles May Be Identical
or Different 190
D. Every Generation Has a Name 191
10.3 The Two Alleles of a Gene End Up in Different
Gametes 192
A. The Simplest Punnett Squares Track the Inheritance of
One Gene 192
B. Meiosis Explains Mendel’s Law of Segregation 193
10.4 Genes on Different Chromosomes Are Inherited
Independently 194
A. Tracking Two-Gene Inheritance May Require Large
Punnett Squares 194
B. Meiosis Explains Mendel’s Law of Independent
Assortment 194
C. The Product Rule Is a Useful Shortcut 196
xxii CONTENTS
10.5 Genes on the Same Chromosome May Be Inherited
Together 196
A. Genes on the Same Chromosome Are Linked 196
B. Studies of Linked Genes Have Yielded Chromosome
Maps 197
10.6 Dominance Relationships Are Rarely Simple 199
A. Incomplete Dominance and Codominance Add
Phenotype Classes 199
B. Some Inheritance Patterns Are Especially Difficult
to Interpret 200
10.7 Sex-Linked Genes Have Unique Inheritance
Patterns 201
A. X and Y Chromosomes Carry Sex-Linked Genes 201
B. X-Linked Recessive Disorders Affect More Males Than
Females 202
C. X Inactivation Prevents “Double Dosing” of Proteins 203
10.8 Pedigrees Show Modes of Inheritance 205
10.9 Most Traits Are Influenced by the Environment and
Multiple Genes 207
A. The Environment Can Alter the Phenotype 207
B. Polygenic Traits Depend on More Than One Gene 207
10.10 Investigating Life: Heredity and the Hungry Hordes 209
11 | DNA Technology 216
11.1 DNA Technology Is Changing the World 218
11.2 DNA Technology’s Tools Apply to Individual Genes or
Entire Genomes 219
A. Transgenic Organisms Contain DNA from Other
Species 219
B. DNA Sequencing Reveals the Order of Bases 222
C. PCR Replicates DNA in a Test Tube 223
D. DNA Profiling Detects Genetic Differences 225
11.3 Stem Cells and Cloning Add New Ways to Copy Cells
and Organisms 226
A. Stem Cells Divide to Form Multiple Cell Types 226
B. Cloning Produces Identical Copies of an Organism 226
11.4 Many Medical Tests and Procedures Use DNA
Technology 229
A. DNA Probes Detect Specific Sequences 229
B. Preimplantation Genetic Diagnosis Can Screen
Embryos for Some Diseases 229
C. Genetic Testing Can Detect Existing Diseases 230
D. Gene Therapy Uses DNA to Treat Disease 230
E. Medical Uses of DNA Technology Raise Many Ethical
Issues 231
11.5 Investigating Life: Weeds Get a Boost from Their
Transgenic Cousins 232
UNIT 3 The Evolution of Life
|
12 The Forces of
Evolutionary
Change 236
©Chris Ryan/Getty Images RF
12.1 Evolution Acts on
Populations 238
12.2 Evolutionary Thought Has Evolved for Centuries 238
A. Many Explanations Have Been Proposed for Life’s
Diversity 238
B. Charles Darwin’s Voyage Provided a Wealth of
Evidence 240
C. On the Origin of Species Proposed Natural Selection as
an Evolutionary Mechanism 240
D. Evolutionary Theory Continues to Expand 243
12.3 Natural Selection Molds Evolution 244
A. Adaptations Enhance Reproductive Success 244
B. Natural Selection Eliminates Poorly Adapted
Phenotypes 244
C. Natural Selection Does Not Have a Goal 246
D. What Does “Survival of the Fittest” Really Mean? 246
12.4 Evolution Is Inevitable in Real Populations 248
A. At Hardy–Weinberg Equilibrium, Allele Frequencies
Do Not Change 248
B. In Reality, Allele Frequencies Always Change 249
12.5 Natural Selection Can Shape Populations in Many
Ways 250
12.6 Sexual Selection Directly Influences Reproductive
Success 252
12.7 Evolution Occurs in Several Additional Ways 253
A. Mutation Fuels Evolution 253
B. Genetic Drift Occurs by Chance 253
C. Nonrandom Mating Concentrates Alleles Locally 255
D. Gene Flow Moves Alleles Between Populations 255
12.8 Investigating Life: Size Matters in Fishing Frenzy 256
13 | Evidence of Evolution 260
13.1 Clues to Evolution Lie in the Earth, Body Structures,
and Molecules 262
13.2 Fossils Record Evolution 264
A. Fossils Form in Many Ways 264
B. The Fossil Record Is Often Incomplete 266
C. The Age of a Fossil Can Be Estimated in Two Ways 266
13.3 Biogeography Considers Species’ Geographical
Locations 268
A. The Theory of Plate Tectonics Explains Earth’s Shifting
Continents 268
B. Species Distributions Reveal Evolutionary Events 268

15 | The Origin and History of Life 304
13.4 Anatomical Comparisons May Reveal Common
Descent 270
A. Homologous Structures Have a Shared Evolutionary
Origin 270
B. Vestigial Structures Have Lost Their Functions 270
C. Convergent Evolution Produces Superficial
Similarities 271
13.5 Embryonic Development Patterns Provide Evolutionary
Clues 272
13.6 Molecules Reveal Relatedness 274
A. Comparing DNA and Protein Sequences May Reveal
Close Relationships 274
B. Molecular Clocks Help Assign Dates to Evolutionary
Events 275
13.7 Investigating Life: Limbs Gained and Limbs Lost 276
14 | Speciation and Extinction 280
14.1 What Is a Species? 282
A. Linnaeus Devised the Binomial Naming System 282
B. Species Can Be Defined Based on the Potential to
Interbreed 282
14.2 Reproductive Barriers Cause Species to Diverge 284
A. Prezygotic Barriers Prevent Fertilization 285
B. Postzygotic Barriers Prevent the Development of Fertile
Offspring 285
14.3 Spatial Patterns Define Three Types of Speciation 286
A. Allopatric Speciation Reflects a Geographic
Barrier 286
B. Parapatric Speciation Occurs in Neighboring
Regions 288
C. Sympatric Speciation Occurs in a Shared Habitat 288
D. Determining the Type of Speciation May Be
Difficult 289
14.4 Speciation May Be Gradual or May Occur in
UNIT 4
Bursts 290
A. Gradualism and Punctuated Equilibrium Are Two
Models of Speciation 290
16 | Viruses 330
B. Bursts of Speciation Occur During Adaptive
Radiation 291
14.5 Extinction Marks the End of the Line 292
A. Many Factors Can Combine to Put a Species at Risk 292
B. Extinction Rates Have Varied over Time 292
14.6 Biological Classification Systems Are Based on
Common Descent 294
A. The Taxonomic Hierarchy Organizes Species into
Groups 294
B. A Cladistics Approach Is Based on Shared Derived
Traits 294
C. Cladograms Depict Hypothesized Evolutionary
Relationships 296
D. Many Traditional Groups Are Not Clades 298
14.7 Investigating Life: Plant Protection Rackets May
Stimulate Speciation 300
CONTENTS xxiii
15.1 Life’s Origin Remains Mysterious 306
A. The First Organic Molecules May Have Formed in a
Chemical “Soup” 306
B. Some Investigators Suggest an “RNA World” 309
C. Membranes Enclosed the Molecules 309
D. Early Life Changed Earth Forever 309
15.2 Complex Cells and Multicellularity Arose over a Billion
Years Ago 311
A. Endosymbiosis Explains the Origin of Mitochondria
and Chloroplasts 311
B. Multicellularity May Also Have Its Origin in
Cooperation 312
15.3 Life’s Diversity Exploded in the Past 500 Million
Years 314
A. The Strange Ediacarans Flourished Late in the
Precambrian 314
B. Paleozoic Plants and Animals Emerged onto Land 314
C. Reptiles and Flowering Plants Thrived During the
Mesozoic Era 317
D. Mammals Diversified During the Cenozoic Era 318
15.4 Fossils and DNA Tell the Human Evolution Story 320
A. Humans Are Primates 320
B. Molecular Evidence Documents Primate
Relationships 322
C. Human Evolution Is Partially Recorded in Fossils 323
D. Environmental Changes Have Spurred Human
Evolution 324
E. Migration and Culture Have Changed Homo
sapiens 325
15.5 Investigating Life: What Makes Us Human? 326
The Diversity of Life
16.1 Viruses Are Genes
Wrapped in a
Protein Coat 332 ©Yuriy Dyachyshyn/AFP/Getty Images
A. Viruses Are Smaller and Simpler Than Cells 332
B. A Virus’s Host Range Consists of the Organisms It
Infects 333
C. Are Viruses Alive? 333
16.2 Viral Replication Occurs in Five Stages 334
16.3 Viruses May Kill Bacteria Immediately or Their DNA
May “Hide” in the Cell 335
16.4 Illnesses Caused by Animal Viruses May Be Mild or
Severe 336
A. Symptoms Result from Cell Death and the Immune
Response 336
B. Some Animal Viruses Linger for Years 336
C. Drugs and Vaccines Help Fight Viral Infections 337
xxiv CONTENTS
16.5 Viruses Cause Diseases in Plants 339
16.6 Viroids and Prions Are Other Noncellular Infectious
Agents 340
A. A Viroid Is an Infectious RNA Molecule 340
B. A Prion Is an Infectious Protein 340
16.7 Investigating Life: Scientific Detectives Follow HIV’s
Trail 341
17 | Bacteria and Archaea 344
17.1 Prokaryotes Are a Biological Success Story 346
17.2 Prokaryote Classification Traditionally Relies on Cell
Structure and Metabolism 347
A. Microscopes Reveal Cell Structures 347
B. Metabolic Pathways May Be Useful in
Classification 349
C. Molecular Data Reveal Evolutionary Relationships 351
D. Horizontal Gene Transfer Complicates
Classification 351
17.3 Prokaryotes Include Two Domains with Enormous
Diversity 352
A. Domain Bacteria Includes Many Familiar Groups 352
B. Many, but Not All, Archaea Are “Extremophiles” 353
17.4 Bacteria and Archaea Are Essential to All Life 354
A. Microbes Form Vital Links in Ecosystems 354
B. Bacteria and Archaea Live in and on Us 354
C. Humans Put Many Prokaryotes to Work 356
17.5 Investigating Life: Bacterial Evolution Goes “Hog
Wild” on the Farm 356
18 | Protists 360
18.1 Protists Lie at the Crossroads Between Simple and
Complex Organisms 362
A. What Is a Protist? 362
B. Protists Are Important in Many Ways 362
C. Protists Have a Lengthy Evolutionary History 362
18.2 Algae Are Photosynthetic Protists 364
A. Euglenoids Are Heterotrophs and Autotrophs 364
B. Dinoflagellates Are “Whirling Cells” 364
C. Golden Algae, Diatoms, and Brown Algae Contain
Yellowish Pigments 365
D. Red Algae Can Live in Deep Water 366
E. Green Algae Are the Closest Relatives of Land
Plants 366
18.3 Some Heterotrophic Protists Resemble Fungi 368
A. Slime Molds Are Unicellular and Multicellular 368
B. Water Molds Are Decomposers and Parasites 368
18.4 Protozoa Are Diverse Heterotrophic Protists 370
A. Several Flagellated Protozoa Cause Disease 370
B. Amoeboid Protozoa Produce Pseudopodia 370
C. Ciliates Are Common Protozoa with Complex
Cells 371
D. Apicomplexans Include Nonmotile Animal Parasites 372
18.5 Protist Classification Is Changing Rapidly 374
18.6 Investigating Life: Shining a Spotlight on Danger 375
19 | Plants 378
19.1 Plants Have Changed the World 380
A. Green Algae Are the Closest Relatives of Plants 380
B. Plants Are Adapted to Life on Land 382
19.2 Bryophytes Are the Simplest Plants 384
A. Bryophytes Lack Vascular Tissue 384
B. Bryophytes Have a Conspicuous Gametophyte 385
19.3 Seedless Vascular Plants Have Xylem and Phloem but
No Seeds 386
A. Seedless Vascular Plants Include Ferns and Their Close
Relatives 386
B. Seedless Vascular Plants Have a Conspicuous
Sporophyte and Swimming Sperm 387
19.4 Gymnosperms Are “Naked Seed” Plants 388
A. Gymnosperms Include Conifers and Three Related
Groups 388
B. Conifers Produce Pollen and Seeds in Cones 389
19.5 Angiosperms Produce Seeds in Fruits 390
A. Most Angiosperms Are Eudicots or Monocots 390
B. Flowers and Fruits Are Unique to the Angiosperm Life
Cycle 390
C. Wind and Animals Often Participate in Angiosperm
Reproduction 392
19.6 Investigating Life: Genetic Messages from Ancient
Ecosystems 394
20 | Fungi 398
20.1 Fungi Are Essential Decomposers 400
A. Fungi Are Eukaryotic Heterotrophs That Digest Food
Externally 400
B. Fungal Classification Is Traditionally Based on
Reproductive Structures 402
20.2 Chytridiomycetes Produce Swimming Spores 403
20.3 Zygomycetes Are Fast-Growing and Prolific 404
20.4 Glomeromycetes Colonize Living Plant Roots 405
20.5 Ascomycetes Are the Sac Fungi 406
20.6 Basidiomycetes Are the Familiar Club Fungi 408
20.7 Fungi Interact with Other Organisms 410
A. Endophytes Colonize Plant Tissues 410
B. Mycorrhizal Fungi Exchange Materials with Roots 410
C. Some Ants Cultivate Fungi 410
D. Lichens Are Dual Organisms 411

20.8 Investigating Life: The Battle for Position in Cacao
Tree Leaves 412
21 | Animals 416
21.1 Animals Live Nearly Everywhere 418
A. What Is an Animal? 418
B. Animal Life Began in the Water 418
C. Animal Features Reflect Shared Ancestry 419
D. Biologists Also Consider Additional
Characteristics 421
21.2 Sponges Are Simple Animals That Lack Differentiated
Tissues 423
21.3 Cnidarians Are Radially Symmetrical, Aquatic
Animals 424
21.4 Flatworms Have Bilateral Symmetry and Incomplete
Digestive Tracts 425
21.5 Mollusks Are Soft, Unsegmented Animals 427
21.6 Annelids Are Segmented Worms 428
21.7 Nematodes Are Unsegmented, Cylindrical Worms 430
21.8 Arthropods Have Exoskeletons and Jointed
Appendages 432
A. Arthropods Have Complex Organ Systems 432
B. Arthropods Are the Most Diverse Animals 434
21.9 Echinoderm Adults Have Five-Part, Radial
Symmetry 436
21.10 Most Chordates Are Vertebrates 438
A. Four Key Features Distinguish Chordates 438
B. Many Features Reveal Evolutionary Relationships
Among Chordates 439
21.11 Tunicates and Lancelets Are Invertebrate
Chordates 442
21.12 Hagfishes and Lampreys Are Craniates Lacking
Jaws 443
21.13 Fishes Are Aquatic Vertebrates with Jaws, Gills,
and Fins 444
A. Cartilaginous Fishes Include Sharks, Skates,
and Rays 444
B. Bony Fishes Include Two Main Lineages 444
C. Fishes Changed the Course of Vertebrate
Evolution 445
21.14 Amphibians Lead a Double Life on Land and in
Water 446
A. Amphibians Were the First Tetrapods 446
B. Amphibians Include Three Main Lineages 446
21.15 Reptiles Were the First Vertebrates to Thrive on Dry
Land 448
A. Nonavian Reptiles Include Four Main Groups 448
B. Birds Are Warm, Feathered Reptiles 450
CONTENTS xxv
21.16 Mammals Are Warm, Furry Milk-Drinkers 451
A. Mammals Share a Common Ancestor with Reptiles 451
B. Mammals Lay Eggs or Bear Live Young 452
21.17 Investigating Life: Sponges Fill Holes in Animal
Evolution 453
UNIT 5 Plant Life
|
22 Plant Form and
Function 460
22.1 Vegetative Plant Parts
©Gustavo Gilabert/Corbis SABA
Include Stems, Leaves,
and Roots 462
22.2 Plant Cells Build Tissues 465
A. Plants Have Several Cell Types 465
B. Plant Cells Form Three Main Tissue Systems 467
22.3 Tissues Build Stems, Leaves, and Roots 469
A. Stems Support Leaves 469
B. Leaves Are the Primary Organs of Photosynthesis 469
C. Roots Absorb Water and Minerals and Anchor the
Plant 470
22.4 Plants Have Flexible Growth Patterns, Thanks to
Meristems 473
A. Plants Grow by Adding New Modules 473
B. Plant Growth Occurs at Meristems 473
C. In Primary Growth, Apical Meristems Lengthen Stems
and Roots 474
D. In Secondary Growth, Lateral Meristems Thicken
Stems and Roots 474
22.5 Investigating Life: An Army of Tiny Watchdogs 477
23 | Plant Nutrition and Transport 482
23.1 Soil and Air Provide Water and Nutrients 484
A. Plants Require 16 Essential Elements 484
B. Soils Have Distinct Layers 484
C. Leaves and Roots Absorb Essential Elements 485
23.2 Water and Minerals Are Pulled Up to Leaves in
Xylem 487
A. Water Evaporates from Leaves in Transpiration 487
B. Water and Dissolved Minerals Enter at the Roots 488
C. Xylem Transport Relies on Cohesion 488
D. The Cuticle and Stomata Help Conserve Water 489
23.3 Sugars Are Pushed in Phloem to Nonphotosynthetic
Cells 490
A. Phloem Sap Contains Sugars and Other Organic
Compounds 490
B. The Pressure Flow Theory Explains Phloem
Function 490
xxvi CONTENTS
23.4 Parasitic Plants Tap into Another Plant’s Vascular
Tissue 492
23.5 Investigating Life: The Hidden Cost of Traps 492
24
|
Reproduction and Development
of Flowering Plants 496
24.1 Angiosperms Reproduce Asexually and Sexually 498
A. Asexual Reproduction Yields Clones 498
B. Sexual Reproduction Generates Variability 499
24.2 The Angiosperm Life Cycle Includes Flowers, Fruits,
and Seeds 500
A. Flowers Are Reproductive Organs 500
B. The Pollen Grain and Embryo Sac Are
Gametophytes 501
C. Pollination Brings Pollen to the Stigma 501
D. Double Fertilization Yields Zygote and Endosperm 502
E. A Seed Is an Embryo and Its Food Supply Inside a Seed
Coat 503
F. The Fruit Develops from the Ovary 504
G. Fruits Protect and Disperse Seeds 505
24.3 Plant Growth Begins with Seed Germination 506
24.4 Hormones Regulate Plant Growth and Development 507
A. Auxins and Cytokinins Are Essential for Plant
Growth 507
B. Gibberellins, Ethylene, and Abscisic Acid Influence
Plant Development in Many Ways 508
C. Biologists Continue to Discover Additional Plant
Hormones 509
24.5 Light Is a Powerful Influence on Plant Life 510
A. Phototropism Is Growth Toward Light 510
B. Phytochrome Regulates Seed Germination, Daily
Rhythms, and Flowering 511
24.6 Plants Respond to Gravity and Touch 513
24.7 Plant Parts Die or Become Dormant 514
24.8 Investigating Life: A Red Hot Chili Pepper Paradox 515
UNIT 6 Animal Life
|
25 Animal Tissues
and Organ
Systems 518
©Daniel Garcia/AFP/Getty Images
25.1 Specialized Cells Build
Animal Bodies 520
25.2 Animals Consist of Four Tissue Types 522
A. Epithelial Tissue Covers Surfaces 522
B. Most Connective Tissues Bind Other Tissues
Together 523
C. Muscle Tissue Provides Movement 524
D. Nervous Tissue Forms a Rapid Communication
Network 525
25.3 Organ Systems Are Interconnected 526
A. The Nervous and Endocrine Systems Coordinate
Communication 526
B. The Skeletal and Muscular Systems Support and Move
the Body 526
C. The Digestive, Circulatory, and Respiratory Systems
Help Acquire Energy 526
D. The Urinary, Integumentary, Immune, and Lymphatic
Systems Protect the Body 527
E. The Reproductive System Produces the Next
Generation 527
25.4 Organ System Interactions Promote Homeostasis 528
25.5 The Integumentary System Regulates Temperature and
Conserves Moisture 529
25.6 Investigating Life: Vitamins and the Evolution of
Human Skin Pigmentation 531
26 | The Nervous System 534
26.1 The Nervous System Forms a Rapid Communication
Network 536
A. Invertebrates Have Nerve Nets, Nerve Ladders, or
Nerve Cords 536
B. Vertebrate Nervous Systems Are Highly Centralized 537
26.2 Neurons Are Functional Units of a Nervous System 538
A. A Typical Neuron Consists of a Cell Body, Dendrites,
and an Axon 538
B. The Nervous System Includes Three Classes of
Neurons 539
26.3 Action Potentials Convey Messages 540
A. A Neuron at Rest Has a Negative Charge 540
B. A Neuron’s Membrane Potential Reverses During an
Action Potential 540
C. The Myelin Sheath Speeds Impulse Conduction 543
26.4 Neurotransmitters Pass the Message from Cell to Cell 544
A. Neurons Communicate at Synapses 544
B. A Neuron Integrates Signals from Multiple Synapses 545
26.5 The Peripheral Nervous System Consists of Nerve Cells
Outside the Central Nervous System 546
26.6 The Central Nervous System Consists of the Spinal
Cord and Brain 548
A. The Spinal Cord Transmits Information Between Body
and Brain 548
B. The Human Brain Is Divided into Several Regions 548
C. Many Brain Regions Participate in Memory
Formation 550
D. Damage to the Central Nervous System Can Be
Devastating 551
26.7 Investigating Life: Scorpion Stings Don’t Faze
Grasshopper Mice 554

27 | The Senses 558
27.1 Diverse Senses Operate by the Same Principles 560
29
A. Sensory Receptors Respond to Stimuli by Generating
Action Potentials 560
B. Continuous Stimulation May Cause Sensory
Adaptation 561
27.2 The General Senses Detect Touch, Temperature, Pain,
and Position 562
27.3 The Senses of Smell and Taste Detect Chemicals 563
A. Chemoreceptors in the Nose Detect Odor Molecules 563
B. Chemoreceptors in the Mouth Detect Taste 564
27.4 Vision Depends on Light-Sensitive Cells 565
A. Invertebrate Eyes Take Many Forms 565
B. In the Vertebrate Eye, Light Is Focused on the Retina 565
C. Signals Travel from the Retina to the Optic Nerve and
Brain 566
27.5 The Senses of Hearing and Equilibrium Begin in the
Ears 568
A. Mechanoreceptors in the Inner Ear Detect Sound
Waves 568
B. The Inner Ear Also Provides the Sense of
Equilibrium 569
27.6 Investigating Life: How Do Whales Taste? 570
28 | The Endocrine System 574
28.1 The Endocrine System Uses Hormones to
Communicate 576
A. Endocrine Glands Secrete Hormones That Interact with
Target Cells 576
B. The Nervous and Endocrine Systems Work
Together 577
30 | The Circulatory System 608
28.2 Hormones Stimulate Responses in Target Cells 578
A. Water-Soluble Hormones Trigger Second Messenger
Systems 578
B. Lipid-Soluble Hormones Directly Alter Gene
Expression 579
28.3 The Hypothalamus and Pituitary Gland Oversee
Endocrine Control 581
A. The Posterior Pituitary Stores and Releases Two
Hormones 581
B. The Anterior Pituitary Produces and Secretes Six
Hormones 581
28.4 Hormones from Many Glands Regulate
Metabolism 582
A. The Thyroid Gland Sets the Metabolic Pace 582
B. The Parathyroid Glands Control Calcium Level 583
C. The Adrenal Glands Coordinate the Body’s Stress
Responses 583
D. The Pancreas Regulates Blood Glucose 584
E. The Pineal Gland Secretes Melatonin 585
CONTENTS xxvii
28.5 Hormones from the Ovaries and Testes Control
Reproduction 586
28.6 Investigating Life: Addicted to Affection 586
|
The Skeletal and Muscular
Systems 590
29.1 Skeletons Take Many Forms 592
29.2 The Vertebrate Skeleton Features a Central Backbone 593
29.3 Bones Provide Support, Protect Internal Organs, and
Supply Calcium 594
A. Bones Consist Mostly of Bone Tissue and
Cartilage 594
B. Bones Are Constantly Built and Degraded 596
C. Bones Help Regulate Calcium Homeostasis 596
D. Bone Meets Bone at a Joint 597
29.4 Muscle Movement Requires Contractile Proteins,
Calcium, and ATP 598
A. Actin and Myosin Filaments Fill Muscle Cells 598
B. Sliding Filaments Are the Basis of Muscle Fiber
Contraction 599
C. Motor Neurons Stimulate Muscle Fiber Contraction 600
29.5 Muscle Fibers Generate ATP in Many Ways 602
29.6 Many Muscle Fibers Combine to Form One Muscle 603
A. Each Muscle May Contract with Variable Force 603
B. Muscles Contain Slow- and Fast-Twitch Fibers 603
C. Exercise Strengthens Muscles 604
29.7 Investigating Life: Did a Myosin Gene Mutation Make
Humans Brainier? 604
30.1 Circulatory Systems Deliver Nutrients and Remove
Wastes 610
A. Circulatory Systems Are Open or Closed 610
B. Vertebrate Circulatory Systems Have Become
Increasingly Complex 611
30.2 Blood Is a Complex Mixture 612
A. Plasma Carries Many Dissolved Substances 612
B. Red Blood Cells Transport Oxygen 613
C. White Blood Cells Fight Infection 613
D. Blood Clotting Requires Platelets and Plasma
Proteins 614
30.3 Blood Circulates Through the Heart and Blood
Vessels 615
30.4 The Human Heart Is a Muscular Pump 616
A. The Heart Has Four Chambers 616
B. The Right and Left Halves of the Heart Deliver Blood
Along Different Paths 616
C. Cardiac Muscle Cells Produce the Heartbeat 617
D. Exercise Strengthens the Heart 618
xxviii CONTENTS
30.5 Blood Vessels Form the Circulation Pathway 619
A. Arteries, Capillaries, and Veins Have Different
Structures 619
B. Blood Pressure and Velocity Differ Among Vessel
Types 620
30.6 The Lymphatic System Maintains Circulation and
Protects Against Infection 623
30.7 Investigating Life: In (Extremely) Cold Blood 624
31 | The Respiratory System 628
31.1 Gases Diffuse Across Respiratory Surfaces 630
A. Some Invertebrates Exchange Gases Across the Body
Wall or in Internal Tubules 631
B. Gills Exchange Gases with Water 632
C. Terrestrial Vertebrates Exchange Gases in Lungs 632
31.2 The Human Respiratory System Delivers Air to the
Lungs 634
A. The Nose, Pharynx, and Larynx Form the Upper
Respiratory Tract 634
B. The Lower Respiratory Tract Consists of the Trachea
and Lungs 635
31.3 Breathing Requires Pressure Changes in the
Lungs 636
31.4 Blood Delivers Oxygen and Removes Carbon
Dioxide 638
A. Blood Carries Gases in Several Forms 638
B. Blood Gas Levels Help Regulate the Breathing
Rate 638
31.5 Investigating Life: Why Do Bugs Hold Their
Breath? 640
32 | Digestion and Nutrition 644
32.1 Digestive Systems Derive Nutrients from Food 646
A. Animals Eat to Obtain Energy and Building
Blocks 646
B. How Much Food Does an Animal Need? 646
C. Animals Process Food in Four Stages 646
D. Animal Diets and Feeding Strategies Vary
Greatly 647
32.2 Animal Digestive Tracts Take Many Forms 648
32.3 The Human Digestive System Consists of Several
Organs 650
A. Digestion Begins in the Mouth 650
B. The Stomach Stores, Digests, and Churns Food 651
C. The Small Intestine Digests and Absorbs
Nutrients 652
D. The Large Intestine Completes Nutrient and Water
Absorption 654
32.4 A Healthy Diet Includes Essential Nutrients and the
Right Number of Calories 656
A. A Varied Diet Is Essential to Good Health 656
B. Body Weight Reflects Food Intake and Activity
Level 658
C. Starvation: Too Few Calories to Meet the Body’s
Needs 659
D. Obesity: More Calories Than the Body Needs 660
32.5 Investigating Life: The Cost of a Sweet Tooth 661
|
33 Regulation of Temperature and Body
Fluids 664
33.1 Animals Regulate Their Internal Temperature 666
A. Heat Gains and Losses Determine an Animal’s Body
Temperature 666
B. Several Adaptations Help an Animal to Adjust Its
Temperature 667
33.2 Animals Regulate Water and Ions in Body Fluids 669
33.3 Nitrogenous Wastes Include Ammonia, Urea, and Uric
Acid 670
33.4 The Urinary System Produces, Stores, and Eliminates
Urine 671
33.5 The Nephron Is the Functional Unit of the Kidney 672
A. Nephrons Interact Closely with Blood Vessels 672
B. Urine Formation Includes Filtration, Reabsorption,
and Secretion 672
C. The Glomerular Capsule Filters Blood 674
D. Reabsorption and Secretion Occur in the Renal
Tubule 674
E. The Collecting Duct Conserves More Water 675
F. Hormones Regulate Kidney Function 675
33.6 Investigating Life: Sniffing Out the Origin of Fur and
Feathers 676
34 | The Immune System 680
34.1 Many Cells, Tissues, and Organs Defend the Body 682
A. White Blood Cells Play Major Roles in the Immune
System 682
B. The Lymphatic System Produces and Transports Many
Immune System Cells 683
C. The Immune System Has Two Main Subdivisions 683
34.2 Innate Defenses Are Nonspecific and Act Early 684
A. External Barriers Form the First Line of Defense 684
B. Internal Innate Defenses Destroy Invaders 684
34.3 Adaptive Immunity Defends Against Specific
Pathogens 686
A. Macrophages Trigger Both Cell-Mediated and Humoral
Immunity 686
B. Cytotoxic T Cells Provide Cell-Mediated Immunity 687
C. B Cells Direct the Humoral Immune Response 687
D. The Immune Response Turns Off Once the Threat Is
Gone 690
E. The Secondary Immune Response Is Stronger Than the
Primary Response 690

34.4 Vaccines Jump-Start Immunity 692
34.5 Several Disorders Affect the Immune System 693
A. Autoimmune Disorders Are Devastating and
Mysterious 693
B. Immunodeficiencies Lead to Opportunistic Infections 693
C. Allergies Misdirect the Immune Response 694
D. A Pregnant Woman’s Immune System May Attack Her
Fetus 695
34.6 Investigating Life: The Hidden Cost of Hygiene 696
35
|
Animal Reproduction and
Development 700
35.1 Animal Development Begins with Reproduction 702
A. Reproduction Is Asexual or Sexual 702
B. Gene Expression Dictates Animal Development 702
C. Development Is Indirect or Direct 703
35.2 Males Produce Sperm Cells 704
A. Male Reproductive Organs Are Inside and Outside the
Body 704
B. Spermatogenesis Yields Sperm Cells 705
C. Hormones Influence Male Reproductive Function 706
35.3 Females Produce Egg Cells 707
A. Female Reproductive Organs Are Inside the Body 707
B. Oogenesis Yields Egg Cells 708
C. Hormones Influence Female Reproductive Function 709
D. Hormonal Fluctuations Can Cause Discomfort 711
35.4 Reproductive Health Considers Contraception and
37 | Populations 748
Disease 711
35.5 The Human Infant Begins Life as a Zygote 714
A. Fertilization Initiates Pregnancy 714
B. The Preembryonic Stage Ends When Implantation Is
Complete 715
C. Organs Take Shape During the Embryonic Stage 716
D. Organ Systems Become Functional in the Fetal
Stage 719
E. Muscle Contractions in the Uterus Drive Childbirth 720
35.6 Birth Defects Have Many Causes 721
35.7 Investigating Life: The Ultimate Sacrifice 723
UNIT 7 The Ecology of Life
36
|
Animal
Behavior 728
36.1 Animal Behaviors
©Gerald Hinde/Gallo Images/
Have Proximate and Getty Images RF
Ultimate Causes 730
36.2 Animal Behaviors Combine Innate and Learned
Components 731
A. Innate Behaviors Do Not Require Experience 731
CONTENTS xxix
B. Learning Requires Experience 732
C. Genes and Environment Interact to Determine
Behavior 733
36.3 Many Behaviors Improve Survival 734
A. Some Animals Can Find Specific Locations 734
B. Animals Balance the Energy Content and Costs of
Acquiring Food 735
C. Avoiding Predation Is Another Key to Survival 736
36.4 Many Behaviors Promote Reproductive Success 738
A. Courtship Sets the Stage for Mating 738
B. Sexual Selection Leads to Differences Between the
Sexes 738
C. Animals Differ in Mating Systems and Degrees of
Parental Care 739
D. Human Reproductive Choices May Reflect Natural
Selection 740
36.5 Social Behaviors Often Occur in Groups 741
A. Group Living Has Costs and Benefits 741
B. Dominance Hierarchies and Territoriality Reduce
Competition 742
C. Kin Selection and Reciprocal Altruism Explain Some
Acts of Cooperation 742
D. Eusocial Animals Have Highly Developed
Societies 743
36.6 Investigating Life: The “Cross-Dressers” of the
Reef 744
37.1 A Population Consists of Individuals of One Species 750
A. Density and Distribution Patterns Are Static Measures
of a Population 750
B. Isolated Subpopulations May Evolve into New
Species 750
37.2 Births and Deaths Help Determine Population
Size 752
A. Births Add Individuals to a Population 752
B. Survivorship Curves Show the Probability of Dying at a
Given Age 753
37.3 Population Growth May Be Exponential or Logistic 754
A. Growth Is Exponential When Resources Are
Unlimited 754
B. Population Growth Eventually Slows 755
C. Many Conditions Limit Population Size 757
37.4 Natural Selection Influences Life Histories 758
A. Organisms Balance Reproduction Against Other
Requirements 758
B. Opportunistic and Equilibrium Life Histories Reflect
the Trade-Off Between Quantity and Quality 758
37.5 The Human Population Continues to Grow 759
A. Birth Rates and Death Rates Vary Worldwide 759
B. The Ecological Footprint Is an Estimate of
Resource Use 761
37.6 Investigating Life: A Toxic Compromise 763
xxx CONTENTS
38 | Communities and Ecosystems 766
38.1 Multiple Species Interact in Communities 768
A. Many Species Compete for the Same Resources 768
B. Symbiotic Interactions Can Benefit or Harm a
Species 769
C. Herbivory and Predation Link Species in Feeding
Relationships 770
D. Closely Interacting Species May Coevolve 771
38.2 Succession Is a Gradual Change in a Community 772
38.3 Ecosystems Require Continuous Energy Input 774
A. Food Webs Depict the Transfer of Energy and
Atoms 774
B. A Keystone Species Has a Pivotal Role in the
Community 776
C. Heat Energy Leaves Each Food Web 776
D. Harmful Chemicals May Accumulate in the Highest
Trophic Levels 777
38.4 Chemicals Cycle Within Ecosystems 778
A. Water Circulates Between the Land and the
Atmosphere 778
B. Autotrophs Obtain Carbon as CO2 780
C. The Nitrogen Cycle Relies on Bacteria 781
D. The Phosphorus Cycle Begins with the Erosion of
Rocks 782
E. Excess Nitrogen and Phosphorus Cause Problems in
Water 782
F. Terrestrial and Aquatic Ecosystems Are Linked in
Surprising Ways 783
38.5 Investigating Life: Two Kingdoms and a Virus Team Up
to Beat the Heat 784
39 | Biomes 788
39.1 The Physical Environment Determines Where Life
Exists 790
39.2 Earth Has Diverse Climates 792
39.3 Terrestrial Biomes Range from the Lush Tropics to the
Frozen Poles 794
A. Towering Trees Dominate the Forests 795
B. Grasslands Occur in Tropical and Temperate
Regions 796
C. Whether Hot or Cold, All Deserts Are Dry 797
D. Fire- and Drought-Adapted Plants Dominate
Mediterranean Shrublands (Chaparral) 798
E. Tundras Occupy High Latitudes and High
Elevations 799
F. Polar Ice Caps Are Cold and Dry 799
39.4 Freshwater Biomes Include Lakes, Ponds, and
Streams 800
A. Lakes and Ponds Contain Standing Water 800
B. Streams Carry Running Water 801
39.5 Oceans Make Up Earth’s Largest Ecosystem 802
A. Land Meets Sea at the Coast 802
B. The Open Ocean Remains Mysterious 803
39.6 Investigating Life: There’s No Place Like Home 804
40 | Preserving Biodiversity 808
40.1 Earth’s Biodiversity Is Dwindling 810
40.2 Many Human Activities Destroy Habitats 811
40.3 Pollution Degrades Habitats 813
A. Water Pollution Threatens Aquatic Life 813
B. Air Pollution Causes Many Types of Damage 814
40.4 Global Climate Change Alters and Shifts Habitats 816
A. Greenhouse Gases Warm Earth’s Surface 816
B. Global Climate Change Has Severe Consequences 817
40.5 Exotic Invaders and Overexploitation Devastate Many
Species 818
A. Invasive Species Displace Native Organisms 818
B. Overexploitation Can Drive Species to Extinction 819
40.6 Some Biodiversity May Be Recoverable 820
A. Protecting and Restoring Habitat Saves Many Species
at Once 820
B. Some Conservation Tools Target Individual
Species 820
C. Conserving Biodiversity Involves Scientists and
Ordinary Citizens 821
40.7 Investigating Life: Up, Up, and Away 822
APPENDIX A Answers to Multiple Choice Questions A-1
APPENDIX B A Brief Guide to Statistical Significance A-2
APPENDIX C Units of Measurement A-5
APPENDIX D Periodic Table of the Elements A-6
APPENDIX E Amino Acid Structures A-7
APPENDIX F Learn How to Learn A-8
Glossary G-1 | Index I-1
BIOLOGY
 1
CHAPTER

The Scientific Study of Life

Undersea World. A coral reef in the Red Sea is home to countless marine species. The prickly animal in the center is a “crown of thorns” sea star.
©Franco Banfi/WaterF/age fotostock

LEARN HOW TO LEARN

Real Learning Takes Time
You got good at basketball, running, dancing, art, music, or video games by putting in lots of practice. Likewise, you will need to commit time to your
biology course if you hope to do well. To get started, look for the “Learn How to Learn” tip in each chapter of this textbook. Each hint is designed to help
you use your study time productively.
With practice, you’ll discover that all concepts in biology are connected. The Survey the Landscape figure in every chapter highlights each chapter’s place
in the “landscape” of the entire unit. Use it, along with the more detailed Pull It Together concept map in the chapter summary, to see how each chapter’s
content fits into the unit’s big picture.
UNIT 1
Life Is Everywhere LEA R NING OU T LI N E
Welcome to biology, the scientific study of life. Living organ- 1.1 What Is Life?
isms surround us. You are alive, and so are your friends, your
1.2 The Tree of Life Includes Three Main Branches
pets, and the plants in your home and yard. Bacteria thrive on and
in your body. Any food you ate today was (until recently, anyway) 1.3 Scientists Study the Natural World
alive. And the news is full of biology-related discoveries about
1.4 Investigating Life: The Orchid and the Moth
fossils, new cancer treatments, genetics, global climate change,
and the environment.
Stories such as these enjoy frequent media coverage be-
cause this is an exciting time to study biology. Not only is the
field changing rapidly, but its new discoveries and applications
might change your life. DNA technology has brought us geneti-
cally engineered bacteria that can manufacture life-saving
drugs—and genetically engineered plants that produce their
own pesticides. This same technology may one day enable phy-
sicians to routinely cure hemophilia, cystic fibrosis, and other
genetic diseases by replacing faulty DNA with a functional
“patch.”
Biology also includes the study of nonhuman life. We exist
only because of our interactions with other species, which
­provide food, oxygen, clean water, clothing, shelter, and other
necessities. Even species that do not directly “serve” us are es-
sential to the ecosystems that sustain all life. Human activities,
however, are pushing many ecosystems dangerously out of
­balance.
Consider the “crown of thorns” sea star shown here. These
animals are notorious for their arsenal of sharp, venomous spines,
which may cause painful wounds. At low population densities,
S U RV E Y T H E L ANDSCAPE
their coral-eating habits help maintain reef biodiversity. Some-
Science, Chemistry, and Cells
times, however, huge numbers of sea stars destroy entire patches
of coral. What causes these infestations? Many researchers point Life is the scientific Biology
to nutrient-polluted runoff from nearby farms and cities. The nu- study of
consists carry
trient influx triggers a population explosion of algae, which sea of units out the
called functions of
star larvae eat as they develop into adults. Removing the adults
ATP makes Respiration
from an infested reef is dangerous and labor-intensive, but help is
Cells uses
coming from an unusual source: Underwater robots have been
programmed to seek out the crown of thorns and deliver a consist of Carbohydrates

­lethal injection. makes

The list of biology-related topics goes on and on: global cli- Molecules include Proteins Photosynthesis
mate change, stem cell therapies, new cancer treatments, infec- consist of encodes such as
tious disease, improved crop plants, synthetic life, i­nfertility
treatment, endangered species, DNA fingerprinting, biofuels, Atoms DNA Enzymes
pollution, the history of life, and more. This book will bring you
a taste of what we know about life and help you make sense of the
Organisms from all three branches of life share a unique combi-
science-related news you see every day. Chapter 1 begins your nation of characteristics. Biologists are scientists who use
journey by introducing the scope of biology and explaining how evidence to test hypotheses about life.
science teaches us what we know about life. For more details, study the Pull It Together feature in the chapter summary.

3
4 UNIT ONE Science, Chemistry, and Cells

1.1 What Is Life?

Biology is the scientific study of life. The second half of this
chapter explores the meaning of the term scientific, but first we
will consider the question “What is life?” We all have an intuitive
sense of what life is. If we see a rabbit on a rock, we know that
the rabbit is alive and the rock is not. But it is difficult to state just
what makes the rabbit alive. Likewise, in the instant after an in-
dividual dies, we may wonder what invisible essence has trans-
formed the living into the dead.
One way to define life is to list its basic components. The cell
is the basic unit of life; every organism, or living individual, con-
sists of one or more cells. Every cell has an outer membrane that
separates it from its surroundings. This membrane encloses the
­water and other chemicals that carry out the cell's functions. One of
those biochemicals, deoxyribonucleic acid (DNA), is the informa-
tional molecule of life (figure 1.1). Cells use genetic instructions—
as encoded in DNA—to produce proteins, which enable cells to
carry out their functions in tissues, organs, and organ systems.
A list of life’s biochemicals, however, provides an unsatisfy-
ing definition of life. After all, placing DNA, water, proteins, and Figure 1.1 Informational Molecule of Life. All cells contain DNA, a
a membrane in a test tube does not create life. And a crushed in- series of “recipes” for proteins that each cell can make.
sect still contains all of the biochemicals that it had immediately ©Scott Camazine/123RF
before it died.
In the absence of a concise definition, scientists have settled form molecules. These molecules can form organelles, which
on five qualities that, in combination, constitute life. Table 1.1 are compartments that carry out specialized functions in cells
summarizes them, and the rest of section 1.1 describes each one (note that not all cells contain organelles). Many organisms con-
in more detail. An organism is a collection of structures that sist of single cells. In multicellular organisms such as the tree
function together and exhibit all of these qualities. Note, h­ owever, ­illustrated in figure 1.2, however, the cells are organized into spe-
that each trait in table 1.1 may also occur in nonliving objects. A cialized tissues that make up organs. Multiple organs are linked
rock crystal is highly organized, but it is not alive. A fork placed into an individual’s organ systems.
in a pot of boiling water absorbs heat energy and passes it to the We have now reached the level of the organism, which may
hand that grabs it, but this does not make the fork alive. A fire can consist of just one cell or of many cells organized into tissues,
“reproduce” and grow very rapidly, but it lacks most of the other organs, and organ systems. Organization in the living world
characteristics of life. It is the combination of these five charac- ­extends beyond the level of the individual organism as well. A
teristics that makes life unique. ­population includes members of the same species occupying the
same place at the same time. A community includes the popula-
tions of different species in a region, and an ecosystem includes
A. Life Is Organized both the living and nonliving components of an area. Finally, the
Just as the city where you live belongs to a county, state, and na- biosphere consists of all parts of the planet that can support life.
tion, living matter also consists of parts organized in a hierarchi- Biological organization is apparent in all life. Humans, eels,
cal pattern (figure 1.2). At the smallest scale, all living structures and evergreens, although outwardly very different, are all orga-
are composed of particles called atoms, which bond together to nized into specialized cells, tissues, organs, and organ systems.

TA B L E 1.1 Characteristics of Life: A Summary

Characteristic Example
Organization Atoms make up molecules, which make up cells, which make up tissues, and so on.

Energy use A kitten uses the energy from its mother’s milk to fuel its own growth.

Maintenance of internal constancy (homeostasis) Your kidneys regulate your body’s water balance by adjusting the concentration of your urine.

Reproduction, growth, and development An acorn germinates, develops into an oak seedling, and, at maturity, reproduces sexually to produce its
own acorns.

Evolution Increasing numbers of bacteria survive treatment with antibiotic drugs.

 CHAPTER 1 The Scientific Study of Life 5

CELL
ORGANELLE The fundamental
A membrane-bounded unit of life. Multicellular
structure that has a specific organisms consist of many
function within a cell. cells; unicellular organisms TISSUE
Example: Chloroplast consist of one cell. A collection of specialized cells
Example: Leaf cell that function in a coordinated
fashion. (Multicellular life only.)
MOLECULE Example: Epidermis of leaf
A group of joined atoms.
Example: DNA
ORGAN
A structure consisting
of tissues organized to
interact and carry
ATOM out specific functions.
The smallest chemical (Multicellular life only.)
unit of a type of pure Example: Leaf
substance (element).
Example: Carbon atom ORGANISM
A single living individual.
Example: One acacia tree

ORGAN SYSTEM
Organs connected
physically or chemically
POPULATION
that function together.
A group of the same species of organism
(Multicellular life only.)
living in the same place and time.
Example: Aboveground
Example: Multiple acacia trees
part of a plant

BIOSPHERE
COMMUNITY The global ecosystem;
All populations that occupy the parts of the planet
the same region. ECOSYSTEM and its atmosphere
Example: All populations The living and nonliving where life is possible.
in a savanna components of an area.
Example: The savanna

Figure 1.2 Life’s Organizational Hierarchy. This diagram applies life’s organizational hierarchy to a multicellular organism (an acacia tree). Green arrows
represent the organizational hierarchy up to the level of the organism; blue arrows represent levels that include multiple organisms.
Photos: (population): ©Gregory G. Dimijian, M.D./Science Source; (community): ©Daryl Balfour/Gallo Images/Getty Images; (ecosystem): ©Bas Vermolen/Getty Images;
(biosphere): ©Corbis RF
6 UNIT ONE Science, Chemistry, and Cells
Brain cell Interacting brain cells
Brain
Emergent property: Memory
Figure 1.3 An Emergent Property—from Cells to Memories. Highly
branched cells interact to form a complex network in the brain. Memories,
consciousness, and other qualities of the mind emerge only when these cells
interact in a certain way.
Single-celled bacteria, although less complex than animals or
plants, still contain DNA, proteins, and other molecules that in-
teract in highly organized ways.
An organism, however, is more than a collection of succes-
sively smaller parts. Emergent properties are new functions that
arise from physical and chemical interactions among a system’s
components, much as flour, sugar, butter, and chocolate can be-
come brownies—something not evident from the parts them-
selves. Figure 1.3 shows another example of emergent properties:
the thoughts and memories produced by interactions among the
neurons in a person’s brain. For an emergent property, the whole
is greater than the sum of the parts.
Emergent properties explain why structural organization is
closely tied to function. Disrupt a structure, and its function ceases.
Brain damage, for instance, disturbs the interactions between brain
cells and can interfere with memory, coordination, and other brain
functions. Likewise, if a function is interrupted, the corresponding
structure eventually breaks down, much as unused muscles begin
to waste away. Biological function and form are interdependent.
B. Life Requires Energy
Inside each cell, countless chemical reactions sustain life. These
reactions, collectively called metabolism, allow organisms to ac-
quire and use energy and nutrients to build new structures, repair
old ones, and reproduce.
Biologists divide organisms into broad categories, based on
their source of energy and raw materials (figure 1.4). Producers,
also called autotrophs, make their own food by extracting energy
and nutrients from nonliving sources. The most familiar produc-
ers are the plants and microbes that capture light energy from the
sun, but some bacteria can derive chemical energy from rocks.
Consumers, in contrast, obtain energy and nutrients by eating
other organisms, living or dead; consumers are also called het-
erotrophs. You are a consumer, relying on energy and atoms from
food to stay alive. Decomposers are heterotrophs that absorb
­energy and nutrients from wastes or dead organisms. These or-
ganisms, which include fungi and some bacteria, recycle nutri-
ents to the nonliving environment.
Within an ecosystem, organisms are linked into elaborate
food webs, beginning with producers and continuing through
several levels of consumers (including decomposers). But energy
transfers are never 100% efficient; some energy is always lost to
the surroundings in the form of heat (see figure 1.4). Because no
organism can use it as an energy source, heat represents a perma-
nent loss from the cycle of life. All ecosystems therefore depend
on a continuous stream of energy from an outside source, usually
the sun. i food webs, section 38.3A
C. Life Maintains Internal Constancy
The conditions inside cells must remain within a constant range,
even if the surrounding environment changes. For example, a cell
must maintain a certain temperature; it will die if it becomes too
Energy Usable energy
from Nutrients
sunlight
Heat
Heat
Consumers obtain
energy and nutrients by
eating other organisms.
Producers extract energy
and nutrients from the
nonliving environment.
Soil
Heat
Decomposers are consumers
that obtain nutrients from dead
organisms and organic wastes.
Figure 1.4 Life Is Connected. All organisms extract energy and nutrients
from the nonliving environment or from other organisms. Decomposers
recycle nutrients back to the nonliving environment. At every stage along the
way, heat is lost to the surroundings.
 CHAPTER 1 The Scientific Study of Life 7

a. b.

a. b. Figure 1.6 Asexual and Sexual Reproduction. (a) Identical plantlets

develop along the runners of a wild strawberry plant. (b) Two swans protect
Figure 1.5 Temperature Homeostasis. (a) Shivering and (b) sweating their offspring, the products of sexual reproduction.
are responses that maintain body temperature within an optimal range. (a): ©Dorling Kindersley/Getty Images; (b): ©Jadranko Markoc/flickr/Getty Images RF
(a): ©Design Pics/Kristy-Anne Glubish RF; (b): ©John Rowley/Getty Images RF

environment where conditions change frequently; it is extremely

hot or too cold. The cell must also take in nutrients, excrete
wastes, and regulate its many chemical reactions to prevent a
shortage or surplus of essential substances. Homeostasis is this
state of internal constancy.
Because cells maintain homeostasis by counteracting c­ hanges
as they occur, organisms must be able to sense and react to stimu-
li. To illustrate this idea, consider the mechanisms that help main-
tain your internal temperature at about 37°C (figure 1.5). When
you go outside on a cold day, you may begin to shiver; heat from
these muscle movements warms the body. In severe cold, your
lips and fingertips may turn blue as your circulatory system sends
blood away from your body’s surface. Conversely, on a hot day,
sweat evaporating from your skin helps cool your body.
D. Life Reproduces, Grows, and Develops
Organisms reproduce, making other individuals that are similar
to themselves (figure 1.6). Reproduction transmits DNA from
generation to generation; this genetic information defines the in-
herited characteristics of the offspring.
Reproduction occurs in two basic ways: asexually and sexu-
ally. In asexual reproduction, genetic information comes from
only one parent, and all offspring are virtually identical
(­figure 1.6a). One-celled organisms such as bacteria reproduce
asexually by doubling and then dividing the contents of the cell.
Many multicellular organisms also reproduce asexually. A straw-
berry plant, for instance, produces “runners” that sprout leaves
and roots, forming new plants that are identical to the parent.
Fungi produce countless asexual spores, visible as the green,
white, or black powder on moldy bread or cheese. Some animals,
including sponges, reproduce asexually when a fragment of the
common among plants, animals, and fungi.
If each offspring is to reproduce, it must grow and develop to
adulthood. Each young swan in figure 1.6b, for example, started as
a single fertilized egg cell. That cell divided over and over, devel-
oping into an embryo. Continued cell division and specialization
yielded the newly hatched swans, which will eventually mature
into adults that can also reproduce—just like their parents.
E. Life Evolves
One of the most intriguing questions in biology is how organisms
become so well suited to their environments. A beaver’s enor-
mous front teeth, which never stop growing, are ideal for gnawing
wood. Tubular flowers have exactly the right shapes for the beaks
of their hummingbird pollinators. Some organisms have color pat-
terns that enable them to fade into the background (­figure 1.7).
These examples, and countless others, illustrate adaptations.
An adaptation is an inherited characteristic or behavior that
­enables an organism to survive and reproduce successfully in
its environment.
Where do these adaptive traits come from? The answer lies in
natural selection. The simplest way to think of natural selection is
Figure 1.7 Hiding in Plain
Sight. This pygmy seahorse
is barely visible in its coral
habitat, thanks to its unique
body shape, skin color, and
texture.
©Mark Webster/Getty Images

parent animal detaches and develops into a new individual.

In sexual reproduction, genetic material from two parents
unites to form an offspring, which has a new combination of in-
herited traits (figure 1.6b). By mixing genes at each generation,
sexual reproduction results in tremendous diversity in a popula-
tion. Genetic diversity, in turn, enhances the chance that some
individuals will survive even if conditions change. Sexual repro-
duction is therefore a very successful strategy, especially in an
8 UNIT ONE Science, Chemistry, and Cells
to consider two facts. First, populations produce many more off-
spring than will survive to reproduce; these organisms must com-
pete for limited resources such as food and habitat. A single mature
oak tree may produce thousands of acorns in one season, but only
a few are likely to germinate, develop, and reproduce. The rest die.
Second, no organism is exactly the same as any other. Genetic
mutations—changes in an organism’s DNA sequence—generate
variability in all organisms, even those that reproduce asexually.
Of all the offspring in a population, which will outcompete the
others and live long enough to reproduce? The answer is those
with the best adaptations to the current environment; conversely,
the poorest competitors are most likely to die before reproducing.
A good definition of natural selection, then, is the enhanced re-
productive success of certain individuals from a population based
on inherited characteristics (figure 1.8). The same principle ap-
plies to all populations. In general, individuals with the best com-
binations of genes survive and reproduce, while those with less
suitable characteristics fail to do so. Over many generations, indi-
viduals with adaptive traits make up most or all of the population.
But the environment is constantly changing. Continents shift,
sea levels rise and fall, climates warm and cool. What happens to
a population when the selective forces that drive natural selection
change? Only some organisms survive: those with the “best” traits
in the new environment. Features that may once have been rare
become more common as the reproductive success of individuals
with those traits improves. Notice, however, that this outcome de-
pends on variability within the population. If no individual can
reproduce in the new environment, the species may go extinct.
Natural selection is one mechanism of evolution, which is a
change in the genetic makeup of a population over multiple gen-
erations. Although evolution can also occur in other ways, natural
selection is the mechanism that selects for adaptations. Charles
Generation 1
Bacterial
cell Time
Bandage
fiber
Staphylococcus aureus
before mutation
a. SEM (false color) 20 μm b.
Figure 1.8 Natural Selection. (a) Staphylococcus aureus (commonly called “staph”) is a bacterium that causes
skin infections. (b) A bacterium undergoes a random genetic mutation that (by chance) makes the cell resistant to an
antibiotic. The presence of the antibiotic increases the reproductive success of the resistant cell and its offspring.
After many generations, nearly all of the bacteria in the population are antibiotic-resistant.
Photo: (a): © Paul Gunning/Science Source
Darwin became famous in the 1860s after the publication of his
book On the Origin of Species by Means of Natural Selection,
which introduced the theory of evolution by natural selection;
another naturalist, Alfred Russel Wallace, independently devel-
oped the same idea at around the same time.
Evolution is the single most powerful idea in biology. As
unit 3 describes in detail, evolution has been operating since life
began, and it explains the current diversity of life. In fact, the
similarities among existing organisms strongly suggest that all
species descend from a common ancestor. Evolution has molded
the life that has populated the planet since the first cells formed
almost 4 billion years ago, and it continues to act today.
1.1 MASTERING CONCEPTS
1. Does any nonliving object possess all of the characteristics of life?
Explain your answer.
2. List the levels of life’s organizational hierarchy from smallest to
largest, starting with atoms and ending with the biosphere.
3. The bacteria in figure 1.8 reproduce asexually, yet they are
evolving. What is their source of genetic variation?
1.2 The Tree of Life Includes
Three Main Branches
Biologists have been studying life for centuries, documenting the
existence of everything from bacteria to blue whales. An endur-
ing problem has been how to organize the ever-growing list of
known organisms into meaningful categories. Taxonomy is the
science of naming and classifying organisms.
Generation 2 Multiple generations later
Antibiotic present
Reproduction
and
selection
Antibiotic absent
Mutation
occurs (red)
 CHAPTER 1 The Scientific Study of Life 9

The basic unit of classification is the species, which desig-
nates a distinctive “type” of organism. Closely related species are
grouped into the same genus. Together, the genus and a specific
descriptor denote the unique, two-word scientific name of each
species. A human, for example, is Homo sapiens. (Note that sci-
entific names are always italicized and that the genus—but not
the specific descriptor—is capitalized.) Scientific names help
taxonomists and other biologists communicate with one another.
But taxonomy involves more than simply naming species.
Taxonomists also strive to classify organisms according to what
we know about evolutionary relationships; that is, how recently
one type of organism shared an ancestor with another type. The
more recently they diverged from a shared ancestor, the more
closely related we presume the two types of organisms to be. Re-
searchers infer these relationships by comparing anatomical, be-
havioral, cellular, genetic, and biochemical characteristics.
Section 14.6 describes the taxonomic hierarchy in more detail.
For now, it is enough to know that genetic evidence suggests that all
species fall into one of three domains, the broadest (most inclusive)
taxonomic category. Figure 1.9 depicts the three domains: Bacte-
ria, Archaea, and Eukarya. The species in domains Bacteria and
Archaea are superficially similar to one another; all are prokaryotes,
meaning that their DNA is free in the cell and not confined to an
organelle called a nucleus. Major differences in DNA sequences
separate these two domains from each other. The third domain, Eu-
karya, contains all species of eukaryotes, which are unicellular or
multicellular organisms whose cells contain a nucleus.
The species in each domain are further subdivided into
­kingdoms; figure 1.9 shows the kingdoms within domain Eukarya.
Three of these kingdoms—Animalia, Fungi, and Plantae—are famil-
iar to most people. Within each one, organisms share the same gen-
eral strategy for acquiring energy. The plant kingdom contains
autotrophs, whereas fungi and animals are consumers that differ in
the details of how they obtain food. But the fourth group of eukary-
otes, the Protista, contains a huge collection of unrelated species. Pro-
tista is a convenient but artificial “none of the above” category for the
many species of eukaryotes that are not plants, fungi, or animals.
1.2 MASTERING CONCEPTS
1. What are the goals of taxonomy?
2. How are domains related to kingdoms?
3. List and describe the four main groups of eukaryotes.

Figure 1.9 Life’s Diversity. The three domains of life arose from a hypothetical common ancestor, shown at the base of the evolutionary tree.
Photos: (Bacteria): ©Heather Davies/Getty Images RF; (Archaea): ©Eye of Science/Science Source; (Protista): ©Melba Photo Agency/PunchStock RF; (Animalia): USDA/ARS/Scott
Bauer; (Fungi): ©Corbis RF; (Plantae) USDA/Keith Weller

DOMAIN BACTERIA DOMAIN ARCHAEA DOMAIN EUKARYA

• Cells lack nuclei (prokaryotic) • Cells lack nuclei (prokaryotic) • Cells contain nuclei (eukaryotic)
• Most are unicellular • Most are unicellular • Unicellular or multicellular

Protista (multiple kingdoms) Kingdom Animalia

• Unicellular or multicellular • Multicellular
• Autotrophs or heterotrophs • Heterotrophs (by ingestion)

TEM (false color) 1 μm SEM (false color) 1 μm

LM 200 μm
Prokaryotes DOMAIN EUKARYA

Kingdom Fungi Kingdom Plantae

Animals
• Most are multicellular • Multicellular
DOMAIN DOMAIN Fungi
BACTERIA ARCHAEA • Heterotrophs (by external • Autotrophs
Plants digestion)

Protista

Common ancestor of all life

10 UNIT ONE Science, Chemistry, and Cells
1.3 Scientists Study the
Natural World
The idea of biology as a “rapidly changing field” may seem
strange if you think of science as a collection of facts. After all,
the parts of a frog are the same now as they were 50 or 100 years
ago. But memorizing frog anatomy is not the same as thinking
scientifically. Scientists use evidence to answer questions about
the natural world. If you compare a frog to a snake, for instance,
can you determine how the frog can live in water and on land,
whereas the snake survives in the desert? Understanding anat-
omy simply gives you the vocabulary you need to ask these and
other interesting questions about life.
A. The Scientific Method Has Multiple
Interrelated Parts
Scientific knowledge arises from application of the scientific
method, which is a general way of using evidence to answer
questions and test ideas (figure 1.10). Although this diagram
may give the impression that science is a tedious, step-by-step
process, that is not at all true. Instead, science combines thinking,
detective work, communicating with other scientists, learning
from mistakes, and noticing connections between seemingly un-
related events. The resulting insights have taught us everything
we know about the natural world.
Observations and Questions The scientific method begins
with observations and questions. The observations may rely on
what we can see, hear, touch, taste, or smell, or they may be
based on existing knowledge and experimental results. Often, a
great leap forward happens when one person makes connections
between previously unrelated observations. Charles Darwin, for
example, developed the idea of natural selection by combining
his understanding of Earth’s long history with his detailed obser-
vations of organisms. Another great advance occurred decades
later, when biologists realized that mutations in DNA generate
the variation that Darwin saw but could not explain.
Hypothesis and Prediction A hypothesis is a tentative
explanation for one or more observations. The hypothesis is the
essential “unit” of scientific inquiry. To be useful, the hypothe-
sis must be testable—that is, there must be a way to collect data
that can support or reject it. Interestingly, a hypothesis cannot
be proven true, because future discoveries may contradict to-
day’s results. Nevertheless, a hypothesis becomes widely ac-
cepted when multiple lines of evidence support it, no credible
data refute it, and plausible alternative hypotheses have been
rejected.
A hypothesis is a general statement that should lead to spe-
cific predictions. Often, the prediction is written as an if–then
statement. As a simple example, suppose you hypothesize that
your lawn mower stopped working because it ran out of gas. A
reasonable prediction would be “If I put fuel into the tank, then
my lawn mower should start.”
Publish
Peer
review
Make
observations
Draw
conclusions Ask a question
Consult prior Consult prior
knowledge knowledge
Collect and
interpret data Formulate
a hypothesis
Make
predictions
Figure 1.10 Scientific Inquiry. This researcher studies tiger sharks; her
observations could lead to questions and testable hypotheses. Additional
data, combined with prior findings, can help support or reject each
hypothesis. Peer review determines w1hether the results are publishable.
Photo: © Jeff Rotman/Getty Images RF
Data Collection Investigators draw conclusions based on
data (figure 1.11). The data may come from careful observa-
tions of the natural world, an approach called discovery sci-
ence. The National Audubon Society’s annual Christmas Bird
Count is a case in point: For more than a century, thousands of
“citizen scientists” have documented the ups and downs of hun-
dreds of bird species nationwide. Another way to gather data is
to carry out an experiment to test a hypothesis under controlled
conditions (section 1.3B explores experimental design in
more detail).
Discovery and experimentation work hand in hand. As just
one example, consider the well-known connection between ciga-
rettes and lung cancer. In the late 1940s, scientists showed that
smokers are far more likely than nonsmokers to develop cancer.
Since that time, countless laboratory experiments have revealed
how the chemicals in tobacco damage living cells.
Analysis and Peer Review After collecting and interpret-
ing data, investigators decide whether the evidence supports or
falsifies the hypothesis. Often, the most interesting results are
those that are unexpected, because they provide new observa-
tions that force scientists to rethink their hypotheses; figure 1.10
shows this feedback loop. Science advances as new information
arises and explanations continue to improve.
Once a scientist has enough evidence to support or reject a
hypothesis, he or she may write a paper and submit it for publica-
tion in a scientific journal. The journal’s editors send the paper to
 CHAPTER 1 The Scientific Study of Life 11

anonymous reviewers who are knowledgeable about the research

topic. In a process called peer review, these scientists indepen-
dently evaluate the validity of the methods, data, and conclu-
sions. Overall, peer review ensures that journal articles—the
tangible products of the global scientific conversation—are of
high quality.

B. An Experimental Design Is a Careful Plan

Scientists test many hypotheses with the help of experiments. An
experiment is an investigation carried out in controlled condi-
tions. This section considers a real study that tested the hypothe-
sis that a new vaccine protects against a deadly virus. The virus,
called rotavirus, causes severe diarrhea and takes the lives of
hundreds of thousands of young children each year. An effective,
inexpensive vaccine would prevent many childhood deaths.

a. Sample Size One of the most important decisions that an in-

vestigator makes in designing an experiment is the sample size,
which is the number of individuals assigned to each treatment.
The sample size in the rotavirus study, for example, was approxi-
mately 100 infants per treatment. In general, the larger the sam-
ple size, the more credible the results of a study.

Variables A systematic consideration of variables is also im-

b.
Figure 1.11 Different Types of Science. (a) Scientists track the number
of migratory birds that visit a wildlife refuge each year—an example of
discovery science. (b) Controlled experiments can help food scientists
objectively compare different techniques for roasting or brewing coffee.
(a): U.S. Fish & Wildlife Service/J&K Hollingsworth; (b): ©Corbis RF
portant in experimental design (table 1.2). A variable is a
changeable element of an experiment, and there are several
types. The independent variable is the factor that an investiga-
tor directly manipulates to determine whether it causes another
variable to change. In the rotavirus study, the independent vari-
able was the dose of the vaccine. The dependent variable is any
response that might depend on the value of the independent vari-
able, such as the number of children with rotavirus-related ill-
ness during the study period.
A standardized variable is anything that the investigator
holds constant for all subjects in the experiment, ensuring the
best chance of detecting the effect of the independent variable.
Because rotavirus infection is most common among very young
children, the test of the new vaccine included only infants ­younger
than 12 weeks. Furthermore, vaccines work best in people with
healthy immune systems, so the study excluded infants who were
ill or had weak immunity. Age and health were therefore among
the study’s standardized variables.

TA B L E 1.2 Types of Variables in an Experiment: A Summary

Type of Variable
Independent variable
Dependent variable
Standardized variable
Definition Example
A variable that an investigator manipulates to determine whether it Dose of vaccine
influences the dependent variable
A variable that an investigator measures to determine whether it is affected Number of children with illness caused by
by the independent variable rotavirus
Any variable that an investigator intentionally holds constant for all subjects Age and health of children in study
in an experiment, including the control group
12 UNIT ONE Science, Chemistry, and Cells

Controls Well-designed experiments compare one or more C. Theories Are Comprehensive

groups undergoing treatment to a group of “normal” individuals.
The experimental control is the untreated group, and it is impor-
Explanations
tant because it provides a basis for comparison in measuring the Outside of science, the word theory is often used to describe an
effect of the independent variable. Ideally, the only difference opinion or a hunch. For instance, immediately after a plane crash,
between the control and any other experimental group is the one experts offer their “theories” about the cause of the disaster.
factor being tested. These tentative explanations are really untested hypotheses.
Experimental controls may take several forms. Sometimes, In science, the word theory has a distinct meaning. Like a hy-
the control group simply receives a “zero” value for the indepen- pothesis, a theory is an explanation for a natural phenomenon, but
dent variable. If a gardener wants to test a new fertilizer in her a theory is typically broader in scope than a hypothesis. For ex-
garden, she may give some plants a lot of fertilizer, others only a ample, the germ theory—the idea that some microorganisms cause
little, and still others—the control plants—none. In medical re- human disease—is the foundation for medical microbiology. Indi-
search, a control group might receive a placebo, an inert substance vidual hypotheses relating to the germ theory are much narrower,
that resembles the treatment given to the experimental group. The such as the suggestion that rotavirus causes illness. Not all theories
control infants in the rotavirus study received a placebo that con- are as “large” as the germ theory, but they generally encompass
tained all components of the vaccine except the active ingredient. multiple hypotheses. Note also that the germ theory does not imply
The investigators in the rotavirus study used a double-blind that all microbes make us sick or that all illnesses have microbial
design, in which neither the researchers nor the participants knew causes. But it does explain many types of human disease.
who received the vaccine and who received the placebo. Double- A second difference between a hypothesis and a theory is
blind studies help avoid bias in medical research. The investiga- acceptance and evidence. A hypothesis is tentative, whereas the-
tors break the “code” of who received which treatment only after ories reflect broader agreement. This is not to imply that theories
the experiment is complete and the data are tabulated. are not testable; in fact, the opposite is true. Every scientific
theory is potentially falsifiable, meaning that a particular set of
Statistical Analysis Once an experiment is complete, the inves- observations could prove the theory wrong. The germ theory re-
tigator compiles the data and decides whether the results support the mains widely accepted because many observations support it and
hypothesis (figure 1.12). The researchers in the rotavirus study con- no reliable tests have disproved it. The same is true for the theory
cluded that the vaccine was effective, but only after applying a sta- of evolution and many other scientific theories.
tistical test. All statistical tests consider both variation and sample Another quality of a scientific theory is its predictive power.
size to estimate the probability that the results arose purely by A good theory not only ties together many existing observations
chance. If this probability is low, then the results are considered but also suggests predictions about phenomena that have yet to be
­statistically significant. Appendix B shows how scientists use error observed. Both Charles Darwin and naturalist Alfred Russel
bars and other notation to illustrate statistical significance in graphs. Wallace used the theory of evolution by natural selection to pre-
dict the existence of a moth that could pollinate orchid flowers
with unusually long nectar tubes (figure 1.13). Decades later,
scientists discovered the long-tongued insect (see section 1.4).
This finding was consistent with evolutionary theory, but the
theory would have been weakened if subsequent research had not
Incidence of illness (cases/100 child-years)

30
25.86
Any rotavirus illness supported its predictions.
Severe rotavirus illness
25 What is the relationship between facts and a theory? One
definition of the word fact is “a repeatable observation that ev-
Dependent variable

20 eryone can agree on.” It is a fact that a dropped pencil falls to-
ward the ground; no reasonable person disagrees with that
15 14.46 statement. Gravity is a fact; gravitational theory explains the
forces that cause pencils and other objects to fall.
10 Biologists also consider biological evolution to be a fact. Yet
6.19 6.86 the phrase “theory of evolution” persists because evolution is
5 both a fact and a theory. Like gravity, evolution is a fact. No one
2.15 2.15
can dispute that antibiotics drive evolutionary change in bacteria
0 0
0 (see figure 1.8). On a broader scale, the combined evidence for
Placebo Low Medium High
(control) genetic change over time is so persuasive and comes from so
many different fields of study that to deny the existence of evolu-
Dose of vaccine
tion is unrealistic. Evolutionary theory explains how life has di-
Independent variable versified since its origin. Note that biologists do not understand
everything about how evolution works. Many questions about
Figure 1.12 Vaccine Test. This graph shows that the rotavirus vaccine is life’s history remain, but the debates swirl around how, not
more effective than a placebo. (The statistical analysis is not shown.) whether, evolution occurs.

Nectar tubes
Figure 1.13 Prediction Confirmed. When Charles Darwin saw this
Madagascar orchid, he predicted that its pollinator would have long, thin
mouthparts that could reach the bottom of the elongated nectar tube.
He was right; the unknown pollinator turned out to be a moth with an
extraordinarily long tongue.
©Kjell Sandved/Alamy
D. Scientific Inquiry Has Limitations
Scientific inquiry is neither foolproof nor always easy to imple-
ment. One problem is that experimental evidence may lead to
multiple interpretations, and even the most carefully designed
experiment can fail to provide a definitive answer (see the Apply
It Now box in this section). Consider the observation that animals
fed large doses of vitamin E live longer than similar animals that
do not ingest the vitamin. So, does vitamin E slow aging?
Burning Question
Why am I here?
The Burning Questions featured in each chapter of this book came
from students. On the first day of class, I always ask students to turn in
a “burning question”—anything they have always wondered about bi-
ology. I answer most of the questions as the relevant topics come up
during the semester.
Why not answer all of the questions? It is because at least one
student often asks something like “Why am I here?” or “What is the
meaning of life?” Such puzzles have fascinated humans throughout the
ages, but they are among the many questions that we cannot approach
scientifically. Biology can explain how you developed after a sperm
from your father fertilized an egg cell from your mother. But no one
can develop a testable hypothesis about life’s meaning or the purpose
of human existence. Science must remain silent on such questions.
Instead, other ways of knowing must satisfy our curiosity about
“why.” Philosophers, for example, can help us see how others have
CHAPTER 1 The Scientific Study of Life 13
Possibly, but excess vitamin E also causes weight loss, and other
research has connected weight loss with longevity. Does
­vitamin E extend life, or does weight loss? The experiment alone
does not distinguish between these possibilities.
Another limitation is that researchers may misinterpret ob-
servations or experimental results. For example, centuries ago,
scientists sterilized a bottle of broth, corked the bottle shut, and
observed bacteria in the broth a few days later. They concluded
that life arose directly from the broth. The correct explanation,
however, was that the cork did not keep airborne bacteria out.
Although scientists may make mistakes in the short term, science
is self-correcting in the long run because it remains open to new
data and new interpretations.
A related problem is that the scientific community may be
slow to accept new evidence that suggests unexpected conclu-
sions. Every investigator should try to keep an open mind about
observations, not allowing biases or expectations to cloud inter-
pretation of the results. But it is human nature to be cautious in
accepting an observation that does not fit what we think we
know. The careful demonstration that life does not arise from
broth surprised many people who believed that mice sprang
from moldy grain and that flies came from rotted beef. More
recently, it took many years to set aside the common belief that
stress caused ulcers. Today, we know that a bacterium causes
most ulcers.
Although science is a powerful tool for answering questions
about the natural world, it cannot answer questions of beauty,
morality, ethics, or the meaning of life (see this section’s B
­ urning
Question). Nor can we directly study some phenomena that oc-
curred long ago and left little physical evidence. Consider the
many experiments that have attempted to recreate the chemical
reactions that might have produced life on early Earth. Although
the experiments produce interesting results and reveal ways that
these early events may have occurred, we cannot know if they
accurately reflect conditions at the dawn of life.
c­onsidered these questions. Religion may
also provide the meaning that many people
seek. Part of the value of higher educa-
tion is to help you acquire the tools you
need to find your own life’s purpose.
In the meantime, this book’s
Burning Questions can help you dis-
cover the answers to many ques-
tions—asked by students just like
you—about human health, environ-
mental quality, life’s diversity, and the
rest of the biological world.
©Getty Images/Photodisc RF
Submit your burning question to
Marielle.Hoefnagels@mheducation.com
14 UNIT ONE Science, Chemistry, and Cells

Apply It Now
It’s Hard to Know What’s Bad for You
You have probably heard reports that a food previously
considered healthy is actually bad for you, or vice
versa. These conflicting reports may tempt you to mis-
takenly conclude that scientific studies are no better than
guesswork.
Instead, the problem lies in the fact that some questions
are extremely hard to answer. Take, for example, the contro-
versy surrounding the artificial sweetener called saccharin.
In 1977, the U.S. Food and Drug Administration (FDA) pro-
posed a ban on saccharin, based on a handful of studies suggest-
ing that the sweetener caused bladder cancer in rats. Congress
opted to require warning labels on products containing ­saccharin.
In 1991, the FDA withdrew its proposed ban, and in 1998,
­saccharin was rated as “not classifiable as to its carcinogenicity
to humans.” Two years later, legislation removed the warning
label requirement.
This tangled history raises an important issue. Why can’t
©Photodisc/Getty Images RF
science reply “yes” or “no” to the seemingly simple question
of whether saccharin is bad for you? To understand the an-
swer, consider one of the studies that prompted the FDA to propose the
ban on saccharin in the first place. Researchers divided 200 rats into
control rats by the end of the study, suggesting that high doses of the
sweetener are toxic. Rather than causing cancer directly, the saccharin
may have simply made the animals more susceptible to disease. More-
over, follow-up studies using other animals were inconclusive.
Perhaps the scientists should have studied the saccharin–­cancer
connection in humans instead. Unfortunately, however, docu-
menting a link between any food and cancer in people is ex-
tremely difficult. One strategy might be to measure the
incidence of cancer in saccharin users versus nonusers. But
with so many other possible causes of cancer—smoking,
poor diet, exposure to job-related chemicals, genetic pre-
disposition—it is difficult to separate out just the effects
of saccharin.
So what are we to make of the mixed news reports?
It is hard to know, but one thing is certain: No matter
what the headlines say, one study, especially a small one,
cannot reveal the whole story.
Male rats Female rats
30 30
27
Saccharin-fed
25 25
Controls
% with tumors

% with tumors
two groups. The control animals ate standard rodent chow, whereas the 20 19 20
experimental group got the same food supplemented with saccharin. 15 15
At reproductive maturity the animals were bred, and the researchers
fed the offspring the same dose of saccharin throughout their lives as 10 10
well. To measure the incidence of cancer, they counted the tumors in 5 3 5 4
both generations of rats for 24 months or until the rats died, whichever 0
0
0
0 0 0
came first. Figure 1.A summarizes the results. Parents Offspring Parents Offspring
At first glance, the conclusion seems inescapable: Saccharin causes
cancer in male lab rats. But closer study reveals several hidden com- Figure 1.A The Saccharin Scare. These graphs summarize the results
plexities that make the data hard to interpret. First, the dose of saccharin of one study examining the link between saccharin and bladder cancer in
was huge: 5% of the rats’ diets, for life. The equivalent dose in humans rats. Sample sizes ranged from 36 to 49 rats per treatment.
would require drinking hundreds of cans of saccharin-sweetened soda Source: Data adapted from Office of Technology Assessment Report, October
every day. In addition, the experimental rats weighed much less than the 1977, Cancer Testing Technology and Saccharin, page 52.

E. Biology Continues to Advance
Science is just one of many ways to investigate the world, but its
strength is its openness to new information. Theories change to
accommodate new knowledge. The history of science is full of
long-established ideas that changed as we learned more about
nature, often thanks to new technology. People thought that Earth
was flat and at the center of the universe before inventions and
data analysis revealed otherwise. Similarly, biologists thought all
organisms were plants or animals until microscopes unveiled a
world of life invisible to our eyes.
Technology is the practical application of scientific knowl-
edge. Science and technology are therefore intimately related.
For example, thanks to centuries of scientific inquiry, we under-
stand many of the differences between humans and bacteria. We
can exploit these differences to invent new antibiotic drugs that
kill germs without harming our own bodies. These antibiotics, in
turn, can be useful tools that help biologists learn even more
about bacterial cells. The new scientific discoveries spawn new
technologies, and so on.
Biology is changing rapidly because technology has e­ xpanded
our ability to spy on living cells, compare DNA sequences, track
wildlife, and make many other types of observations. Scientists
can now answer questions about the natural world that previous
generations could never have imagined.
1.3 MASTERING CONCEPTS
1. Identify the elements of the experiment summarized in the
Apply It Now box.
2. What is a statistically significant result?
3. What is the difference between a hypothesis and a theory, and why
are some theories regarded as facts?
4. What are some limitations of scientific inquiry?
5. Compare and contrast science and technology.
Another random document with
no related content on Scribd:
The “Essay on Man” has been praised and admired by men of the
most opposite beliefs, and men of no belief at all. Bishops and free-
thinkers have met here on a common ground of sympathetic
approval. And, indeed, there is no particular faith in it. It is a droll
medley of inconsistent opinions. It proves only two things beyond a
question: that Pope was not a great thinker; and that wherever he
found a thought, no matter what, he would express it so tersely, so
clearly, and with such smoothness of versification, as to give it an
everlasting currency. Hobbes’s unwieldy “Leviathan,” left stranded on
the shore of the last age and nauseous with the stench of its
selfishness—from this Pope distilled a fragrant oil with which to fill
the brilliant lamps of his philosophy, lamps like those in the tombs of
alchemists, that go out the moment the healthy air is let in upon
them. The only positive doctrine in the poem is the selfishness of
Hobbes set to music, and the pantheism of Spinoza brought down
from mysticism to commonplace. Nothing can be more absurd than
many of the dogmas taught in the “Essay on Man.”

The accuracy on which Pope prided himself, and for which he is

commended, was not accuracy of thought so much as of expression.
But the supposition is that in the “Essay on Man” Pope did not know
what he was writing himself. He was only the condenser and
epigrammatizer of Bolingbroke—a fitting St. John for such a gospel.
Or if he did know, we can account for the contradictions by
supposing that he threw in some of the commonplace moralities to
conceal his real drift. Johnson asserts that Bolingbroke in private
laughed at Pope’s having been made the mouthpiece of opinions
which he did not hold. But this is hardly probable when we consider
the relations between them. It is giving Pope altogether too little
credit for intelligence to suppose that he did not understand the
principles of his intimate friend.

Dr. Warburton makes a rather lame attempt to ward off the charge of
Spinozism from the “Essay on Man.” He would have found it harder
to show that the acknowledgment of any divine revelation would not
overthrow the greater part of its teachings. If Pope intended by his
poem all that the Bishop takes for granted in his commentary, we
must deny him what is usually claimed as his first merit—clearness.
If we did not, we grant him clearness as a writer at the expense of
sincerity as a man. Perhaps a more charitable solution of the
difficulty is that Pope’s precision of thought was not equal to his
polish of style.

But it is in his “Moral Essays” and part of his “Satires” that Pope
deserves the praise which he himself desired—

Happily to steer
From grave to gay, from lively to severe.
Correct with spirit, eloquent with ease,
Intent to reason, or polite to please.

Here Pope must be allowed to have established a style of his own, in

which he is without a rival. One can open upon wit and epigram at
every page.

In his epistle on the characters of woman, no one who has ever

known a noble woman will find much to please him. The climax of his
praise rather degrades than elevates:

O blest in temper, whose unclouded ray

Can make to-morrow cheerful as to-day,
She who can love a sister’s charms, or hear
Sighs for a daughter with unwounded ear,
She who ne’er answers till a husband cools,
Or if she rules him, never shows she rules,
Charms by accepting, by submitting sways,
Yet has her humor most when she obeys;
Let fops or fortune fly which way they will,
Disdains all loss of tickets, or codille,
Spleen, vapors, or smallpox, above them all;
And mistress of herself though china fall.
The last line is very witty and pointed; but consider what an ideal of
womanly nobleness he must have had who praises his heroine for
not being jealous of her daughter.

It is very possible that the women of Pope’s time were as bad as

they could be, but if God made poets for anything it was to keep
alive the traditions of the pure, the holy, and the beautiful. I grant the
influence of the age, but there is a sense in which the poet is of no
age, and Beauty, driven from every other home, will never be an
outcast and a wanderer while there is a poet-nature left; will never
fail of the tribute at least of a song. It seems to me that Pope had a
sense of the nice rather than of the beautiful. His nature delighted in
the blemish more than in the charm.

Personally, we know more about Pope than about any of our poets.
He kept no secret about himself. If he did not let the cat out of the
bag, he always contrived to give her tail a pinch so that we might
know she was there. In spite of the savageness of his satires, his
disposition seems to have been a truly amiable one, and his
character as an author was as purely fictitious as his style. I think
that there was very little real malice in him.

A great deal must be allowed to Pope for the age in which he lived,
and not a little, I think, for the influence of Swift. In his own province
he still stands unapproachably alone. If to be the greatest satirist of
individual men rather than of human nature; if to be the highest
expression which the life of court and the ball-room has ever found in
verse; if to have added more phrases to our language than any other
but Shakspeare; if to have charmed four generations makes a man a
great poet, then he is one. He was the chief founder of an artificial
style of writing which in his hand was living and powerful because he
used it to express artificial modes of thinking and an artificial state of
society. Measured by any high standard of imagination, he will be
found wanting; tried by any test of wit, he is unrivaled.

To what fatuities his theory of correctness led in the next generation,

when practised upon by men who had not his genius, I shall
endeavor to show in my next lecture.
 LECTURE X
POETIC DICTION

(Friday Evening, February 9, 1855)

X
No one who has read any early poems, of whatever nation, can have
failed to notice a freshness in the language—a sort of game flavor,
as it were—that gradually wastes out of it when poetry becomes
domesticated, so to speak, and has grown to be a mere means of
amusement both to writers and readers, instead of answering a
deeper necessity in their natures. Our Northern ancestors
symbolized the eternal newness of song by calling it the Present,
and its delight by calling it the drink of Odin.

There was then a fierce democracy of words; no grades had then

been established, and no favored ones advanced to the Upper
House of Poetry. Men had a meaning, and so their words had to
have one, too. They were not representatives of value, but value
itself. They say that Valhalla was roofed with golden shields; that
was what they believed, and in their songs they called them golden
shingles. We should think shields the more poetical word of the two;
but to them the poetry was in the thing, and the thought of it and the
phrase took its life and meaning from them.

It is one result of the admixture of foreign words in our language that

we use a great many phrases without knowing the force of them.
There is a metaphoric vitality hidden in almost all of them, and we
talk poetry as Molière’s citizen did prose, without ever suspecting it.
Formerly men named things; now we merely label them to know
them apart. The Vikings called their ships sea-horses, just as the
Arabs called their camels ships of the desert. Capes they called sea-
noses, without thinking it an undignified term which the land would
resent. And still, where mountains and headlands have the luck to be
baptized by uncultivated persons, Fancy stands godmother. Old
Greylock, up in Berkshire, got his surname before we had State
geologists or distinguished statesmen. So did Great Haystack and
Saddle-Mountain. Sailors give good names, if they have no
dictionary aboard, and along our coasts, here and there, the word
and the thing agree, and therefore are poetical. Meaning and poetry
still cling to some of our common phrases, and the crow-foot,
mouse-ear, goat’s-beard, day’s-eye, heart’s-ease, snow-drop, and
many more of their vulgar little fellow-citizens of the wood and
roadsides are as happy as if Linnæus had never been born. Such
names have a significance even to one who has never seen the
things they stand for, but whose fancy would not be touched about a
pelargonium unless he had an acquired sympathy with it. Our
“cumulus” language, heaped together from all quarters, is like the
clouds at sunset, and every man finds something different in a
sentence, according to his associations. Indeed, every language that
has become a literary one may be compared to a waning moon, out
of which the light of beauty fades more and more. Only to poets and
lovers does it repair itself from its luminous fountains.

The poetical quality of diction depends on the force and intensity of

meaning with which it is employed. We are all of us full of latent
significance, and let a poet have but the power to touch us, we
forthwith enrich his word with ourselves, pouring into his verse our
own lives, all our own experience, and take back again, without
knowing it, the vitality which we had given away out of ourselves. Put
passion enough into a word, and no matter what it is it becomes
poetical; it is no longer what it was, but is a messenger from original
man to original man, an ambassador from royal Thee to royal Me,
and speaks to us from a level of equality. Pope, who did not scruple
to employ the thoughts of Billingsgate, is very fastidious about the
dress they come in, and claps a tawdry livery-coat on them, that they
may be fit for the service of so fine a gentleman. He did not mind
being coarse in idea, but it would have been torture to him to be
thought commonplace. The sin of composition which he dreaded
was,

Lest ten low words should creep in one dull line.

But there is no more startling proof of the genius of Shakspeare than

that he always lifts the language up to himself, and never thinks to
raise himself atop of it. If he has need of the service of what is called
a low word, he takes it, and it is remarkable how many of his images
are borrowed out of the street and the workshop. His pen ennobled
them all, and we feel as if they had been knighted for good service in
the field. Shakspeare, as we all know (for does not Mr. Voltaire say
so?), was a vulgar kind of fellow, but somehow or other his genius
will carry the humblest things up into the air of heaven as easily as
Jove’s eagle bore Ganymede.

Whatever is used with a great meaning, and conveys that meaning

to others in its full intensity, is no longer common and ordinary. It is
this which gives their poetic force to symbols, no matter how low
their origin. The blacksmith’s apron, once made the royal standard of
Persia, can fill armies with enthusiasm and is as good as the
oriflamme of France. A broom is no very noble thing in itself, but at
the mast-head of a brave old De Ruyter, or in the hands of that awful
shape which Dion the Syracusan saw, it becomes poetical. And so
the emblems of the tradesmen of Antwerp, which they bore upon
their standards, pass entirely out of the prosaic and mechanical by
being associated with feelings and deeds that were great and
momentous.

Mr. Lowell here read a poem by Dr. Donne entitled “The Separation.”
As respects Diction, that becomes formal and technical when poetry
has come to be considered an artifice rather than an art, and when
its sole object is to revive certain pleasurable feelings already
conventional, instead of originating new sources of delight. Then it is
truly earth to earth; dead language used to bury dead emotion in.
This kind of thing was carried so far by the later Scandinavian poets
that they compiled a dictionary of the metaphors used by the elder
Skalds (whose songs were the utterance of that within them which
would be spoken), and satisfied themselves with a new arrangement
of them. Inspiration was taught, as we see French advertised to be,
in six lessons.

In narrative and descriptive poetry we feel that proper keeping

demands a certain choice and luxury of words. The question of
propriety becomes one of prime importance here. Certain terms
have an acquired imaginative value from the associations they
awake in us. Certain words are more musical than others. Some
rhymes are displeasing; some measures wearisome. Moreover,
there are words which have become indissolubly entangled with
ludicrous or mean ideas. Hence it follows that there is such a thing
as Poetic Diction, and it was this that Milton was thinking of when he
spoke of making our English “search her coffers round.”

I will illustrate this. Longfellow’s “Evangeline” opens with a noble

solemnity:

This is the forest primeval; the murmuring pines and the hemlocks,
Bearded with moss and in garments green, indistinct in the twilight,
Stand like the Druids of eld, with voices sad and prophetic,
Stand like harpers hoar, with beards that rest on their bosoms.
Loud from its rocky caverns the deep-voiced neighboring ocean
Speaks, and in accents disconsolate answers the wail of the forest.

There is true feeling here, and the sigh of the pines is heard in the
verses. I can find only one epithet to hang a criticism on, and that is
the “wail of the forest” in the last line, which is not in keeping with the
general murmur. Now I do not suppose that the poet turned over any
vocabulary to find the words he wanted, but followed his own poetic
instinct altogether in the affair. But suppose for a moment, that
instead of being a true poet, he had been only a gentleman
versifying; suppose he had written, “This is the primitive forest.” The
prose meaning is the same, but the poetical meaning, the music, and
the cadence would be gone out of it, and gone forever. Or suppose
that, instead of “garments green,” he had said “dresses green”; the
idea is identical, but the phrase would have come down from its
appropriate remoteness to the milliner’s counter. But not to take such
extreme instances, only substitute instead of “harpers hoar,” the
words “harpers gray,” and you lose not only the alliteration, but the
fine hoarse sigh of the original epithet, which blends with it the
general feeling of the passage. So if you put “sandy beaches” in the
place of “rocky caverns,” you will not mar the absolute truth to
nature, but you will have forfeited the relative truth to keeping.

When Bryant says so exquisitely,

Painted moths
Have wandered the blue sky and died again,

we ruin the poetry, the sunny spaciousness of the image, without

altering the prose sense, by substituting

Have flown through the clear air.

But the words “poetic diction” have acquired a double meaning, or

perhaps I should say there are two kinds of poetic diction, the one
true and the other false, the one real and vital, the other mechanical
and artificial. Wordsworth for a time confounded the two together in
one wrathful condemnation, and preached a crusade against them
both. He wrote, at one time, on the theory that the language of
ordinary life was the true dialect of poetry, and that one word was as
good as another. He seemed even to go farther and to adopt the
Irishman’s notion of popular equality, that “one man is as good as
another, and a dale better, too.” He preferred, now and then, prosaic
words and images to poetical ones. But he was not long in finding
his mistake and correcting it. One of his most tender and pathetic
poems, “We are Seven,” began thus in the first edition:

A simple child, dear brother Jim.

All England laughed, and in the third edition Wordsworth gave in and
left the last half of the line blank, as it has been ever since. If the
poem had been a translation from the Turkish and had begun,

A simple child, dear Ibrahim,

there would have been nothing unpoetical in it; but the “dear brother
Jim,” which would seem natural enough at the beginning of a familiar
letter, is felt to be ludicrously incongruous at the opening of a poem.

To express a profound emotion, the simpler the language and the

less removed from the ordinary course of life the better. There is a
very striking example of this in Webster’s tragedy of “The Duchess of
Malfy.” The brother of the Duchess has procured her murder, and
when he comes in and sees the body he merely says:

“Cover her face; mine eyes dazzle; she died young.”

Horror could not be better expressed than in these few words, and
Webster has even taken care to break up the verse in such a way
that a too entire consciousness of the metre may not thrust itself
between us and the bare emotion he intends to convey.
In illustration, Mr. Lowell quoted from Shakspeare (“Henry V”),
Marlowe, Chapman, Dunbar, Beaumont and Fletcher, Waller, Young,
and Cawthorn.

These men [the poets of the eighteenth century] were perfectly

conscious of the fact that poetry is not produced under an ordinary
condition of the mind, and accordingly, when they begin to grind their
barrel-organs, they go through the ceremony of invoking the Muse,
talk in the blandest way of divine rages and sacred flames, and one
thing or another, and ask for holy fire to heat their little tea-urns with
as coolly as one would borrow a lucifer. They appeal ceremoniously
to the “sacred Nine,” when the only thing really necessary to them
was the ability to count as high as the sacred ten syllables that
constituted their verse. If the Muse had once granted their prayer, if
she had once unveiled her awful front to the poor fellows, they would
have hidden under their beds, every man John of them.

The eighteenth century produced some true poets, but almost all,
even of them, were infected by the prevailing style. I cannot find any
name that expresses it better than the “Dick Swiveller style.” As Dick
always called wine the “rosy,” sleep the “balmy,” and so forth, so did
these perfectly correct gentlemen always employ either a fluent
epithet or a diffuse paraphrasis to express the commonest emotions
or ideas. If they wished to say tea they would have done it thus:

Of China’s herb the infusion hot and mild.

Coffee would be

The fragrant juice of Mocha’s kernel gray,

or brown or black, as the rhyme demanded. A boot is dignified into

The shining leather that the leg encased.

Wine is

The purple honor of th’ ambrosial vine.

All women are “nymphs,” carriages are “harnessed pomps,” houses

are sumptuous or humble “piles,” as the case may be, and
everything is purely technical. Of nature there seems to have been
hardly a tradition.

But instead of attempting to describe in prose the diluent diction

which passed for poetic under the artificial system—which the
influence of Wordsworth did more than anything else to abolish and
destroy—I will do it by a few verses in the same style. Any subject
will do—a Lapland sketch, we will say:

Where far-off suns their fainter splendors throw

O’er Lapland’s wastes of uncongenial snow,
Where giant icebergs lift their horrent spires
And the blank scene a gelid fear expires,
Where oft the aurora of the northern night
Cheats with pale beams of ineffectual light,
Where icy Winter broods o’er hill and plain,
And Summer never comes, or comes in vain;
Yet here, e’en here, kind Nature grants to man
A boon congenial with her general plan.
Though no fair blooms to vernal gales expand,
And smiling Ceres shuns th’ unyielding land,
Behold, even here, cast up a monstrous spoil,
The sea’s vast monarch yields nutritious oil,
Escaped, perchance, from where the unfeeling crews
Dart the swift steel, and hempen coils unloose,
He whirls impetuous through the crimson tide,
Nor heeds the death that quivers in his side;
Northward he rushes with impulsive fin,
Where shores of crystal groan with ocean’s din,
Shores that will melt with pity’s glow more soon
Than the hard heart that launched the fierce harpoon.
In vain! he dies! yet not without avail
The blubbery bulk between his nose and tail.
Soon shall that bulk, in liquid amber stored,
Shed smiling plenty round some Lapland board.
Dream not, ye nymphs that flutter round the tray
When suns declining shut the door of day,
While China’s herb, infused with art, ye sip,
And toast and scandal share the eager lip.
Dream not to you alone that Life is kind,
Nor Hyson’s charms alone can soothe the mind;
If you are blest, ah, how more blest is he
By kinder fate shut far from tears and tea,
Who marks, replenished by his duteous hand,
Dark faces oleaginously expand;
And while you faint to see the scalding doom
Invade with stains the pride of Persia’s loom,
Happier in skins than you in silks perhaps,
Deals the bright train-oil to his little Lap’s.
 LECTURE XI
WORDSWORTH

(Tuesday Evening, February 13, 1855)

XI
A few remarks upon two of the more distinguished poets of the
eighteenth century will be a fitting introduction to Wordsworth, and,
indeed, a kind of commentary on his poetry. Of two of these poets
we find very evident traces in him—Thomson and Cowper—of the
one in an indiscriminating love of nature, of the other in a kind of
domestic purity, and of both in the habit of treating subjects
essentially prosaic, in verse; whence a somewhat swelling wordiness
is inevitable.

Thomson had the good luck to be born in Scotland, and to be

brought up by parents remarkable for simplicity and piety of life.
Living in the country till he was nearly twenty, he learned to love
natural beauty, and must have been an attentive student of scenery.
That he had true instincts in poetry is proved by his making Milton
and Spenser his models. He was a man of force and originality, and
English poetry owes him a large debt as the first who stood out both
in precept and practice against the vicious artificial style which then
reigned, and led the way back to purer tastes and deeper principles.
He was a man perfectly pure in life; the associate of eminent and
titled personages, without being ashamed of the little milliner’s shop
of his sisters in Edinburgh; a lover of freedom, and a poet who never
lost a friend, nor ever wrote a line of which he could repent. The
licentiousness of the age could not stain him. His poem of “Winter”
was published a year before the appearance of the “Dunciad.”

Thomson’s style is not equal to his conceptions. It is generally

lumbering and diffuse, and rather stilted than lofty. It is very likely that
his Scotch birth had something to do with this, and that he could not
write English with that unconsciousness without which elegance is
out of the question—for there can be no true elegance without
freedom. Burns’s English letters and poems are examples of this.

But there are passages in Thomson’s poems full of the truest

feelings for nature, and gleams of pure imagination.

Mr. Lowell here read a passage from “Summer,” which, he said,

illustrated better than almost any other his excellences and defects.
It is a description of a storm, beginning:

At first heard solemn o’er the verge of Heaven

The tempest growls.

This is fustian patched with cloth of gold. The picture, fine as it is in

parts, is too much frittered with particulars. The poet’s imagination
does not seem powerful enough to control the language. There is no
autocratic energy, but the sentences are like unruly barons, each
doing what he likes in his own province. Many of them are prosaic
and thoroughly unpicturesque, and come under the fatal
condemnation of being flat. Yet throughout the passage,

The unconquerable genius struggles through

half-suffocated in a cloud of words.

But the metre is hitchy and broken, and seems to have no law but
that of five feet to the verse. There is no Pegasean soar, but the
unwieldy gallop of an ox. The imagination, which Thomson
undoubtedly had, contrasted oddly with the lumbering vehicle of his
diction. He takes a bushel-basket to bring home an egg in. In him
poetry and prose entered into partnership, and poetry was the
sleeping partner who comes down now and then to see how the
business is getting on. But he had the soul of a poet, and that is the
main thing.

Of Gray and Collins there is no occasion to speak at length in this

place. Both of them showed true poetic imagination. In Gray it was
thwarted by an intellectual timidity that looked round continually for
precedent; and Collins did not live long enough to discharge his mind
thoroughly of classic pedantry; but both of them broke away from the
reigning style of decorous frigidity. Collins’s “Ode to Evening” is
enough to show that he had a sincere love of nature—but generally
the scenery of both is borrowed from books.

In Cowper we find the same over-minuteness in describing which

makes Thomson wearisome, but relieved by a constant vivacity of
fancy which in Thomson was entirely wanting. But Cowper more
distinctly preluded Wordsworth in his delight in simple things, in
finding themes for his song in the little incidents of his own fireside
life, or his daily walks, and especially in his desire to make poetry a
means of conveying moral truth. The influence of Cowper may be
traced clearly in some of Wordsworth’s minor poems of pure fancy,
and there is one poem of his—that on “Yardly Oak”—which is almost
perfectly Wordsworthian. But Cowper rarely rises above the region of
fancy, and he often applied verse to themes that would not sing. His
poetry is never more than agreeable, and never reaches down to the
deeper sources of delight. Cowper was one of those men who,
wanting a vigorous understanding to steady the emotional part of his
nature, may be called peculiar rather than original. Great poetry can
never be made out of a morbid temperament, and great wits are
commonly the farthest removed from madness. But Cowper had at
least the power of believing that his own thoughts and pleasures
were as good, and as fit for poetry, as those of any man, no matter
how long he had enjoyed the merit of being dead.
The closing years of the eighteenth century have something in
common with those of the sixteenth. The air was sparkling with moral
and intellectual stimulus. The tremble of the French Revolution ran
through all Europe, and probably England, since the time of the great
Puritan revolt, had never felt such a thrill of national and indigenous
sentiment as during the Napoleonic wars. It was a time fitted to give
birth to something original in literature. If from the collision of minds
sparks of wit and fancy fly out, the shock and jostle of great events,
of world-shaping ideas, and of nations who do their work without
knowing it, strike forth a fire that kindles heart and brain and tongue
to more inspired conceptions and utterances.

It was fortunate for Wordsworth that he had his breeding in the

country, and not only so, but among the grandest scenery of
England. His earliest associates were the mountains, lakes, and
streams of his native district, and the scenery with which his mind
was stored during its most impressionable period was noble and
pure. The people, also, among whom he grew up were a simple and
hardy race, who kept alive the traditions and many of the habits of a
more picturesque time. There was also a general equality of
condition which kept life from becoming conventional and trite, and
which cherished friendly human sympathies. When death knocked at
any door of the hamlet, there was an echo from every fireside; and a
wedding dropped an orange blossom at every door. There was not a
grave in the little churchyard but had its story; not a crag or glen or
aged tree without its legend. The occupations of the people, who
were mostly small farmers and shepherds, were such as fostered
independence and originality of character. And where everybody
knew everybody, and everybody’s father had known everybody’s
father, and so on immemorially, the interest of man in man was not
likely to become a matter of cold hearsay and distant report. It was
here that Wordsworth learned not only to love the simplicity of
nature, but likewise that homely and earnest manliness which gives
such depth and sincerity to his poems. Travel, intercourse with
society, scholarly culture, nothing could cover up or obliterate those
early impressions. They widened with the range of his knowledge
and added to his power of expression, but they never blunted that
fine instinct in him which enables him always to speak directly to
men and to gentleman, or scholar, or citizen. It was this that enabled
his poetry afterwards to conquer all the reviews of England. The
great art of being a man, the sublime mystery of being yourself, is
something to which one must be apprenticed early.

Mr. Lowell here gave an outline of Wordsworth’s personal history and

character.

As a man we fancy him just in the least degree uninteresting—if the

horrid word must come out—why, a little bit of a bore. One must
regard him as a prophet in order to have the right kind of feeling
toward him; and prophets are excellent for certain moods of mind,
but perhaps are creatures

Too bright and good

For human nature’s daily food.

I fancy from what I have heard from those who knew him that he had
a tremendous prose-power, and that, with his singing-robes off, he
was dry and stiff as a figure-head. He had a purity of mind
approaching almost to prudery, and a pupil of Dr. Arnold told me he
had heard him say once at dinner that he thought the first line of
Keats’s ode to a “Grecian Urn” indecorous. The boys considered him
rather slow. There was something rocky and unyielding in his mind;
something that, if we found it in a man we did not feel grateful to and
respect, we should call hard. Even his fancy sometimes is glittering
and stiff, like crystallizations in granite. But at other times how tender
and delicate and dewy from very contrast, like harebells growing in a
crag-cleft!

There seem to have been two distinct natures in him—Wordsworth

the poet, and Wordsworth the man who used to talk about
Wordsworth the poet. One played a kind of Baruch to the other’s
Jeremiah, and thought a great deal of his master the prophet.
Baruch was terrifically uninspired, and was in the habit of repeating
Jeremiah’s poems at rather more length than was desired, selecting
commonly the parts which pleased him, Baruch, the best. Baruch
Wordsworth used to praise Jeremiah Wordsworth, and used to tell
entertaining anecdotes of him,—how he one day saw an old woman
and the next did not, and so came home and dictated some verses
on this remarkable phenomenon; and how another day he saw a
cow.

But in reading Wordsworth we must skip all the Baruch

interpolations, and cleave wholly to Jeremiah, who is truly inspired
and noble—more so than any modern. We are too near him,
perhaps, to be able wholly to separate the personal from the
poetical. I acknowledge that I reverence the noble old man both for
his grand life and his poems, that are worthy expressions of it. But a
lecturer is under bonds to speak what he believes to be the truth.
While I think that Wordsworth’s poetry is a thing by itself, both in its
heights and depths, something sacred and apart, I cannot but
acknowledge that his prosing is sometimes a gift as peculiar to
himself. Like old Ben Jonson, he apparently wished that a great deal
of what he wrote should be called “works.” Especially is this true of
his larger poems, like the “Excursion” and the “Prelude.” However
small, however commonplace the thought, the ponderous machine
of his verse runs on like a railway train that must start at a certain
hour though the only passenger be the boy that cries lozenges. He
seems to have thought that inspiration was something that could be
turned on like steam. Walter Savage Landor told me that he once
said to Wordsworth: “Mr. Wordsworth, a man may mix as much
poetry with prose as he likes, and it will make it the better; but the
moment he mixes a bit of prose with his poetry, it precipitates the
whole.” Wordsworth, he added, never forgave him.

There was a great deal in Wordsworth’s character that reminds us of

Milton; the same self-reliance, the same purity and loftiness of
purpose, and, I suspect, the same personal dryness of temperament
and seclusion of self. He seems to have had a profounder
imagination than Milton, but infinitely less music, less poetical faculty.
I am not entirely satisfied of the truth of the modern philosophy
which, if a man knocks another on the head, transfers all the guilt to
some peccant bump on his own occiput or sinciput; but if we
measure Wordsworth in this way, I feel as if he had plenty of
forehead, but that he wanted hind-head, and would have been more
entirely satisfactory if he had had one of the philo-something-or-
other.

It cannot be denied that in Wordsworth the very highest powers of

the poetical mind were associated with a certain tendency to the
diffuse and commonplace. It is in the Understanding (always prosaic)
that the great golden veins of his imagination are embedded. He
wrote too much to write always well; for it is not a great Xerxes army
of words, but a compact Greek ten thousand that march safely down
to posterity. He sets tasks to the divine faculty, which is much the
same as trying to make Jove’s eagle do the service of a clucking
hen. Throughout the “Prelude” and the “Excursion,” he seems
striving to bind the wizard imagination with the sand-ropes of dry
disquisition, and to have forgotten the potent spell-word which would
make the particulars adhere. There is an arenaceous quality in the
style which makes progress wearisome; yet with what splendors of
mountain-sunsets are we not rewarded! What golden rounds of
verse do we not see stretching heavenward, with angels ascending
and descending! What haunting melodies hover around us, deep
and eternal, like the undying barytone of the sea! And if we are
compelled to fare through sands and desert wilderness, how often
do we not hear airy shapes that syllable our names with a startling
personal appeal to our highest consciousness and our noblest
aspiration, such as we might wait for in vain in any other poet.

Take from Wordsworth all which an honest criticism cannot but allow,
and what is left will show how truly great he was. He had no humor,
no dramatic power, and his temperament was of that dry and
juiceless quality that in all his published correspondence you shall
not find a letter, but only essays. If we consider carefully where he
was most successful, we shall find that it was not so much in
description of natural scenery, or delineation of character, as in vivid
expression of the effect produced by external objects and events
upon his own mind. His finest passages are always monologues. He
had a fondness for particulars, and there are parts of his poems
which remind us of local histories in the undue importance given to
trivial matter. He was the historian of Wordsworthshire. This power of
particularization (for it is as truly a power as generalization) is what
gives such vigor and greatness to single lines and sentiments of
Wordsworth, and to poems developing a single thought or word. It
was this that made him so fond of the sonnet. His mind had not that
reach and elemental movement of Milton’s which, like the trade-
winds, gathered to itself thoughts and images like stately fleets from
every quarter; some, deep with silks and spicery, come brooding
over the silent thunders of their battailous armaments, but all swept
forward in their destined track, over the long billows of his verse,
every inch of canvas strained by the unifying breath of their common
epic impulse. It was an organ that Milton mastered, mighty in
compass, capable equally of the trumpet’s ardors, or the slim
delicacy of the flute; and sometimes it bursts forth in great crashes
through his prose, as if he touched it for solace in the intervals of his
toil. If Wordsworth sometimes puts the trumpet to his lips, yet he lays
it aside soon and willingly for his appropriate instrument, the pastoral
reed. And it is not one that grew by any vulgar stream, but that which
Apollo breathed through tending the flocks of Admetus, that which
Pan endowed with every melody of the visible universe, the same in
which the soul of the despairing nymph took refuge and gifted with
her dual nature, so that ever and anon, amid notes of human joy and
sorrow, there comes suddenly a deeper and almost awful tone,
thrilling us into dim consciousness of a forgotten divinity.

Of no other poet, except Shakspeare, have so many phrases

become household words as of Wordsworth. If Pope has made
current more epigrams of worldly wisdom, to Wordsworth belongs
the nobler praise of having defined for us, and given us for a daily
possession, those faint and vague suggestions of other-worldliness
of whose gentler ministry with our baser nature the hurry and bustle
of life scarcely ever allowed us to be conscious. He has won for
himself a secure immortality by a depth of intuition which makes only
the best minds at their best hours worthy, or indeed capable, of his
companionship, and by a homely sincerity of human sympathy which
reaches the humblest heart. Our language owes him gratitude for the