Understanding Biology 4th Edition

Kenneth Mason
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 Understanding
Biology
Fourth Edition

Kenneth A. Mason
University of Iowa

Tod Duncan

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

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 UNDERSTANDING BIOLOGY

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

This book is printed on acid-free paper.

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

ISBN 978-1-266-10043-7
MHID 1-266-10043-1

Cover Image: Anna Veselova/Shutterstock

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

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

mheducation.com/highered

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 Brief Contents
About the Authors iv Part V The Diversity of Life 474
Changes to This Edition vi 22 Systematics and Phylogeny 474
23 Prokaryotes and Viruses 494
Acknowledgments viii
24 Protists 520
A Learning Path to Understanding Biology ix 25 Fungi 547
Contents xv 26 Plants 568
27 Animal Diversity 593
Part I The Molecular Basis of Life 1 28 Vertebrates 625

1 The Science of Biology 1

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

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

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

Part IV Evolution 404

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

iii

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

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

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

iv

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

about the authors v

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 Changes to This Edition
A new pedagogical feature called “Concept Overview” was
added to all chapters in this fourth edition of Understanding
Biology. The chapter opening page contains a diagram laying out
the main concepts in the chapter. In the online eBook in Connect,
Progressive Concept Overviews are inserted in relevant places
in the chapter and provide a visual presentation of concept
statements that support the main concepts. These Progressive
Concept Overviews are available in the Instructor Resources
for students who are using a printed text. All of the Progressive
Concept Overviews are combined in an end-of-chapter diagram
that presents a conceptual hierarchy of the chapter, highlighting
the important concepts.
Also, McGraw Hill is dedicated to creating products that foster
a culture of belonging and are accessible to all the diverse global
customers we serve. Within this edition, content has been reviewed
to implement inclusive content guidelines around topics including
generalizations and stereotypes, gender, abilities/disabilities, race/
ethnicity, sexual orientation, diversity of names, and age. Similarly,
the illustrations throughout were evaluated and revised to ensure
meaningful text and images are distinguishable and perceivable by
users with limited color vision and moderately low vision.
Part I The Molecular Basis of Life
Chapter 1 A number of edits were made for clarity. The section
on the process of science was edited for clarity and
content, including the social context of science.
Chapter 2 Some minor edits were made for clarity.
Chapter 3 Edits were made to clarify the structure of nucleic
acids, and the process of protein folding. One figure
was converted into two figures to better fit the text
and to increase student understanding.
Part II Biology of the Cell
Chapter 4 The section on prokaryotic cell structure was
rewritten to incorporate new information on
organization, substructure, and compartments.
This provides a different view of the complexity
of prokaryotic cells. A new figure was added
comparing bacterial flagella and the archaeal
archaellum.
Chapter 5 A variety of edits were made for clarity and student
comprehension.
Chapter 6 The material on the Second Law of Thermodynamics
was rewritten for clarity and accessibility. This includes
one new figure to clarify the exergonic and endergonic
reactions. The nature of enzymes and how they
function was also rewritten for clarity.
Chapter 7 Light edits were made for clarity and student
understanding.
Chapter 8 The section on the light-independent reactions
was rewritten for clarity and accuracy. This
included edits to one figure for accuracy and
understanding. This also makes this chapter more
vi
mas39023_FM_i-xxiv.indd 6
congruent with chapter 7 in terms of names of
compounds that have multiple names.
Chapter 9 The chapter was edited for clarity, including
content on the nature of steroid hormones, and
how their structure affects their function.
Chapter 10 The section on eukaryotic chromosome structure
was further updated to reflect current thinking on the
organization of chromatin in the nucleus. This update
includes a new figure on chromosome organization.
Part III Genetics and Molecular Biology
Chapter 11 Minor edits for clarity, including in several figures,
should improve student understanding. The
material on segregation without crossing over was
deleted as outside the scope of a majors course.
Chapter 12 The historical material in the beginning of the
chapter was streamlined, although not removed. The
material on extensions to Mendel was reorganized
and rewritten for both clarity and updated content.
The material on quantitative traits was updated to
provide a more modern view. A new figure was
added to replace an old one on epistasis with a more
approachable example for students.
Chapter 13 The chapter was reorganized to both provide a more
logical flow of topics, and integrate new material that
provides a more modern view of human genetics.
Chapter 14 The presentation of experiments underlying DNA
structure was edited for clarity. The descriptions
of DNA structure were edited to emphasize
structure–function relationships.
Chapter 15 The chapter was edited for clarity and some content
was updated. The discussion of the genetic code
was rewritten for clarity, and information on the
genome, transcriptome, and proteome was edited
for currency.
Chapter 16 The introduction on transcription factors was edited
for clarity and content. The influence of chromatin
structure on gene regulation was updated. Material
on the ubiquitin/proteasome regulation of protein
turnover was added, including one new figure.
Chapter 17 The section on genetic tools and modern medicine
was completely rewritten. New material on
molecular tests for infectious diseases, including
COVID-19, was added. This also clarifies the use of
different kinds of tests.
Chapter 18 The chapter was edited extensively to update
content on a very fast-moving field. The human
genome section was rewritten to update all
content. Discussion was added of reference
genomes, and current knowledge of the human
genome. The wheat genome project was also
rewritten to include new material on sequencing
complex genomes. The content on what makes us
human was also updated.
04/10/22 12:47 PM
 Part IV Evolution
Chapter 19 The section on genetic variation in populations
was revised reflecting new information based
on wide­spread genomic investigation. Genomic
variation in humans is now discussed in detail,
quantifying the extent of variation that exists
and how that variation is apportioned within and
between populations.
Part V The Diversity of Life
Chapter 22 Information on roots of the eukaryotic tree was
revised to reflect changes in the classification of
the protists.
Chapter 23 The chapter was edited to reflect new information
on the organization of prokaryotic cells, including
compartmentalization. The material on prokaryotic
genetics was compressed, and the figures were
edited. New material on SARS-CoV-2 was added,
including a figure showing the virus life cycle.
Chapter 24 Many changes were made to the chapter to
improve clarity and readability.
Chapter 25 The chapter was edited for clarity and readability.
Chapter 26 The chapter was edited for clarity and readability.
Chapter 27 The discussion of relationships at the base of the
phylogeny for all animals was revised to reflect
new understand­ing and debate about relationships
among sponges, ctenophores, and other animals.
Additional changes were made to reflect other
changes in understanding of phylogenetic
relationships among animal taxa, such as the
position of chaetognaths and relationships among
protostome taxa. Aspects of taxonomy and natural
history were updated in line with new findings.
Chapter 28 The discussion of human evolution was revised in
light of new discoveries.
Part VI Plant Form and Function
Chapter 29 Minor edits were made throughout for readability,
clarity, and accuracy.
Chapter 30 The chapter was edited for clarity and readability.
Chapter 31 The chapter was edited throughout for clarity,
accuracy, and readability.
mas39023_FM_i-xxiv.indd 7
Part VII Animal Form and Function
Chapter 32 The chapter was edited for clarity. The material on
bone structure and development was rewritten to
improve student understanding.
Chapter 33 The description of sensory systems, including
receptor types, was updated to make it more
consistent and to improve student understanding.
The material on the retina was edited extensively
for clarity, and one figure was added.
Chapter 34 The section on lung function was edited for clarity.
Information was added on how lung function is
assessed. The cardiac cycle was edited for clarity
and student understanding.
Chapter 35 The material on hormones was edited to provide
a better focus on structure–function relationships.
The material on fluid balance was edited to
emphasize homeostatic mechanisms. The section
on adaptive immunity was rewritten to include
more history of vaccination and a more worldwide
perspective. Material on vaccination and how
the immune system responds was added. This
includes a discussion of vaccination and the
COVID pandemic.
Chapter 36 The material on nuclear reprogramming was
edited to provide a better historical perspective.
The material on cloning in mammals was edited to
include more detail. The material on contraception
was edited for clarity.
Part VIII Ecology and Behavior
Chapter 37 Several Review of Concept questions were
replaced to better assess student understanding.
Chapter 38 Human population trends and other timely data
were updated to stay current. Also, a new section
(38.7) was added on pandemics and human health
that covers the general topic, but extensively
details the population biology of the COVID-19
pandemic.
Chapter 40 Data on biosphere impacts of humans were
updated to stay current.
changes to this edition vii
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 Acknowledgments
A revision of this scope relies on the talents and efforts
of many people working behind the scenes, and we have
benefited greatly from their assistance.
The copy editor, Beth Bulger, labored many hours and always
improved the clarity and consistency of the text. She has made
a tremendous contribution to the quality of the final product.
We were fortunate to work with MPS Limited to update the
art program and improve the layout of the pages. Our close
collaboration resulted in a text that is pedagogically effective as
well as more beautiful than any other biology text on the market.
We have the continued support of an excellent team at
McGraw Hill. Lora Neyens, the Portfolio Manager for Biology,
has been a steady leader during a time of change. Senior
Product Developer, Liz Sievers, provided support in so many
ways, it would be impossible to name them all. Kelly Hart, Lead
Content Project Manager, and David Hash, Designer, ensured
our text was elegantly designed. Kelly Brown, Senior Marketing
Manager, is always a sounding board for more than just
marketing, and many more people behind the scenes have all
contributed to the success of our text. This includes the digital
team, to whom we owe a great deal for their efforts to continue
improving our Connect assessment tools.
Throughout this edition we have had the support of spouses
who have seen less of us than they might have liked because
of the pressures of getting this revision completed. They
have adapted to the many hours this book draws us away
from them, and, even more than us, looked forward to its
completion.
In the end, the people we owe the most are the generations
of students who have passed through our lecture halls. They
have taught us at least as much as we have taught them, and
viii
mas39023_FM_i-xxiv.indd 8
their questions and suggestions continue to improve the text
and supplementary materials.
Finally, we need to thank instructors from across the country
who are continually sharing their knowledge and experience
with us through market feedback, reviews, and symposia. Their
feedback shaped this edition. All of these people took time to
share their ideas and opinions to help us build a better edition of
Understanding Biology for the next generation of introductory
biology students, and they have our heartfelt thanks.
Reviewers for Understanding Biology, 4th edition
Christopher Allen Lone Star College, University Park
Lauri Carey Illinois Valley Community College
Christopher Chamberlain St. Cloud Technical
& Community College
Mary Colon Seminole State College
Carolyn Danna Stevenson University
Jacqueline Dartley Bergen Community College
Sandra Fox-Moon Anne Arundel Community College
Robin Graham Dallas College
Linda Johnson University of Maryland Eastern Shore
Olga Ruiz Kopp Utah Valley University
Kimberly Maznicki Seminole State College
Rosa Moscarella University of Massachusetts
Terina Nusinov Seminole State College
Helene Peters Brewton-Parker College
Debra Rinne Seminole State College
Sherry Stewart Navarro College
viii
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 A Learning Path to Understanding Biology
Understanding Biology and its online assets have been carefully thought out and crafted to help students and professors work efficiently
and effectively through the material in the course, making the most of study time and furthering instructional goals.

The Learning Path

glycolysis needs to be transported into the mitochondria by active

The Total Energy Yield of
7.5 transport, which costs one ATP per NADH transported. This
reduces the predicted yield for eukaryotes to 30 ATP.
Aerobic Respiration Far Exceeds
C o n c e p t O vThat
e rv of
i eGlycolysis
w

7
Calculation of P/O Ratios Has
How Cells Harvest Energy
Changed
This Concept Overview diagrams the key concepts that were discussed in this chapter. over Time
Ever since the link was discovered between electron transport
and the proton gradient used by ATP synthase, biochemists have LEARNING OBJECTIVE 7.5.2 Explain how our understanding
Life is possible due to energy transformations
attempted to determine the number of ATPs produced per NADH of the P/O ratio has changed over time.
Le arning Path feeding electrons into electron transport. This number has proved
7.1 Cells Harvest Energy from 7.5 The Total Energy Yield of to be surprisingly elusive. Early estimates were based on errone- The value for the amount of ATP synthesized per O2 molecule
Organic Compounds by Aerobic Respiration Far
ous assumptions, but we now have both theoretical and calculated reduced is called the phosphate-to-oxygen ratio (P/O ratio).
Oxidation Exceeds That of Glycolysis
Biological values
energythat
can are
takeinvarious
agreement. Both theoretical calculations and direct measurement of this
7.2 Glycolysis Splits Glucose and 7.6 Aerobic Respiration Is Cells store and use energy as ATP Enzymes are specific biological catalysts
forms value have been contentious issues. When theoretical calcula-
Yields a Small Amount of ATP Regulated by Feedback
Inhibition tions were first made, we lacked detailed knowledge of the
7.3 The Citric Acid Cycle Is the The Theoretical Yield for Eukaryotes Is respiratory chain and the mechanism for coupling electron
Oxidative Core of Cellular 7.7 Some Forms of Energy
Respiration Metabolism Do Not Require O2
Laws of
30 Molecules of ATP perATP
Free energy in a
Glucose
is continually made and used transport to ATP synthesis. Because redox reactions occur
Even exergonic Metabolism includes
in a cycle at three sitesreactions
for NADH and two sites for FADH 2 , it was
7.4 Electrons Removed by 7.8 Carbohydrates Are Not the thermodynamicsMolecule system can do have all reactions
2 ATP Glycolysis Figure 7.14 Overview
Oxidation Pass Along an
Electron Transport Chain
Heterotrophs
of
Only Energy Source Used by
determine how work assumed that three energy
activation molecules of ATP were produced per
energy changesLEARNING OBJECTIVE 7.5.1 Calculate the number of ATP NADH and two per FADH2. We now know that assumption
Glucose NADH aerobic respiration in the Energy from
produced by a cell via aerobic respiration. ATP synthesiswas overly simplistic. Anabolic reactions
Catalysts speed
mitochondria. The entire Free energy ATP hydrolysis requires energy Understanding that a proton gradientbuild
reactions by lowering
is themolecules
link between
The number of molecules
changes determineof ATP drives endergonic
produced by ATP synthasefromper
exergonicelectron transport andenergy
ATP synthesis changed the nature of the
Pyruvate ErikAgar/Getty Images
reactions reactions activation
process of aerobic respiration is Energy is notmolecule of theglucose
directiondepends
of on the number of protons trans- calculations. We need to know the number of protons
Catabolic pumped
reactions
created orported acrossreactions
the inner membrane and the number of protons during electron transport: 10 H+ per NADH, 6 H+ per
degradeand molecules
shown in cellular context. destroyed but

Pyruvate
C onc e pt Ove rvie w
Glycolysis occurs in the
This Concept Overview provides a diagram of the overarching concepts that are covered in this chapter. These concepts will be expanded upon
in the Concept Overview at the end of the chapter. form
▲
can change
needed per ATP synthesized. The ATP
per NADH and FADH2 is 10 and 6immediate
synthesized requires 4 H+, leading
number

to 10/4
of protons transported
provides
H+, respectively. Each ATP
= 2.5 ATP/NADH,
FADH2. Then we need to know the number of protons needed per
ATP.
Enzymes bind

depends
Because ATP
to specific
Enzymes have
synthase is a
optimal pH and
on the numbertemperatures
rotary motor,
Enzymes
of binding sites for ATP
this calculation
work
and the number
energy for cells substrates
NADH cytoplasm with the pyruvate and
Every concept is broken down into sections that
together in
Oxidation and 6/4 = 1.5Exergonic
ATP/FADH2. of protons required for rotation. We know thatbiochemical
ATP synthase has
CO2 Organisms convert chemical energy to ATP through cellular respiration
reactions release
Oxidizing glucose to pyruvate via glycolysis yields 2 ATP three binding sites for ATP. If 12 protons are used per rotation,
Acetyl-CoA
NADH produced entering the No energy free energy and
directly, and 2 × 2.5 = 5 ATPHydrolysis
from NADH. of twoThe oxidation of
ATP powers
Enzymes
you getconform
the value
Activators
+
of 4 Hinhibitors
per ATP
and pathways
canused in the previous calcula-
cover skills or ideas students should master. Learning
transaction is are spontaneous to
Cellular respiration
extracts energy from
mitochondria. Here, pyruvate is
Glycolysis splits glucose
The citric acid cycle
oxidizes 2-carbon
Oxidative phosphorylation
uses chemiosmosis to
Energy metabolism
varies across
pyruvate to acetyl-CoA yields terminal
100% efficient anotherphosphates
2 × 2.5 = 5 ATP biosynthesis,
from tion. Actual measurements
the shape of affectofenzyme
the P/O ratio have been problem-
and yields a little ATP
NADH. Finally, the citric acid cycle in ATPproduces
releases 2 ATP movement,
directly, and atic,their
but they be at most 2.5. Pathways can be
now appear toactivity
organic molecules units make ATP living systems substrates
oxidized and fed into the citric energy transport
objectives at the beginning of each section help identify
6 × 2.5 = 15Endergonic
ATP from NADH, and 2 × 1.5 = 3 ATP from FADH regulated by
2. We can also calculate how efficiently respiration captures
CO2 NADH acid cycle to complete the The entropy Summing
of reactions require
all of these leads to 32 ATP for respiration (figure 7.16). the free released by the oxidation offeedback
energy bind
Substrates glucose in inhibition
the form of
H+ Citric Introduc tion the universe is Thisenergy
number and are
is accurate for bacteria, but it does not hold ATP.enzyme
The amount
activeof free energy released by the oxidation of glu-
important concepts.
not spontaneous
Acid oxidation process. All the
Life is driven by energy. All the activities carried out by organisms use energy—the swimming of bacteria, the purring of a cat, and even the thoughts
you are forming to process these words. In this chapter, we discuss the processes all cells use to obtain chemical energy from organic molecules,
increasingfor eukaryotes, because the NADH produced in the cytoplasm by cose is sites
686 with
kcal/mol, and the free energy stored in each ATP is
28 ATP e−is used to synthesize ATP. Then, in chapter 8, we will examine
which photosynthesis, which uses light energy to make chemical energy. We consider
energetic the plant,electrons
a photosynthesizer, andharvested
induced fit
Cycle FADH2
the conversion of chemical energy to ATP first because all organisms—including the caterpillar feeding on the
2 ATP plant, pictured in the photo—are capable of harvesting energy from chemical bonds. Obtaining energy via respiration is an ancient and universal Figure 7.16 Theoretical Glucose
process. by oxidations in the overall ATP yield. The theoretical
2 ATP
e− 2 ATP
2H+ process are transferred by yield of ATP harvested from Glycolysis
+ H 2O
glucose by aerobic respiration
c. An enzyme does not change as a result of the reaction.Pyruvate c. the sum of ΔG for 2 ATP hydrolysis and5ΔG for ATPthe reaction
1/ O NADH and FADH2 to the totals 32 molecules. In
NADH
2 2
e− electron transport chain. The Assessing the Learning Path
d. An enzyme works in both the forward and reverse
eukaryotes this is reduced to
directions of a reaction.
30 because it takes 1 ATP to
is positive.
d. ATP hydrolysis raises the activation energy for the Chemiosmosis
8. Which statement about the influence of temperature on Pyruvate oxidation reaction. 2 NADH 5 ATP
electron transport chain uses transport each molecule of
enzymes is NOT true? 6. The enzyme aromatase is found in the cytosol of some cells
NADH that is generated by
the energy released during a. All enzymes have the same intrinsic optimal temperature.
Understand glycolysis in the cytoplasm into
and converts testosterone to estrogen. You decide to test
4. An endergonic reaction has which of the following properties?

Q ▲ electron transport to pump

b. Raising the temperature may increase the activity of an
1. A covalent
c. Some
the mitochondria.
enzyme. bond between a hydrogen atom and an oxygen
enzymeswhat are stable at energy?
the boiling point of water.
aromatase from a particular cell type, and2your ATP
admits +ΔG
a. that heand
+ΔG of
tubes.b.Which and
the reaction
drastically
thethe
increased
reaction
following
is spontaneous.
is aislikely
not spontaneous.
result?
lab partner
the pH in all the test
protons across the inner atom represents kind of −ΔG and
c. enzyme the reaction is and
spontaneous.
C
Every chapter opens with a Learning Path that walks students d. Raising the temperature may decrease the activity of Citric Acid
an a. The will be denatured the substrate
a. Kinetic energy 6 NADH 15 ATP will not
Cycle d. −ΔG and the reaction is not spontaneous.
H+ membrane. This creates an b. enzyme.
Potential energy bind to its active site.
H+ 9. Coenzymes
c. Mechanical energy
b.5. The
Theenzyme
molecule willATP
convert testosterone
is less stable than to estrogen
ADP + at Chemiosmosis
Pi abecause
H+ through the main concepts in the chapter to enable their
electrochemical gradient that a. can be metal ions.
d. Solar energy
b. can bind in active sites and participate directly in a
faster
c. The
a. therate.
negatively
b. mistake
2 FADH
will have
the positively
charged
charged
2
no effect,
phosphates 3 repel
as enzymes
phosphates
ATP each other.
repel areeach
not other.
contains potential energy. The 2. During a redox
catalytic reaction, the molecule that gains an electron is
reaction. sensitive
c. ATPtoischanges in pH.than ADP and P .
understanding of where the material fits in the context of
enzyme ATP synthase uses this a. are
c. reduced
b. All
d. oxidized
and now
sometimes
of the and
above
has a higher energy level.
vitamins.
now has a lower energy level.
d. The d. ΔG
the will
much larger
be lowered
adenine
spontaneously.
inTotal
ATP and
net isATPtheyield
(30 in eukaryotes)
reaction
charged.
i
= 32 will proceed

gradient to phosphorylate ADP 6. Whatthe is activation energy?pathway below. It is regulated

other concepts in the chapter. A new Concept Overview
c. competitive
10. In reduced and now has a lower energy level.
inhibition, 7. Consider enzyme-catalyzed
d. oxidized
a. two 146
enzymes Part
nowIIhas
andcompete Biology of the
a higher
with each Cell level.
energy
other for a substrate. a. The inhibition
by feedback thermal energy of enzymeassociated
1. Which with random
of the movements of
following
to form ATP. molecules
3. b. an inhibitor
When a bear eats molecule bindssome
a salmon, to anofallosteric
the energysitestored
on an in the statements is NOT accurate regarding this pathway?
provides a graphical presentation of the Learning Path, enzyme,
salmon
c. the
of an inhibitor
causing
is used a change
by the
molecule in
energy originally binds
in its
bear for theactivities
the to
active site.
the active
salmon
and growth. Much
site of an as heat.
is dissipated
b. The energy released through breaking chemical bonds
Enzyme Enzyme Enzyme Enzyme
c. The difference in free energy between reactants and Enzyme
enzyme, so the substrate cannot bind. products
1 2 3 4 5

highlighting the main concepts in the chapter. This

d.
a. the
is an example
the products
11. Anabolism
conversion
of
of aofreaction
kineticboth
energy
is Law of Thermodynamics.
compete for the energy.
to potential active site. d. TheA energy B required C to initiate D a chemical
E reaction
F
b. the Second 7. Which of the following is NOT a property of an enzyme?
a. Compound F binds to the active site of enzyme 1.
a. the gain of a proton.
c. 100% efficient energy conversion. a. An enzyme reduces the activation energy of F. a reaction.
b. metabolism in animals. b. Enzyme 1 contains an allosteric site for compound
d. a conversion of potential energy to kinetic energy. c. Whenb. Ancompound
enzyme lowersF is made the in free
highenergy
enough of levels,
the reactants.
it shuts
c. the buildup of molecules.
off its own synthesis.
H+ This complex can be dissociated into two subportions: the Chapter 6 Energy and Metabolism 129
F0 membrane-bound complex, and the F1 complex composed of Apply
ATP the stalk and a knob, or head domain. 1. When a hibernating animal uses its stored fat to power basic Synthesize
body functions (for example, breathing), it is
1. Some people argue that evolution, which is generally
The F1 complex has enzymatic activity. The F0 complex con- a. converting kinetic energy to potential energy.
associated with progressive increases in the complexity
b. converting kinetic energy to chemical energy.
tains a channel through which protons move across the membrane c. converting potential energy to kinetic energy.
(order) of organisms, cannot occur because entropy (disorder)
ADP + Pi is increasing in the universe. Is this argument valid? Explain.
down their concentration gradient. As they do so, their movement d. converting chemical energy to potential energy.
2. On summer nights in many parts of the country, one can
2. During certain stages of cellular respiration, electrons are
causes part of the F0 complex and the stalk to rotate relative to the transferred from glucose molecules to a molecule called
often see fireflies glowing briefly in the dark. Do you suppose
producing this light requires energy? If so, where might the
nicotinamide adenine dinucleotide (NAD+). During this reaction,
knob. The mechanical energy of this rotation is used to change the a. glucose is oxidized and NAD+ is reduced.
energy come from? How would you test your hypothesis?
3. Examine the graph shown here. Describe what happens
conformation of the catalytic domain in the F1 complex. b. glucose is reduced and NAD+ is oxidized.
to this human enzyme protein’s structure when the body’s
c. both glucose and NAD+ have gained protons.
Thus, the synthesis of ATP is achieved by a tiny rotary motor, d. glucose has gained protons and NAD+ has lost protons. temperature is raised above 40°C.
Stalk
the rotation of which is driven directly by a gradient of protons. 3. Sodium ions (Na+) can move through channel proteins
across some biological membranes. If Na+ is present in a Optimum temperature
Rate of reaction

The flow of protons is like that of water in a hydroelectric power higher concentration on one side of a membrane, the ions for human enzyme
will tend to move across the membrane until they are equally
plant. As the flow of water driven by gravity causes a turbine to distributed on both sides of the membrane. This process
rotate and generate electric current, the proton gradient produces a. results in a gain of potential energy for the cell.
b. results in a decrease in entropy.
the energy that drives the rotation of the ATP synthase generator. c. follows the Second Law of Thermodynamics. 30 40 50
d. All of the above Temperature of reaction
4. If the products of a chemical reaction have higher free energy
4. Phosphofructokinase functions to add a phosphate group
REVIEW OF CONCEPT 7.4 than the reactants, this reaction
a. will not proceed spontaneously. to fructose-6-phosphate. This enzyme functions early in
b. will proceed spontaneously. glycolysis (refer to chapter 7). The enzyme’s active site binds
H+ H+
H+ H+ The electron transport chain receives electrons from NADH c. must have increased the total energy in the universe. to fructose and ATP. An allosteric inhibitory site can also bind
d. must have decreased the total energy in the universe. ATP when ATP levels are high.
and FADH2 and passes them down the chain to oxygen, using a. Predict the rate of the reaction if with low ATP levels.
H+ 5. ATP can be used to drive an endergonic reaction if
b. Predict the rate of the reaction if with high ATP levels.
H+ the energy from electron transfer to pump protons across the a. ATP hydrolysis lowers the activation energy for the reaction.
b. the sum of ΔG for ATP hydrolysis and ΔG for the reaction c. Describe what is happening to the enzyme with high levels
membrane, creating an electrochemical gradient. The enzyme is negative. of ATP.

he ATP rotary engine. Protons move across ATP synthase uses this gradient to drive the endergonic reac-
own their concentration gradient. The energy
the rotor and stalk structures to rotate. This
tion of phosphorylating ADP to ATP. ▲
■ How would poking a small hole in the outer membrane
gy alters the conformation of the ATP synthase
affect ATP synthesis?
Chapter concepts are assessed at three different levels at the
yze the formation of ATP.
end of the chapter. On a first pass through the chapter prior to
Chapter 7 How Cells Harvest Energy 145
▲ class, students might focus on the “Understand” level. As they
At the end of each section, Review of Concept questions gain greater mastery of the material, they should challenge
allow students to check their understanding before themselves with “Apply” and “Synthesize” questions that
moving on to the next concept. require higher cognitive skills.

ix

mas39023_FM_i-xxiv.indd 9 04/10/22 12:47 PM

 Think Like a Scientist

Scientific Thinking figures throughout Inquiry & Analysis

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

Inquiry & Analysis

pH Sensitivity of the Glu-GABA Antiporter

a scientific experiment, laying out Resists the Acid Environment

of the Stomach
300
GadC
AdiC
all chapters explore a
250

the Hypothesis, Predictions, Test scientific investigation in

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

Procedures, Results, and Conclusion. more detail, presenting

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

Some also challenge the students to experimental results and

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

devise further experiments. challenging students to

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

interpret the data.

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

environments, investigators compared its activity at a vari-

electrons in the porphyrin ring, which are b. Compare the amount of substrate accumulated
Light

y through the alternating carbon single- and
Different small side groups attached to the
ter the absorption properties of the molecule
ds of chlorophyll (figure 8.6). The precise
m is also influenced by the local microenvi-
the association of chlorophyll with different
ctrum of photosynthesis—that is, the relative
ferent wavelengths of light in promoting
esponds to the absorption spectrum for chlo-
amously demonstrated in the classic experi-
gure 8.7. All plants, algae, and cyanobacteria
their primary pigments. This action spec-
ened with the help of accessory pigments
s section.
Filament of green algae ety of pHs with that of a different amino acid antiporter
called AdiC, which transports arginine out of cells under a by AdiC in 10 minutes at pH 9.0 with that
accumulated at pH 5.0. What fraction of the
broad range of conditions. The results of monitoring trans-
low pH activity is observed at the higher pH?
port for 10 minutes are presented in the graph.
low c. In a similar fashion, compare the amount of
substrate accumulated by GadC at pH 9.0
with that accumulated at pH 5.0. What
Result: The bacteria move to regions of high O2 , or regions of most fraction of the low pH activity is observed at
active photosynthesis. These are in the purple/blue and red regions
Outside Inside the higher pH?
cell cell
3. Making Inferences Would you say that the GadC
of the spectrum. antiporter exhibits the same pH dependence as the
Conclusion: All wavelengths are not equally effective at promoting AdiC antiporter? If not, which antiporter is less
GABA active at nonacid pHs?
photosynthesis. The most effective constitute the action spectrum
4. Drawing Conclusions Is the glutamate-GABA
for photosynthesis. antiporter GadC active at nonacid pHs?
Further Experiments: How does the action spectrum relate to the 5. Further Analysis The GadC antiporter also
Glutamate transports the amino acid glutamine (Gln). Do you
various absorption spectra in figure 8.5?
think this activity has any role to play in combating
low pH environments? How would you test this
hypothesis?
Figure 8.7 Determination of an action spectrum for
photosynthesis. Chapter 5 Membranes 111

Chlorophyll a: R = CH3 It is reasonable to ask why these photosynthetic organisms

H R
Chlorophyll b: R = CHO do not use a pigment like retinal (the pigment in our eyes), which
has a broad absorption spectrum that covers the range of 500 to
CH2CH3
600 nm. The most likely hypothesis involves photoefficiency.
N N
Although retinal absorbs a broad range of wavelengths, it does so
Mg H with relatively low efficiency. Chlorophyll, in contrast, absorbs in
N N
CH3
only two narrow bands but does so with high efficiency. For this
reason, plants and other photosynthetic organisms achieve far
Concept Overviews
H higher overall energy capture rates with chlorophyll than they

2
H
CO2CH3
O Each Concept Overview, a new feature in Understanding Biology, 4th edition, is a graphic representation of the Learning Path that
would with other pigments.

Carotenoids Are Accessory Pigments

guides students through the concepts in the chapter. Concept Overviews provide a conceptual framework of the chapter. When
2

LEARNING OBJECTIVE 8.3.3 Explain the role of accessory

concept statements are placed in the context of a flow diagram, students see relationships and connections between concepts.
2
Figure 8.6 Chlorophyll. pigments.
H3 Chlorophyll molecules consist of a
2
porphyrin head and a hydrocarbon Carotenoids are pigment molecules that, like chlorophyll, consist
2
2
tail that anchors the pigment of carbon rings linked to hydrocarbon chains, but in this case
CH3 molecule to hydrophobic regions chains with alternating single and double bonds. Carotenoids
2 of proteins embedded within the can absorb photons with a wide range of energies,
Flowering althoughstrategies
plant reproductive they ensure their success and diversity Flowering plant reproductive strategies ensure their success and diversity
2
2
thylakoid membrane. The only are not always highly efficient in transferring this energy. Carot-
CH3 difference between the two enoids assist in photosynthesis by capturing energy from light
2 chlorophyll molecules is the composed of wavelengths that are not efficiently absorbed by
Flowering is the first step Double fertilization is followed Plant growth starts with Self-pollination can be favored
2
substitution of a —CHO (aldehyde) chlorophylls
Flowering is(figure
the first8.8).
step Double fertilization is followed Plant growth starts with Self-pollination can be favored
in plant reproduction in stable environments
inCarotenoids
plant reproduction in stable environments by embryogenesis germination
2
group in chlorophyll b for a —CH3 byrole
also perform a valuable embryogenesis
in scavenging free germination
CH3
3 (methyl) group in chlorophyll a. radicals. The oxidation–reduction reactions that occur in the
y of the Cell Flowering leads to gamete Fertilization Seeds play roles Germination Life spans vary Asexual reproduction

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

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

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

chapter contains three to five broad concept statements regulated by

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

Progressive Concept
procambium Emerging cloned in the lab
▲

are embryo by wind, develops into a reproduce the

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

in the eBook, at the end and disperse

pollen

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

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

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

x a learning path to understanding biology

mas39023_FM_i-xxiv.indd 10 04/10/22 12:48 PM

 Strengthen Problem-Solving Skills with Connect®
Detailed Feedback in Connect®
Learning is a process of iterative development, of making
mistakes, reflecting, and adjusting over time. The question
bank and test bank in Connect® for Understanding Biology,
4th edition, are more than standard assessments; they are
self-contained learning experiences that systematically build
student learning over time.
For many students, choosing the right answer is not
necessarily based on applying content correctly; it is more
a matter of increasing their statistical odds of guessing. A
major fault with this approach is students don’t learn how to
process the information correctly, mostly because they are
repeating and reinforcing their mistakes rather than reflecting
and learning from them. To help students develop problem-
solving skills, all higher-level Bloom’s questions in Connect
are supported with hints, to help students focus on important
information for answering the questions, as well as detailed
feedback that walks students through the problem-solving
process, using Socratic questions in a decision-tree-style
mas39023_FM_i-xxiv.indd 11
framework to scaffold learning, where each step models and
reinforces the learning process.
The feedback for each higher-level Bloom’s question (Apply,
Analyze, Evaluate) follows a similar process: Clarify Question,
Gather Content, Choose Answer, Reflect on Process.
Unpacking the Concepts
We’ve taken problem solving a step further. In each chapter, two
higher-level Bloom’s questions in the question and test banks
are broken out by the steps of the detailed feedback. Rather than
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answer questions about earlier steps, such as “What is the key
concept addressed by the question?” before proceeding to
answer the question. A professor can choose which version of
the question to include in the assignment based on the problem-
solving skills of the students. The Unpacking the Concepts
questions are found under the Coursewide Content in Connect.
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 Data and Graphing Interactives and the opportunity to manipulate variables in the
interactive, analyze or evaluate data, or view different
To help students develop analytical skills, Connect® for
aspects of the problem. A series of questions follows the
Understanding Biology, 4th edition, is enhanced with Data
activity to assess whether the student understands and
and Graphing Interactives found under the Coursewide
is able to interpret the data and results.
Content. Students are presented with a scientific problem

Quantitative Reasoning Question Bank
Under the Coursewide Content in Connect, the Quantitative
Reasoning question bank provides more challenging
algorithmic questions, intended to help students practice their
quantitative reasoning skills. Hints and stepped-out solutions
walk students through the problem-solving process.

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are now often being asked to deliver some of their lab compo-
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These simulations help each student learn the practical
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McGraw Hill
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Connect’s SmartBook 2.0 provides an adaptive learning expe-
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Fundamentals of Student Success, help students prepare for
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under Adaptive Learning Assignment, and post-lab a ­ ssessment
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From the instructor’s perspective, these simulations may
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Seamlessly integrated within Connect, these services ­allow
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 Contents
About the Authors iv PART II Biology
Changes to This Edition vi of the Cell 65
Acknowledgments viii Dr. Gopal Murti/Science Source

A Learning Path to Understanding Biology ix 4 Cell Structure 65

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

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

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

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

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

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

19 Genes Within Populations 404

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

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

PART V The Diversity 25 Fungi 547

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

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

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

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

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

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

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 Part I The Molecular Basis of Life

1
The Science of Biology

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

Soames Summerhays/Natural Visions

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

Biology is the study of living things

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

In tr oduct ion
You are about to embark on a journey of discovery about the nature of life. Almost two centuries ago, a young English naturalist named Charles
Darwin set sail on a similar journey on board H.M.S. Beagle; the photo on this page shows a replica of this ship. Darwin’s observations during
the voyage of the Beagle influenced his development of the theory of evolution by natural selection, which has become the core of the science of
biology. Before we begin, however, let’s take a moment to think about what biology is and why it’s important.

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 1.1 The Diversity of Life
Is Overwhelming
Biology is the study of living things—literally the science of life.
The living world is teeming with a breathtaking variety of creatures—
whales, butterflies, mushrooms, plants, bacteria—which despite
their obvious differences share features common to all living
­organisms. We will begin with a brief history of how we classify this
diversity.

Domain Archaea. This Domain Bacteria. This

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

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 Protists. Once considered a
kingdom, this term is still used
informally to refer to unicellular
eukaryotes.
Fungi. This kingdom contains
mostly multicellular
nonphotosynthetic organisms
that digest their food
externally, such as mushrooms.
Plantae. This kingdom contains
photosynthetic multicellular
organisms that are terrestrial,
such as the flowering plant
pictured here.
Animalia. Organisms in this
kingdom are nonphotosynthetic
multicellular organisms that
digest their food internally,
such as this ram.

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

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

Chapter 1 The Science of Biology 3

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 Living Systems Show Hierarchical
Organization
LEARNING OBJECTIVE 1.2.2 Describe the hierarchical nature
of living systems.

Life’s organisms interact with each other at many levels, in

ways simple and complex. A key factor organizing these inter-
actions is their degree of complexity. The organization of the
biological world is hierarchical—that is, each level builds on
the level below it, from the very simplest level of individual

Figure 1.4 Energy utilization. This kingfisher obtains the
energy it needs to move, grow, and carry out its body processes
by eating fish. It harvests the energy from food using chemical
processes that occur within cells.
Armin Floreth/imagebroker/Alamy Stock Photo
what an individual organism will be like is contained in a code
dictated by the order of the subunits making up the DNA
molecule. Because DNA is passed from one generation to the
next, any change in a gene can be preserved and passed on to
future generations. The transmission of characteristics from
parent to offspring is a process called heredity. All organisms
interact with other organisms and the nonliving environment
in ways that influence their survival, and as a consequence,
organisms evolve adaptations to their environments.
Figure 1.5 Hierarchical organization of living systems.
Life is highly organized, from the simplest atoms to complex,
multicellular organisms. Along this hierarchy of structure, atoms
form molecules, which are used to form organelles, which in turn
form the functional subsystems within cells. Cells are organized
into tissues, then into organs and organ systems such as the
nervous system. This organization extends beyond individual
organisms to populations, communities, ecosystems, and finally
the entire biosphere.
(organelle): Keith R. Porter/Science Source; (cell): STEVE GSCHMEISSNER/
SCIENCE PHOTO LIBRARY/Alamy Stock Photo; (tissue): Ed Reschke/Getty Images;
(organism): Russell Illig/Getty Images; (population): George Ostertaga/
gefotostock/Alamy Stock Photo; (species top): USDA Natural Resources
Conservation Service; (species bottom): U.S. Department of Agriculture (USDA);
(community): Ryan McGinnis/Alamy Stock Photo; (ecosystem): Steven P. Lynch/
McGraw Hill; (biosphere): Goddard Space Flight Center/NASA

CELLULAR LEVEL

1 Atoms 2 Molecule 3 Macromolecule 4 Organelle 5 Cell 6 Tissue 7 Organ

H
N

H
N
C

O
0.2 µm 100 µm

4 Part I The Molecular Basis of Life

mas39023_ch01_001-020.indd 4 28/11/22 12:17 PM

Another random document with
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The Project Gutenberg eBook of Natural history
of intellect, and other papers
 This ebook is for the use of anyone anywhere in the United
States and most other parts of the world at no cost and with
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laws of the country where you are located before using this
eBook.

Title: Natural history of intellect, and other papers

Author: Ralph Waldo Emerson

Release date: September 4, 2023 [eBook #71558]

Language: English

Original publication: Boston: Houghton, Mifflin and Company,

1893

Credits: Emmanuel Ackerman, A. Marshall, and the Online

Distributed Proofreading Team at https://www.pgdp.net
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*** START OF THE PROJECT GUTENBERG EBOOK NATURAL

HISTORY OF INTELLECT, AND OTHER PAPERS ***
 TRANSCRIBER’S NOTE
Footnote anchors are denoted by [number], and the footnotes have been placed
at the end of the chapter.
Some minor changes to the text are noted at the end of the book.
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domain.
N AT U R A L H I S TORY OF
INTELLECT

A N D O T H E R PA P E R S

BY

RALPH WALDO EMERSON

BOSTON AND NEW YORK

HOUGHTON, MIFFLIN AND COMPANY
The Riverside Press, Cambridge
1893
 Copyright, 1893,
By EDWARD W. EMERSON.

All rights reserved.

The Riverside Press, Cambridge, Mass., U. S. A.

Electrotyped and Printed by H. O. Houghton & Company
 PREFATORY NOTE.

The first two pieces in this volume are lectures from the “University
Courses” on philosophy, given at Harvard College in 1870 and 1871,
by persons not members of the Faculty. “The Natural History of the
Intellect” was the subject which Emerson chose. He had from his
early youth cherished the project of a new method in metaphysics,
proceeding by observation of the mental facts, without attempting an
analysis and coördination of them, which must, from the nature of
the case, be premature. With this view, he had, at intervals from
1848 to 1866, announced courses on the “Natural History of
Intellect,” “The Natural Method of Mental Philosophy,” and
“Philosophy for the People.” He would, he said, give anecdotes of
the spirit, a calendar of mental moods, without any pretense of
system.
None of these attempts, however, disclosed any novelty of
method, or, indeed, after the opening statement of his intention, any
marked difference from his ordinary lectures. He had always been
writing anecdotes of the spirit, and those which he wrote under this
heading were used by him in subsequently published essays so
largely that I find very little left for present publication. The lecture
which gives its name to the volume was the first of the earliest
course, and it seems to me to include all that distinctly belongs to the
particular subject.
The lecture on “Memory” is from the same course; that on
“Boston” from the course on “Life and Literature,” in 1861. The other
pieces are reprints from the “North American Review” and the “Dial.”
J. E. Cabot.
September 9, 1893.
 CONTENTS.

PAGE
Natural History of Intellect 7
Memory 55
Boston 73
Michael Angelo 97
Milton 121
Papers from The Dial 147
I. Thoughts on Modern Literature 149
II. Walter Savage Landor 168
III. Prayers 177
IV. Agriculture of Massachusetts 183
V. Europe and European Books 187
VI. Past and Present 197
VII. A Letter 206
VIII. The Tragic 216
NATURAL HISTORY OF INTELLECT.
 NATURAL HISTORY OF INTELLECT.

I have used such opportunity as I have had, and lately[1] in London

and Paris, to attend scientific lectures; and in listening to Richard
Owen’s masterly enumeration of the parts and laws of the human
body, or Michael Faraday’s explanation of magnetic powers, or the
botanist’s descriptions, one could not help admiring the irresponsible
security and happiness of the attitude of the naturalist; sure of
admiration for his facts, sure of their sufficiency. They ought to
interest you; if they do not, the fault lies with you.
Then I thought—could not a similar enumeration be made of the
laws and powers of the Intellect, and possess the same claims on
the student? Could we have, that is, the exhaustive accuracy of
distribution which chemists use in their nomenclature and anatomists
in their descriptions, applied to a higher class of facts; to those laws,
namely, which are common to chemistry, anatomy, astronomy,
geometry, intellect, morals, and social life;—laws of the world?
Why not? These powers and laws are also facts in a Natural
History. They also are objects of science, and may be numbered and
recorded, like stamens and vertebræ. At the same time they have a
deeper interest, as in the order of nature they lie higher and are
nearer to the mysterious seat of power and creation.
For at last, it is only that exceeding and universal part which
interests us, when we shall read in a true history what befalls in that
kingdom where a thousand years is as one day, and see that what is
set down is true through all the sciences; in the laws of thought as
well as of chemistry.
In all sciences the student is discovering that nature, as he calls it,
is always working, in wholes and in every detail, after the laws of the
human mind. Every creation, in parts or in particles, is on the method
and by the means which our mind approves as soon as it is
thoroughly acquainted with the facts; hence the delight. No matter
how far or how high science explores, it adopts the method of the
universe as fast as it appears; and this discloses that the mind as it
opens, the mind as it shall be, comprehends and works thus; that is
to say, the Intellect builds the universe and is the key to all it
contains. It is not then cities or mountains, or animals, or globes that
any longer command us, but only man; not the fact, but so much of
man as is in the fact.
In astronomy, vast distance, but we never go into a foreign
system. In geology, vast duration, but we are never strangers. Our
metaphysic should be able to follow the flying force through all
transformations, and name the pair identical through all variety.
I believe in the existence of the material world as the expression of
the spiritual or the real, and in the impenetrable mystery which hides
(and hides through absolute transparency) the mental nature, I await
the insight which our advancing knowledge of material laws shall
furnish.
Every object in nature is a word to signify some fact in the mind.
But when that fact is not yet put into English words, when I look at
the tree or the river and have not yet definitely made out what they
would say to me, they are by no means unimpressive. I wait for
them, I enjoy them before they yet speak. I feel as if I stood by an
ambassador charged with the message of his king, which he does
not deliver because the hour when he should say it is not yet arrived.
Whilst we converse with truths as thoughts, they exist also as
plastic forces; as the soul of a man, the soul of a plant, the genius or
constitution of any part of nature, which makes it what it is. The
thought which was in the world, part and parcel of the world, has
disengaged itself and taken an independent existence.
 My belief in the use of a course on philosophy is that the student
shall learn to appreciate the miracle of the mind; shall learn its subtle
but immense power, or shall begin to learn it; shall come to know
that in seeing and in no tradition he must find what truth is; that he
shall see in it the source of all traditions, and shall see each one of
them as better or worse statement of its revelations; shall come to
trust it entirely, as the only true; to cleave to God against the name of
God. When he has once known the oracle he will need no priest.
And if he finds at first with some alarm how impossible it is to accept
many things which the hot or the mild sectarian may insist on his
believing, he will be armed by his insight and brave to meet all
inconvenience and all resistance it may cost him. He from whose
hand it came will guide and direct it.
Yet these questions which really interest men, how few can
answer. Here are learned faculties of law and divinity, but would
questions like these come into mind when I see them? Here are
learned academies and universities, yet they have not propounded
these for any prize.
Seek the literary circles, the stars of fame, the men of splendor, of
bon-mots, will they afford me satisfaction? I think you could not find a
club of men acute and liberal enough in the world. Bring the best wits
together, and they are so impatient of each other, so vulgar, there is
so much more than their wit,—such follies, gluttonies, partialities,
age, care, and sleep, that you shall have no academy.
There is really a grievous amount of unavailableness about men of
wit. A plain man finds them so heavy, dull and oppressive, with bad
jokes and conceit and stupefying individualism, that he comes to
write in his tablets, Avoid the great man as one who is privileged to
be an unprofitable companion. For the course of things makes the
scholars either egotists or worldly and jocose. In so many hundreds
of superior men hardly ten or five or two from whom one can hope
for a reasonable word.
 Go into the scientific club and hearken. Each savant proves in his
admirable discourse that he and he only knows now or ever did
know anything on the subject: “Does the gentleman speak of
anatomy? Who peeped into a box at the Custom House and then
published a drawing of my rat?” Or is it pretended discoveries of new
strata that are before the meeting? This professor hastens to inform
us that he knew it all twenty years ago, and is ready to prove that he
knew so much then that all further investigation was quite
superfluous;—and poor nature and the sublime law, which is all that
our student cares to hear of, are quite omitted in this triumphant
vindication.
Was it better when we came to the philosophers, who found
everybody wrong; acute and ingenious to lampoon and degrade
mankind? And then was there ever prophet burdened with a
message to his people who did not cloud our gratitude by a strange
confounding in his own mind of private folly with his public wisdom?
But if you like to run away from this besetting sin of sedentary
men, you can escape all this insane egotism by running into society,
where the manners and estimate of the world have corrected this
folly, and effectually suppressed this overweening self-conceit. Here
each is to make room for others, and the solidest merits must exist
only for the entertainment of all. We are not in the smallest degree
helped. Great is the dazzle, but the gain is small. Here they play the
game of conversation, as they play billiards, for pastime and credit.
Yes, ’tis a great vice in all countries, the sacrifice of scholars to be
courtiers and diners-out, to talk for the amusement of those who
wish to be amused, though the stars of heaven must be plucked
down and packed into rockets to this end. What with egotism on one
side and levity on the other we shall have no Olympus.
But there is still another hindrance, namely, practicality. We must
have a special talent, and bring something to pass. Ever since the
Norse heaven made the stern terms of admission that a man must
do something excellent with his hands or feet, or with his voice,
eyes, ears, or with his whole body, the same demand has been
made in Norse earth.
 Yet what we really want is not a haste to act, but a certain piety
toward the source of action and knowledge. In fact we have to say
that there is a certain beatitude—I can call it nothing less—to which
all men are entitled, tasted by them in different degrees, which is a
perfection of their nature, and to which their entrance must be in
every way forwarded. Practical men, though they could lift the globe,
cannot arrive at this. Something very different has to be done,—the
availing ourselves of every impulse of genius, an emanation of the
heaven it tells of, and the resisting this conspiracy of men and
material things against the sanitary and legitimate inspirations of the
intellectual nature.
What is life but the angle of vision? A man is measured by the
angle at which he looks at objects. What is life but what a man is
thinking of all day? This is his fate and his employer. Knowing is the
measure of the man. By how much we know, so much we are.

The laws and powers of the Intellect have, however, a stupendous

peculiarity, of being at once observers and observed. So that it is
difficult to hold them fast, as objects of examination, or hinder them
from turning the professor out of his chair. The wonder of the science
of Intellect is that the substance with which we deal is of that subtle
and active quality that it intoxicates all who approach it. Gloves on
the hands, glass guards over the eyes, wire-gauze masks over the
face, volatile salts in the nostrils, are no defence against this virus,
which comes in as secretly as gravitation into and through all
barriers.
Let me have your attention to this dangerous subject, which we
will cautiously approach on different sides of this dim and perilous
lake, so attractive, so delusive. We have had so many guides and so
many failures. And now the world is still uncertain whether the pool
has been sounded or not.
My contribution will be simply historical. I write anecdotes of the
intellect; a sort of Farmer’s Almanac of mental moods. I confine my
ambition to true reporting of its play in natural action, though I should
get only one new fact in a year.
I cannot myself use that systematic form which is reckoned
essential in treating the science of the mind. But if one can say so
without arrogance, I might suggest that he who contents himself with
dotting a fragmentary curve, recording only what facts he has
observed, without attempting to arrange them within one outline,
follows a system also,—a system as grand as any other, though he
does not interfere with its vast curves by prematurely forcing them
into a circle or ellipse, but only draws that arc which he clearly sees,
or perhaps at a later observation a remote curve of the same orbit,
and waits for a new opportunity, well-assured that these observed
arcs will consist with each other.
I confess to a little distrust of that completeness of system which
metaphysicians are apt to affect. ’Tis the gnat grasping the world. All
these exhaustive theories appear indeed a false and vain attempt to
introvert and analyze the Primal Thought. That is upstream, and
what a stream! Can you swim up Niagara Falls?
We have invincible repugnance to introversion, to study of the
eyes instead of that which the eyes see; and the belief of men is that
the attempt is unnatural and is punished by loss of faculty. I share
the belief that the natural direction of the intellectual powers is from
within outward, and that just in proportion to the activity of thoughts
on the study of outward objects, as architecture, or farming, or
natural history, ships, animals, chemistry,—in that proportion the
faculties of the mind had a healthy growth; but a study in the
opposite direction had a damaging effect on the mind.
Metaphysic is dangerous as a single pursuit. We should feel more
confidence in the same results from the mouth of a man of the world.
The inward analysis must be corrected by rough experience.
Metaphysics must be perpetually reinforced by life; must be the
observations of a working-man on working-men; must be biography,
—the record of some law whose working was surprised by the
observer in natural action.
 I think metaphysics a grammar to which, once read, we seldom
return. ’Tis a Manila full of pepper, and I want only a teaspoonful in a
year. I admire the Dutch, who burned half the harvest to enhance the
price of the remainder.
I want not the logic but the power, if any, which it brings into
science and literature; the man who can humanize this logic, these
syllogisms, and give me the results. The adepts value only the pure
geometry, the aerial bridge ascending from earth to heaven with
arches and abutments of pure reason. I am fully contented if you tell
me where are the two termini.
My metaphysics are to the end of use. I wish to know the laws of
this wonderful power, that I may domesticate it. I observe with
curiosity its risings and settings, illumination and eclipse; its
obstructions and its provocations, that I may learn to live with it
wisely, court its aid, catch sight of its splendor, feel its approach,
hear and save its oracles and obey them. But this watching of the
mind, in season and out of season, to see the mechanics of the
thing, is a little of the detective. The analytic process is cold and
bereaving and, shall I say it? somewhat mean, as spying. There is
something surgical in metaphysics as we treat it. Were not an ode a
better form? The poet sees wholes and avoids analysis; the
metaphysician, dealing as it were with the mathematics of the mind,
puts himself out of the way of the inspiration; loses that which is the
miracle and creates the worship.
I think that philosophy is still rude and elementary. It will one day
be taught by poets. The poet is in the natural attitude; he is believing;
the philosopher, after some struggle, having only reasons for
believing.

What I am now to attempt is simply some sketches or studies for

such a picture; Mémoires pour servir toward a Natural History of
Intellect.
 First I wish to speak of the excellence of that element, and the
great auguries that come from it, notwithstanding the impediments
which our sensual civilization puts in the way.
Next I treat of the identity of the thought with Nature; and I add a
rude list of some by-laws of the mind.
Thirdly I proceed to the fountains of thought in Instinct and
Inspiration, and I also attempt to show the relation of men of thought
to the existing religion and civility of the present time.

I. We figure to ourselves Intellect as an ethereal sea, which ebbs

and flows, which surges and washes hither and thither, carrying its
whole virtue into every creek and inlet which it bathes. To this sea
every human house has a water front. But this force, creating nature,
visiting whom it will and withdrawing from whom it will, making day
where it comes and leaving night when it departs, is no fee or
property of man or angel. It is as the light, public and entire to each,
and on the same terms.
What but thought deepens life, and makes us better than cow or
cat? The grandeur of the impression the stars and heavenly bodies
make on us is surely more valuable than our exact perception of a
tub or a table on the ground.
To Be is the unsolved, unsolvable wonder. To Be, in its two
connections of inward and outward, the mind and nature. The
wonder subsists, and age, though of eternity, could not approach a
solution. But the suggestion is always returning, that hidden source
publishing at once our being and that it is the source of outward
nature. Who are we and what is Nature have one answer in the life
that rushes into us.
In my thought I seem to stand on the bank of a river and watch the
endless flow of the stream, floating objects of all shapes, colors and
natures; nor can I much detain them as they pass, except by running
beside them a little way along the bank. But whence they come or
whither they go is not told me. Only I have a suspicion that, as
geologists say every river makes its own valley, so does this mystic
stream. It makes its valley, makes its banks and makes perhaps the
observer too. Who has found the boundaries of human intelligence?
Who has made a chart of its channel or approached the fountain of
this wonderful Nile?
I am of the oldest religion. Leaving aside the question which was
prior, egg or bird, I believe the mind is the creator of the world, and is
ever creating;—that at last Matter is dead Mind; that mind makes the
senses it sees with; that the genius of man is a continuation of the
power that made him and that has not done making him.
I dare not deal with this element in its pure essence. It is too rare
for the wings of words. Yet I see that Intellect is a science of
degrees, and that as man is conscious of the law of vegetable and
animal nature, so he is aware of an Intellect which overhangs his
consciousness like a sky, of degree above degree, of heaven within
heaven.
Every just thinker has attempted to indicate these degrees, these
steps on the heavenly stair, until he comes to light where language
fails him. Above the thought is the higher truth,—truth as yet
undomesticated and therefore unformulated.

It is a steep stair down from the essence of Intellect pure to

thoughts and intellections. As the sun is conceived to have made our
system by hurling out from itself the outer rings of diffuse ether which
slowly condensed into earths and moons, by a higher force of the
same law the mind detaches minds, and a mind detaches thoughts
or intellections. These again all mimic in their sphericity the first
mind, and share its power.
Life is incessant parturition. There are viviparous and oviparous
minds; minds that produce their thoughts complete men, like armed
soldiers, ready and swift to go out to resist and conquer all the
armies of error, and others that deposit their dangerous unripe
thoughts here and there to lie still for a time and be brooded in other
minds, and the shell not be broken until the next age, for them to
begin, as new individuals, their career.
The perceptions of a soul, its wondrous progeny, are born by the
conversation, the marriage of souls; so nourished, so enlarged. They
are detached from their parent, they pass into other minds; ripened
and unfolded by many they hasten to incarnate themselves in action,
to take body, only to carry forward the will which sent them out. They
take to themselves wood and stone and iron; ships and cities and
nations and armies of men and ages of duration; the pomps of
religion, the armaments of war, the codes and heraldry of states;
agriculture, trade, commerce;—these are the ponderous
instrumentalities into which the nimble thoughts pass, and which
they animate and alter, and presently, antagonized by other thoughts
which they first aroused, or by thoughts which are sons and
daughters of these, the thought buries itself in the new thought of
larger scope, whilst the old instrumentalities and incarnations are
decomposed and recomposed into new.
Our eating, trading, marrying, and learning are mistaken by us for
ends and realities, whilst they are properly symbols only; when we
have come, by a divine leading, into the inner firmament, we are
apprised of the unreality or representative character of what we
esteemed final.
So works the poor little blockhead manikin. He must arrange and
dignify his shop or farm the best he can. At last he must be able to
tell you it, or write it, translate it all clumsily enough into the new sky-
language he calls thought. He cannot help it, the irresistible
meliorations bear him forward.

II. Whilst we consider this appetite of the mind to arrange its

phenomena, there is another fact which makes this useful. There is
in nature a parallel unity which corresponds to the unity in the mind
and makes it available. This methodizing mind meets no resistance