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Cell Unit
• Chapter 4. General Features of Cells: This chapter has an
expanded discussion of how to calculate the surface area-to-­
volume ratio, depending on the shape of a cell.
• Chapter 5. Membrane Structure, Synthesis, and Transport:
The section of membrane transport has been revised to better
­distinguish simple diffusion from facilitated diffusion.
• Chapter 6. An Introduction to Energy, Enzymes, and
Metabolism: This chapter has a new BioTIPS feature that
helps students understand how a mutation could affect an
­allosteric site in an enzyme.
• Chapter 7. Cellular Respiration and Fermentation: The
­section on oxidative phosphorylation has been divided into two
sections, the second of which takes a closer look at ATP synthase.
• Chapter 8. Photosynthesis: This chapter has a new BioTIPS
feature that asks students to distinguish between the light-­
harvesting complex, P680, and the primary electron acceptor.
• Chapter 9. Cell Communication: In Biology, 4th edition, the
topic of apoptosis has been expanded to include a discussion of
how it plays a role in certain diseases (see Table 9.2).
• Chapter 10. Multicellularity: The section comparing animal
and plant cells has been streamlined. The discussion of plasmo-
desmata in plants has also been revised.
Genetics Unit
• Chapter 11. Nucleic Acid Structure, DNA Replication, and
Chromosome Structure: This chapter has a new BioTIPS
feature that pertains to the role of DNA ligase during DNA
replication. The illustration regarding the Avery, MacLeod,
and McCarty experiment has been revised (see Figure 11.2).
• Chapter 12. Gene Expression at the Molecular Level: The
section on RNA modification (formerly RNA processing) has
been reorganized.
• Chapter 13. Gene Regulation: A new section has been added
(Section 13.5) on Epigenetic Gene Regulation, including a new
figure (Figure 13.21) and two new tables (Tables 13.1 and 13.2).
• Chapter 14. Mutation, DNA Repair, and Cancer: Section 14.1
that was in Biology, 3rd edition, is now subdivided into two sec-
tions on the Causes of Mutation and the Consequences of
Mutation.
• Chapter 15. The Eukaryotic Cell Cycle, Mitosis, and
­Meiosis: The Genomes & Proteomes Connection has been
changed to the topic of how mitosis in eukaryotes evolved from
binary fission that occurs in prokaryotic cells (see Figure 15.10).
• Chapter 16. Simple Patterns of Inheritance: This chapter has
a new BioTIPS feature that helps students learn how to use prob-
ability and a Punnett square to predict the outcome of crosses.
• Chapter 17. Complex Patterns of Inheritance: Building on
the BioTIPS feature in Chapter 17, Chapter 18 has a BioTIPS
feature that focuses on maternal inheritance.
• Chapter 18. Genetics of Viruses and Bacteria: The first part
of Chapter 18 has been reorganized by adding a new section
(Section 18.2) on viral reproductive cycles.
• Chapter 19. Developmental Genetics: In many places, the
information in this chapter has been broken down into bulleted
and numbered lists to make it easier for students to follow. The
illustration of the effects of the bicoid mutation has been revised
(see Figure 19.8).
• Chapter 20. Genetic Technology: In Chapter 20 of Biology, 4th
edition, a new subsection has been added on how genetically modi-
fied organisms can make products that are useful to people (also see
Table 20.1).
• Chapter 21. Genomes, Proteomes, and Bioinformatics: The
rapidly changing information on the properties of bacterial,
archaeal, and eukaryotic genomes has been updated.
Evolution Unit
• Chapter 22. The Origin and History of Life: This chapter has
a new BioTIPS feature that helps students to learn how the cal-
culation of radioisotope decay is used to date fossils.
• Chapter 23. An Introduction to Evolution: The topic of whale
evolution from tetrapod mammals has been added, along with a
new figure (see Figure 23.7).
• Chapter 24. Population Genetics: An example has been added
to help students use the Hardy-Weinberg equation to calculate
allele and genotype frequencies.
• Chapter 25. Origin of Species and Macroevolution: This
chapter has a new BioTIPS feature that helps students learn how
to design an experiment to determine if two populations consti-
tute different species.
• Chapter 26. Taxonomy and Systematics: The information on
eukaryotic supergroups has been updated. The chapter also
includes an example to help students learn how to assess evolu-
tionary trees using the principle of parsimony.
Diversity Unit
• Chapter 27. Archaea and Bacteria: Phylogenetic history of
major lineages of Bacteria and Archaea has been updated to
reflect recent progress, including identification of archaea most
closely related to eukaryotes.
• Chapter 28. Protists: The phylogenetic history of major lin-
eages of protists has been updated to reflect recent progress,
including identification of the root of the eukaryotic tree of life.
A new Feature Investigation describes an experiment that tests
hypotheses of protist fossil identifications and provides results
useful in industrial applications of materials produced by com-
mon protists.
• Chapter 29. Plants and the Conquest of Land: Table 29.1 that
summarizes the features of major groups of land plants has been
reorganized to enhance reader access.
• Chapter 30. The Evolution and Diversity of Modern
Gymnosperms and Angiosperms: To emphasize the
­
­importance of whole genome duplication (WGD) events in the
­evolutionary history of seed plants, major WGD events have been
mapped onto phylogenetic diagrams.
• Chapter 31. Fungi: Phylogenetic history of the fungi has been
updated to reflect recent progress. Table 31.1 has been simplified
to include major derived fungal lineages of most concern to
human society.
• Chapter 32. An Introduction to Animal Diversity: The
chapter opening photograph has been changed to illustrate a
IMPROVING BIOLOGY EDUCATION vii
 naked mole rat. The introductory paragraph discusses some of
the unusual features of naked mole rat biology as well as the
potential value of naked mole rats to medicine. Some data in
Table 32.3 have been updated.
• Chapter 33. The Invertebrates: The sections on both cnidarians
and flatworms have been simplified. A new photograph has been
provided to illustrate a molluscan veliger larva. The sections on
the annelid body plan and crustacean diversity have been rewrit-
ten. The major echinoderm classes are now illustrated using five
new photographs.
• Chapter 34. The Vertebrates: The section introducing the ver-
tebrates has been simplified. The material on the chondrchthy-
ans has been completely re-organized. New photographs have
been used to illustrate reptile diversity.
Flowering Plants Unit
• Chapter 38. Flowering Plants: Transport: The Feature Inves-
tigation has been replaced. The new Feature Investigation dis-
cusses new ways to genetically modify crop plants for enhanced
drought resistance. The Genomes & Proteomes feature was also
replaced. The new Genomes & Proteomes feature explores the
evolutionary history of plant resistance to drought. Both of these
are related to the increasing need to feed populations living in
arid lands and the need to cope with climate change.
Animal Unit
• Chapter 40. Animal Bodies and Homeostasis: A new confo-
cal micrograph showing the complexity of nervous tissue has
been added, and other figures have been modified for additional
clarity. Several assessments have been added that require mak-
ing predictions and analyzing data, including a new BioConnec-
tions question. A new BioTIPS feature related to body
temperature homeostasis has been added.
• Chapter 41. Neuroscience I: Cells of the Nervous System:
The material on resting membrane potential and action potentials
has been subdivided into seven shorter sections to allow this chal-
lenging material to be more easily assimilated by beginning stu-
dents. Several new assessments, BioConnections, and SciSKILLS
have been added that require critical thinking and the ability to
make predictions and perform calculations. A new figure illus-
trates the structural features of nerves. A new BioTIPS feature
related to the ionic basis of action potentials has been added.
• Chapter 42. Neuroscience II: Evolution, Structure, and Func-
tion of the Nervous System: Several figures have been revised
for clarity and new detail, particularly that of the structures of the
human brain. Test questions have been updated to include higher
Bloom’s level questions. The description of the evolution of the
brain has been updated, as have the discussions of the formation of
memories and the effects of training on brain plasticity. The discus-
sion of the anatomy of the vertebrate central nervous system has
been reorganized and broken into smaller segments for greater
clarity and manageability. A new BioTIPS feature related to the
anatomical features of the vertebrate brain has been added.
• Chapter 43. Neuroscience III: Sensory Systems: The chapter
has been reorganized by the addition of numerous subheadings
to help break up complex areas of text into more manageable
viii IMPROVING BIOLOGY EDUCATION
portions for the reader. Ten figures have been revised and
updated with additional labeling and text boxes to help the
reader navigate through complex topics. Many of the test ques-
tions at the end of the chapter have been revised, updated, and,
where appropriate, changed to a higher Bloom’s level. The
chapter has been tightened to focus the reader on fundamental
features of sensory biology. A new BioTIPS feature asks stu-
dents to make conclusions about the relation between structure
and function in different organs.
• Chapter 44. Muscular-Skeletal Systems and Locomotion:
The chapter has been reorganized to include more subhead-
ings and bulleted lists to assist the reader in navigating
through complex material, particularly the material on the
sarcomere and its bands, and material on the cross bridge
cycle. Numerous figures have been revised for clarity and
improved labeling and visibility. A new discussion and figure
relating to myostatin have been added. Many of the test ques-
tions at the end of the chapter have been updated or revised
including the addition of higher level Bloom’s questions. A
new BioTIPS feature asks students to make calculations and
interpret graphical data related to the metabolic cost of flying.
• Chapter 45. Nutrition and Animal Digestive Systems: The
chapter has been consolidated and shortened to focus less on
feeding behavior and more on the processes of digestion and
absorption. The chapter has been reorganized with eleven new
subheadings for easier navigation through complex material. A
new figure illustrating protein digestion and absorption in the
small intestine has been added, and several figures have been
improved with additional labeling and text. A new Feature Inves-
tigation (Marshall and Warren’s investigation of the relation of
H. pylori to stomach ulcers) has been added. Test questions at the
end of the chapter and in the Feature Investigation have been
modified to include higher order Bloom’s questions. A new
BioTIPS feature asks students to predict the benefits of segre-
gated function in the alimentary canal.
• Chapter 46. Control of Energy Balance, Metabolic Rate,
and Body Temperature: Numerous figures and test questions
have been modified and updated, and the text has been reorga-
nized to include several new subheadings. The chapter has been
shortened to better focus on key, fundamental features of energy
balance and thermal regulation. A new BioTIPS feature has been
added that asks students to propose an hypothesis regarding the
experimental effects of the hormone leptin on body mass in mice.
• Chapter 47. Circulatory Systems: The chapter has been short-
ened and consolidated to better focus on the similarities and dif-
ferences between open and closed circulations, and single and
double circulations. The events of the ECG have been reorga-
nized into a bulleted list for emphasis and clarity. The table
describing comparative features of mammalian hearts has been
expanded. Numerous new subheadings have been added, as have
several new test questions and Concept Checks. A new BioTIPS
feature has been added that requires students to describe the
sequence of steps in the vertebrate heart cycle.
• Chapter 48. Respiratory Systems: Several new subheadings
have been added, as have numerous new or revised test ques-
tions, Concept Checks, BioConnections, and collaborative ques-
 tions. A new figure illustrating asthma and bronchoconstriction
has been added. A new Feature Investigation describing the
effectiveness of surfactant in human newborns has been added. A
new photomicrograph of an emphysematous lung has been
added. A new BioTIPS feature has been added that requires
­students to make a graph and make calculations regarding the
relation between altitude and oxygen pressure.
• Chapter 49. Excretory Systems and the Homeostasis of Inter-
nal Fluids: The title has been changed to reflect the important role
of excretory systems in homeostasis. The concept of osmolarity is
now introduced very early in the text, to lay the groundwork for
later discussions in the chapter. Numerous bulleted lists (for exam-
ple, in the discussion about individual parts of a nephron) and sub-
headings have been added to break up complex material and
provide a useful study guide. Many of the test questions have been
updated or revised, including BioConnection questions and Fea-
ture Investigation questions, in some cases where appropriate at
higher Bloom’s levels. A new BioTIPS feature has been added that
requires students to relate the structure of a nephron to its function.
• Chapter 50. Endocrine Systems: Eleven figures have been
revised with improved labeling and text box detail where appro-
priate, to better help the student work through illustrations of
complex processes. Numerous subheadings and revised test
questions, including Feature Investigation questions, have been
added. The introductory material on the chemical nature and
mechanisms of actions of hormones has been consolidated and
tightened to focus on major concepts. The material on endocrine
disruptors has been updated. A new BioTIPS feature has been
added that requires students to predict the outcome of a scenario
in which Ca2+ homeostasis may be compromised.
• Chapter 51. Animal Reproduction: New conceptual, collab-
orative, and multiple choice questions have been added to the
end of the chapter. Several new subheadings and six revised fig-
ures have been added. A new BioTIPS feature has been added
that requires students to predict what might happen in a person
who takes supplements of the androgen testosterone.
• Chapter 52. Animal Development: New BioConnection ques-
tions, Concept Checks, and multiple choice questions have been
added. The figures on fertilization, cleavage, and neurulation
have been revised. The section on morphogenesis and control of
development has been shortened, consolidated and reorganized
to reduce redundancy and improve clarity. A new BioTIPS fea-
ture has been added that asks students to place various events in
their correct stage of development.
• Chapter 53. Immune Systems: Eleven figures have been
revised and updated (such as the latest figures on people living
with HIV worldwide). New subheadings have been added to
shorten complex regions of text (for example, the discussion of
the production and recirculation of lymphocytes). Several new
collaborative questions and Concept Checks have been added.
A new figure of a micrograph of a mast cell has been
added. The discussion of the different cell types of the immune
system now includes a bulleted list as a visual aid to the reader
and a handy study reference. A new BioTIPS feature has been
added that requires students to make a prediction about
immune responses in the absence of memory cells.
Ecology Unit
• Chapter 54. An Introduction to Ecology and Biomes: Section
54.2, Ecological Methods, has been completely rewritten. Instead
of using a hypothetical example of locust ecology, we have instead
used a real life example of oak winter moth ecology. We discuss
how knowledge of the oak winter moth life cycle was vital in
addressing control of this important apple orchard pest in the U.S.
northwest and western Canada. Several new figures have been pre-
pared to help in illustrating main points of this section. In Section
54.3, The Environment’s Effect on the Distribution of Organisms,
we have provided a new example of the effects of cold temperatures
on organismal distribution by including a figure on how the fre-
quency of the cyanide-­producing form of white clover in Europe is
affected by temperature. In the section on Global Warming, data on
selected greenhouse gases have been updated in Table 54.2. Many
new photographs have been included in Section 54.5 on major
biomes. Finally, in Section 54.6, Continental Drift and Biogeogra-
phy, we have included a new figure on the distribution of fossil land
plants and animals to illustrate the concept of continental drift.
• Chapter 55. Behavioral Ecology: In Section 55.3, Foraging
Behavior, we have illustrated the concept of optimal foraging by
including new information, and a new figure, on optimal forag-
ing by leafcutter ants, and the disruption of this foraging by para-
sitic flies. We have also simplified the chapter by eliminating the
section on sexual selection and mate choice, which is already
covered in Chapter 24.
• Chapter 56. Ecology: The section on Quantifying Population
Density has been divided into more easily digestible parts by the
use of subheadings. The section of Human Population Growth
has been updated.
• Chapter 57. Species Interactions: The section on Predation,
Herbivory, and Parasitism has been substantially rewritten and
simplified. The example of cactus control by the cactus moth,
Cactoblastis cactorum, has been added to this section. Section
57.4, on bottom-up and top-down control, has been simplified
and a new figure on the nitrogen content of plants and animals
has been added. In addition, a new figure illustrates the tro-
phic interactions that can result from the effects of wolf addi-
tion, using data from Banff National Park, Canada.
• Chapter 58. Community Ecology: In Section 58.3, Calculat-
ing Species Diversity, we introduce the new concept of effective
number of species and provide details of how to calculate it.
This measure of species diversity permits easy and meaningful
comparisons between different communities.
• Chapter 59. Ecosystem Ecology: The distinction between
decomposers and detritivores has been made clearer throughout
the chapter. Figure 59.19, which concerns the increase in atmo-
sphere CO2 levels and temperatures due to the burning of fossil
fuels, has been updated.
• Chapter 60. Biodiversity and Conservation Biology: Section
60.2, Why Conserve Biodiversity, has been updated. Tables 60.1
and 60.2, which provide data on examples of the world’s ecosys-
tem services and the valuation of those services, have been com-
pletely rewritten and modernized using the latest data. We have
also added an additional question to the Feature Investigation.
IMPROVING BIOLOGY EDUCATION ix
 A NOTE FROM THE AUTHORS
The lives of most science-textbook authors do not revolve around an analysis of
writing techniques. Instead, we are people who understand science and are inspired
by it, and we want to communicate that information to our students. Simply put,
we need a lot of help to get it right.
Editors are a key component that help the authors modify the content of their
book so it is logical, easy to read, and inspiring. The editorial team for this Biol-
ogy textbook has been a catalyst that kept this project rolling. The members
played various roles in the editorial process. Rebecca Olson and following her,
Justin Wyatt, Brand Managers (Major Biology) did outstanding jobs overseeing
the fourth edition. Their insights with regard to pedagogy, content, and organiza-
tion have been invaluable. Elizabeth Sievers, Lead Product Developer, has been
the master organizer. Liz’s success at keeping us on schedule is greatly appreci-
ated. We would also like to acknowledge our copy editor, Jane Hoover, for keep-
ing our grammar on track.
Another important aspect of the editorial process is the actual design, presentation,
and layout of materials. It’s confusing if the text and art aren’t on the same page, or
if a figure is too large or too small. We are indebted to the tireless efforts of Jayne
Klein, Content Project Manager; and David Hash, Senior Designer at McGraw-
Hill. Likewise, our production company, Lachina Publishing Services, did an
excellent job with the paging, revision of existing art, and the creation of new art
for the fourth edition. Their artistic talents, ability to size and arrange figures, and
attention to the consistency of the figures have been remarkable.
We would like to acknowledge the ongoing efforts of the superb marketing staff
at McGraw-Hill. Special thanks to Patrick Reidy, Executive Marketing Manager-
Life Sciences, for his ideas and enthusiasm for this book.
Finally, other staff members at McGraw-Hill Higher Education have ensured that
the authors and editors were provided with adequate resources to achieve the goal
of producing a superior textbook. These include Kurt Strand, Senior Vice Presi-
dent, Products & Markets, and Marty Lange, Vice President, General Manager,
Products & Markets, and Michael Hackett, Director for Life Sciences.

EMPHASIZING PROBLEM-SOLVING AND SKILLS DEVELOPMENT

Skills Development SciSKILLS is a mental action such as analyze data, form hypoth-
eses, make predictions, or perform calculations. These are skills
At the beginning of each section, Learning Outcomes inform
scientists generally perform and students should practice.
students of concepts they should understand and new to the
The emphasis on skills development continues in the F ­ eature
fourth edition are skills-based Learning Outcomes. Labeled as
Investigations. Feature Investigations provide a complete descrip-
SciSKILLS, these Learning Outcomes are specific to the skills
tion of experiments, including data analysis, so students can
students will acquire when mastering the material and provide
understand how experimentation leads to an understanding of bio-
a specific understanding of how such skills may be assessed.
logical concepts.

x IMPROVING BIOLOGY EDUCATION

Problem Solving
A new feature added to Biology, 4th edition, will help students
develop their problem solving skills. Chapters 2 through 60 con-
tain solved problems called BioTIPS, which stands for Topic,
Information, and Problem Solving Strategy. These solved prob-
lems follow a consistent pattern in which students are given
advice on how to solve problems in biology using different types
of problem solving strategies. These strategies include: Make a
drawing; Compare and contrast; Relate structure and function;
Sort out the steps in a complicated process; Propose a hypothesis;
Design an experiment; Predict the outcome; Interpret data; Use
statistics; Make a calculation; and Search the literature.
A biological question related to chapter content is posed. The
BioTIPS then walks the student through the process of answer-
ing the question. First helping the student identify the topic of
the question—what is really being asked in the question? Then
helping the student collect information that was presented in the
chapter that is related to the question. Finally helping the student
settle on one or more strategies that can be followed to answer the
question. The answers are provided to complete the problem solv-
ing process.
The emphasis on developing problem solving skills is carried
over to the digital content, as described later.

USING STUDENT USAGE DATA TO MAKE IMPROVEMENTS

To help guide the revision for the fourth edition, student usage Below is an example of one of the heat maps from Chapter 8.
data and input were used, derived from thousands of SmartBook® The color-coding of highlighted sections indicates the various
users of the 3rd edition. SmartBook “heat maps” provided a quick levels of difficulty students experienced in learning the material,
visual snapshot of chapter usage data and the relative difficulty topics highlighted in red being the most challenging for students.
students experienced in mastering the content. These data directed
the authors to evaluate text content that was particularly challeng-
ing for students. These same data were also used to revise the
SmartBook probes.
• If the data indicated that the subject was more difficult than
other parts of the chapter, as evidenced by a high proportion of
students responding incorrectly to the probes, the authors revised
or reorganized the content to be as clear and illustrative as pos-
sible by rewriting the section, providing additional examples or
revised figures to assist visual learners, etc.
• In other cases, one or more of the SmartBook probes for a
section was not as clear as it might be or did not appropriately
reflect the content in the chapter. In these cases the probe, rather
than the text, was edited.

IMPROVING BIOLOGY EDUCATION xi

MAKING CONNECTIONS

Principles of Biology are introduced

in Chapter 1 and are then threaded
throughout the entire textbook. This
is achieved in two ways. First, the
principles are explicitly stated in
selected figure legends for figures in
which the specific principle is illus-
trated. The legends that relate to a
Principle of Biology are highlighted
with an icon.

In addition, a Conceptual
Question at the end of
each chapter is directly
aimed at exploring a par-
ticular principle related to
the content of the chapter.

 ioConnections BioConnections are questions found in

B
selected figure legends in each chapter that help students
make connections between biological concepts. BioCon-
nections help students understand that their study of biol-
ogy involves linking concepts together and building on
previously learned information. Answers to the BioCon-
nections are found in Appendix B.

•Unit openers serve two purposes. They allow the student to see
the big picture of the unit. In addition, the unit openers draw
attention to the principles of biology that will be emphasized in
that unit.

xii IMPROVING BIOLOGY EDUCATION

 Strengthen Problem Solving Skills and Key
Concept Development with Connect®
Problem Solving Skills Key Concept Development
Detailed Feedback in Connect® SmartBook with Learning Resources
Learning is a process of iterative development, of making mis- To help students understand key concepts, SmartBook® for
takes, reflecting, and adjusting over time. The question and test ­Biology, 4th edition, is enhanced with Learning Resources. Based
banks in Connect® for Biology, 4th edition, are more than direct on student usage data, derived from thousands of SmartBook
assessments; they are self-contained learning experiences that users of the third edition, concepts that proved more challenging
systematically build student learning over time. for students are supported with Learning Resources to enhance
For many students, choosing the right answer is not necessarily the textbook presentation. Learning Resources, such as anima-
based on applying content correctly; it is more a matter of increasing tions and tutorials, are indicated in Smartbook adjacent to the
their statistical odds of guessing. A major fault with this approach is textbook content. If a student is struggling with a concept based
students don’t learn how to process the questions correctly, mostly on his/her performance of the SmartBook questions, the student
because they are repeating and reinforcing their mistakes rather is given an option to review the Learning Resource or the student
than reflecting and learning from them. To help students develop can click on the Learning Resources at any time.
problem solving skills, all higher level Blooms questions in Con-
nect are supported with hints, to help students focus on important
information for answering the questions, and detailed feedback that
walks students through the problem solving process, using Socratic
questions in a decision tree-style framework to scaffold learning,
where each step models and reinforces the learning process.
The feedback for each higher level Blooms question (Apply,
Analyze, Evaluate) follows a similar process: Clarify Question,
Gather Content, Consider Possibilities, Choose Answer, Reflect
on Process.

Unpacking the Concepts

We’ve taken problem solving a step further. In each chapter, 3 to
5 higher level Blooms questions in the question and test banks are
broken out by the steps in detailed feedback. Rather than leaving it
up to the student to work through the detailed feedback, a second
version of the question is presented in a stepwise format. Follow-
ing the problem solving steps, students need to answer questions
about earlier steps, such as “What is the key concept addressed by
the question?” before proceeding to answer the question. A pro-
fessor can choose which version of the question to include in the
assignment based on the problem solving skills of the students.

xiii
 Using Connect and Biology, 4th edition
Biology, 4th edition, and its online assets have been carefully crafted to help professors and students, work efficiently and effectively
through the material in the course, making the most of instructional and study time.

PREPARE FOR THE COURSE

Prepare for
the course
Many biology students struggle the first few weeks of class.
Many institutions expect students to start majors biology hav-
ing a working knowledge of basic chemistry and cellular biol-
ogy. LearnSmart Prep is now available in Connect. Professors
can assign modules in LearnSmart Prep to help students get up to
speed on core concepts or students can access LearnSmart Prep
directly through the LearnSmart Prep link. Prepare before
class
LearnSmart Prep is an adaptive
learning tool designed to increase
student success and aid retention
through the first few weeks of class. Using this digital tool, Majors
Biology students can master some of the most fundamental and
challenging principles of biology before they begin to struggle in Tying it all
the first few weeks of class together develops Engage in class
critical-thinking
skills

1 A diagnostic establishes your baseline comprehension and

knowledge; then the program generates a learning plan tailored to
your academic needs and schedule. 2 As you work through the learning plan, the program asks you
questions and tracks your mastery of concepts. If you answer
questions about a particular concept incorrectly, the program will
provide a learning resource (ex. animation or tutorial) on that
concept, then ensure that you understand the concept by asking
you more questions. Didn’t get it the first time? Don’t worry—
LearnSmart Prep will keep working with you!

3 Using LearnSmart Prep, you can identify the content you

don’t understand, focus your time on content you need to know
but don’t, and therefore improve your chances of success in your
majors biology course.

xiv
 Prepare Before Class
Prepare for Students who are most successful in college are those who have developed effective study
the course skills and who use those skills, before, during, and after class.

Students can maximize time in class by previewing the material before stepping into the
lecture hall. Biology, 4th edition, is available in two formats: the printed text as well as the
online SmartBook. Students can use either of these options to preview the material before
lecture. Becoming familiar with terminology and basic concepts will allow students to fol-
low along in class and engage in the content in a way that allows for better retention.
Prepare before
class
Professors can help students prepare for class by making assignments—SmartBook
assignments are effective for introducing terminology and general concepts.

Tying it all
together develops Engage in class
critical-thinking
skills

SmartBook provides a personalized, adaptive reading experience.

Powered by an intelligent diagnostic and adaptive engine, SmartBook

facilitates the reading process by identifying what content a student knows
and doesn’t know through adaptive assessments.

The reports in SmartBook

help identify topics where
you need more work.

▲
The SmartBook
experience starts by
previewing key con-
cepts from the chapter
and ensuring that you
▲
understand the big
ideas. SmartBook asks you questions that identify gaps in your
knowledge. The reading experience then continuously adapts
in response to the assessments—highlighting the material you
need to review based on what you don’t know.

USING CONNECT AND BIOLOGY, 4TH EDITION xv

 Engage in Class
Prepare for McGraw-Hill Connect provides online presentation, assignment, and
the course assessment solutions. It connects students with the tools and resources
they’ll need to achieve success. A robust set of questions and activities is
presented in the Question Bank and a separate set of questions to use for
exams are presented in the Test Bank. Instructors can edit existing ques-
tions and author entirely new problems. Track individual student perfor-
mance—by question, assignment, or in relation to the class overall—with
detailed grade reports.
Prepare before
class

Tying it all
together develops Engage in class
critical-thinking
skills

1 Pre-class assignments to help students engage in the content during class.

Assignments are accessed ▶

through Connect and
could include homework
assignments, quizzes,
SmartBook assignments,
and other resources.

◀ Interactive and traditional

questions help assess your
knowledge of the material.

xvi USING CONNECT AND BIOLOGY, 4TH EDITION

 2 Connect Insight is Connect,s visual analytics dashboard for instructors and students.

◀ Provides at-a-glance student

performance on assignments.
Instructors can use the information
for a just-in-time approach to
teaching.

◀ Connect Insight presents data that

empowers students to improve
performance that is efficient and
effective.

Tying It All Together

Follow up class with assessment that helps students develop prob-
Prepare for
the course
lem solving skills. Set up assignments from the various assessment
banks in Connect.

The Question and Test Banks contain higher order critical think-
ing questions that require students to demonstrate a more in-depth
understanding of the concepts—instructors can quickly and easily
filter the banks for these questions using higher level Blooms tags.
Prepare before
class

◀ Detailed Feedback All higher

level Blooms questions, that
involve problem solving,
contain detailed feedback in
Connect. The feedback walks
Tying it all students through the steps of
together develops Engage in class
critical-thinking the problem solving process
skills and helps them evaluate their
scientific thinking skills.

Many chapters also contain a Quantitative Question ▶

Bank. These are more challenging algorithmic
questions, intended to help your students practice
their quantitative reasoning skills. Hints and guided
solution options step students through a problem.

USING CONNECT AND BIOLOGY, 4TH EDITION xvii

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 Contents

CONTENTS
Chapter 1 Feature Investigation: Anfinsen Showed That
the Primary Structure of Ribonuclease Determines
Its Three-Dimensional Structure 60
An Introduction to Biology 1
Genomes & Proteomes Connection: Proteins Contain
1.1 Principles of Biology and the Levels of Biological Functional Domains 62
Organization 2 3.7 Nucleic Acids 63
1.2 Unity and Diversity of Life 6
Genomes & Proteomes Connection: The Study of Genomes
and Proteomes Provides an Evolutionary Foundation for Our
UNIT II  Cell
Understanding of Biology 9
1.3 Biology as a Scientific Discipline 14
Feature Investigation: Observation and Experimentation Form
the Core of Biology 18

UNIT I  Chemistry

Chapter 4

General Features of Cells 68

Chapter 2
The Chemical Basis of Life I:
Atoms, Molecules, and Water 23
2.1 Atoms 23
Feature Investigation: Rutherford Determined the Modern
Model of the Atom 24
2.2 Chemical Bonds and Molecules
2.3 Properties of Water 35
4.1 Microscopy 68
4.2 Overview of Cell Structure 71
Genomes & Proteomes Connection: The Characteristics
of a Cell Are Largely Determined by the Proteins
It Makes 72
4.3 The Cytosol 76
4.4 The Nucleus and Endomembrane System 81
Feature Investigation: Palade Discovered That Proteins
Destined for Secretion Move Sequentially Through Organelles
of the Endomembrane System 84
4.5 Semiautonomous Organelles 89
29 4.6 Protein Sorting to Organelles 92
4.7 Systems Biology of Cells: A Summary 96

Chapter 3 Chapter 5
The Chemical Basis of Life II: Membrane Structure, Synthesis,
Organic Molecules 44 and Transport 100
3.1 The Carbon Atom 44 5.1 Membrane Structure 101
3.2 Formation of Organic Molecules and Genomes & Proteomes Connection: Approximately 20–30% of
Macromolecules 47 All Genes Encode Transmembrane Proteins 102
3.3 Overview of the Four Major Classes of Organic 5.2 Fluidity of Membranes 104
Molecules Found in Living Cells 47 5.3 Synthesis of Membrane Components
3.4 Carbohydrates 47 in Eukaryotic Cells 106
3.5 Lipids 51 5.4 Overview of Membrane Transport 108
3.6 Proteins 55

xxi
 5.5 Transport Proteins 111 Chapter 9
Feature Investigation: Agre Discovered That Osmosis Occurs
More Quickly in Cells with a Channel That Allows the Cell Communication 175
CONTENTS

Facilitated Diffusion of Water 112

5.6 Exocytosis and Endocytosis 117 9.1 General Features of Cell Communication 175
9.2 Cellular Receptors and Their Activation 179
Chapter 6 9.3 Signal Transduction and the Cellular Response 182
9.4 Hormonal Signaling in Multicellular Organisms 187
An Introduction to Energy, Enzymes, Genomes & Proteomes Connection: A Cell’s Response to
Hormones and Other Signaling Molecules Depends on the
and Metabolism 121
Proteins It Makes 188
6.1 Energy and Chemical Reactions 121 9.5 Apoptosis: Programmed Cell Death 188
Genomes & Proteomes Connection: Many Proteins Use ATP as Feature Investigation: Kerr, Wyllie, and Currie Found That
a Source of Energy 124 Hormones May Control Apoptosis 189
6.2 Enzymes and Ribozymes 125
Feature Investigation: The Discovery of Ribozymes by Sidney Chapter 10
Altman Revealed That RNA Molecules May Also Function as
Catalysts 129 Multicellularity 194
6.3 Overview of Metabolism 131
6.4 Recycling of Organic Molecules 134 10.1 Extracellular Matrix and Cell Walls 195
Genomes & Proteomes Connection: Collagens Are a Family
of Proteins That Give the ECM of Animals a Variety of
Chapter 7 Properties 197
10.2 Cell Junctions 200
Cellular Respiration and Fermentation 138
Feature Investigation: Loewenstein and Colleagues Followed
7.1 Overview of Cellular Respiration 138 the Transfer of Fluorescent Dyes to Determine the Size of ­
Gap-Junction Channels 203
7.2 Glycolysis 140
10.3 Tissues 206
Genomes & Proteomes Connection: Cancer Cells Usually
Exhibit High Levels of Glycolysis 141
7.3 Breakdown of Pyruvate 143 UNIT III  Genetics
7.4 Citric Acid Cycle 144
7.5 Overview of Oxidative Phosphorylation 146
7.6 A Closer Look at ATP Synthase 148
Feature Investigation: Yoshida and Kinosita Demonstrated That
the γ Subunit of ATP Synthase Spins 150
7.7 Connections Among Carbohydrate, Protein, and Fat
Metabolism 151
7.8 Anaerobic Respiration and Fermentation 152

Chapter 8
Photosynthesis 156
8.1 Overview of Photosynthesis 156
8.2 Reactions That Harness Light Energy 159
Genomes & Proteomes Connection: The Cytochrome
Complexes of Mitochondria and Chloroplasts Contain
Evolutionarily Related Proteins 163
8.3 Molecular Features of Photosystems 163
8.4 Synthesizing Carbohydrates via the Calvin Cycle
Feature Investigation: The Calvin Cycle Was Determined by
Isotope-Labeling Methods 168
8.5 Variations in Photosynthesis 170
Chapter 11
Nucleic Acid Structure, DNA Replication,
and Chromosome Structure 211
11.1 Biochemical Identification of the Genetic Material 211
Feature Investigation: Avery, MacLeod, and McCarty Used
Purification Methods to Reveal That DNA Is the Genetic
Material 213
166 11.2 Nucleic Acid Structure 215
11.3 Overview of DNA Replication 219

xxii CONTENTS
11.4 Molecular Mechanism of DNA Replication 222 Chapter 15
Genomes & Proteomes Connection: DNA Polymerases Are a
Family of Enzymes with Specialized Functions 227 The Eukaryotic Cell Cycle, Mitosis,

11.5 Molecular Structure of Eukaryotic Chromosomes 229
Chapter 12
Gene Expression at the Molecular Level 234
12.1 Overview of Gene Expression 235
12.2 Transcription 238
12.3 RNA Modification in Eukaryotes 240
12.4 Translation and the Genetic Code 242
Feature Investigation: Nirenberg and Leder Found That RNA
Triplets Can Promote the Binding of tRNA to Ribosomes 245
12.5 The Machinery of Translation 247
Genomes & Proteomes Connection: Comparisons of Small
Subunit rRNAs Among Different Species Provide a Basis for
Establishing Evolutionary Relationships 250
12.6 The Stages of Translation 251
Chapter 13
Gene Regulation 257
13.1 Overview of Gene Regulation 257
13.2 Regulation of Transcription in Bacteria 260
Feature Investigation: Jacob, Monod, and Pardee Studied
a Constitutive Mutant to Determine the Function of the Lac
Repressor 264
13.3 Regulation of Transcription in Eukaryotes: Roles
of Transcription Factors and Mediator 269
13.4 Regulation of Transcription in Eukaryotes: Changes in
Chromatin Structure and DNA Methylation 271
13.5 Epigenetic Gene Regulation 274
13.6 Regulation of RNA Modification and Translation in
Eukaryotes 276
Genomes & Proteomes Connection: Alternative Splicing Tends
to Be More Prevalent in Complex Eukaryotic Species 277
Chapter 14
Mutation, DNA Repair, and Cancer 281
14.1 Consequences of Mutations 281
14.2 Causes of Mutations 285
Feature Investigation: The Lederbergs Used Replica Plating to
Show That Mutations Are Random Events 285
14.3 DNA Repair 289
14.4 Cancer 291
Genomes & Proteomes Connection: Mutations in
Approximately 300 Human Genes May Promote Cancer
CONTENTS
and Meiosis 300
15.1 The Eukaryotic Cell Cycle 300
Feature Investigation: Masui and Markert’s Study of Oocyte
Maturation Led to the Identification of Cyclins and Cyclin-
Dependent Kinases 305
15.2 Mitotic Cell Division 306
Genomes & Proteomes Connection: Mitosis in Eukaryotes
Evolved from the Binary Fission That Occurs in Prokaryotic
Cells 310
15.3 Meiosis and Sexual Reproduction 311
15.4 Variation in Chromosome Structure
and Number 318
Chapter 16
Simple Patterns of Inheritance 325
16.1 Mendel’s Laws of Inheritance 326
16.2 The Chromosome Theory of Inheritance 331
16.3 Pedigree Analysis of Human Traits 334
16.4 Sex Chromosomes and X-Linked Inheritance
Patterns 335
Feature Investigation: Morgan’s Experiments Showed a
Correlation Between a Genetic Trait and the Inheritance
of a Sex Chromosome in Drosophila 337
16.5 Variations in Inheritance Patterns and Their Molecular
Basis 339
Genomes & Proteomes Connection: Recessive
Alleles That Cause Diseases May Have Multiple Effects
on Phenotype 340
16.6 Genetics and Probability 343
Chapter 17
Complex Patterns of Inheritance 347
17.1 Gene Interaction 348
17.2 Genes on the Same Chromosome: Linkage,
Recombination, and Mapping 350
Feature Investigation: Bateson and Punnett’s Cross of
Sweet Peas Showed That Genes Do Not Always Assort
Independently 350
17.3 Extranuclear Inheritance: Organelle
Genomes 354
Genomes & Proteomes Connection: Chloroplast and
Mitochondrial Genomes Are Relatively Small, but Contain
Genes That Encode Important Proteins 354
17.4 Epigenetic Inheritance 357
298

CONTENTS xxiii
 Chapter 18
UNIT IV  Evolution
Genetics of Viruses and Bacteria 364
CONTENTS

18.1 Genetic Properties of Viruses 365

18.2 Viral Reproductive Cycles 368
18.3 Viroids and Prions 373
18.4 Genetic Properties of Bacteria 375
18.5 Gene Transfer Between Bacteria 378
Feature Investigation: Lederberg and Tatum’s Work with
E. coli Demonstrated Gene Transfer Between Bacteria and
Led to the Discovery of Conjugation 379
Genomes & Proteomes Connection: Horizontal Gene
Transfer Can Occur Within a Species or Between Chapter 22
Different Species 382
The Origin and History of Life on Earth 445
Chapter 19 22.1 Origin of Life on Earth 446
Feature Investigation: Bartel and Szostak Demonstrated
Developmental Genetics 386 Chemical Evolution in the Laboratory 450
22.2 The Fossil Record 453
19.1 General Themes in Development 386
22.3 History of Life on Earth 455
19.2 Development in Animals 391
Genomes & Proteomes Connection: The Origin of Eukaryotic
Genomes & Proteomes Connection: A Homologous Group of
Cells Involved a Union Between Bacterial and Archaeal
Homeotic Genes Is Found in Nearly All Animals 396
Cells 459
Feature Investigation: Davis, Weintraub, and Lassar Identified
Genes That Promote Muscle Cell Differentiation 400
19.3 Development in Plants 402 Chapter 23

An Introduction to Evolution 467

Chapter 20
23.1 Overview of Evolution 468
Genetic Technology 406 Feature Investigation: The Grants Observed Natural Selection
in Galápagos Finches 471
20.1 Gene Cloning 406
23.2 Evidence of Evolutionary Change 473
20.2 Genomics: Techniques for Studying
23.3 The Molecular Processes That Underlie Evolution 481
Genomes 412
Genomes & Proteomes Connection: Gene Duplications Produce
Genomes & Proteomes Connection: A Microarray Can Identify
Gene Families 482
Which Genes Are Transcribed by a Cell 415
20.3 Biotechnology 416
Feature Investigation: Blaese and Colleagues Performed the
Chapter 24
First Gene Therapy to Treat ADA Deficiency 422
Population Genetics 487
Chapter 21 24.1 Genes in Populations 488
Genomes & Proteomes Connection: Genes Are Usually
Genomes, Proteomes, and Bioinformatics 426 Polymorphic 488
24.2 Natural Selection 491
21.1 Bacterial and Archaeal Genomes 427
24.3 Sexual Selection 495
Feature Investigation: Venter, Smith, and Colleagues
Sequenced the First Genome in 1995 428 Feature Investigation: Seeha and van Alphen Found That
Male Coloration in African Cichlids Is Subject to Female
21.2 Eukaryotic Genomes 430 Choice 497
21.3 Proteomes 435 24.4 Genetic Drift 499
21.4 Bioinformatics 438 24.5 Migration and Nonrandom Mating 501

xxiv CONTENTS
 Chapter 25 27.5 Ecological Roles and Biotechnology Applications 557
Feature Investigation: Dantas and Colleagues Found That Many
Origin of Species and Macroevolution 505 Bacteria Can Break Down and Consume Antibiotics as a Sole

CONTENTS
Carbon Source 557
25.1 Identification of Species 506 Genomes & Proteomes Connection: The Evolution of Bacterial
25.2 Mechanisms of Speciation 511 Pathogens 561
Feature Investigation: Podos Found That an Adaptation
for Feeding May Have Promoted Reproductive Isolation in Chapter 28
Finches 513
25.3 The Pace of Speciation 517 Protists 565
25.4 Evo-Devo: Evolutionary Developmental Biology 518
Genomes & Proteomes Connection: The Study of the Pax6 28.1 An Introduction to Protists 565
Gene Indicates That Different Types of Eyes Evolved from One 28.2 Evolution and Relationships 568
Simple Form 521 Genomes & Proteomes Connection: Genome Sequences Reveal
the Different Evolutionary Pathways of Trichomonas vaginalis
and Giardia intestinalis 570
Chapter 26
28.3 Nutritional and Defensive Adaptations 577
Taxonomy and Systematics 525 Feature Investigation: Cook and Colleagues Demonstrated
That Cellulose Helps Green Algae Avoid Chemical
26.1 Taxonomy 526 Degradation 578
26.2 Phylogenetic Trees 528 28.4 Reproductive Adaptations 580
26.3 Cladistics 532
Feature Investigation: Cooper and Colleagues Compared DNA Chapter 29
from Extinct Flightless Birds and Existing Species to Propose a
New Phylogenetic Tree 536 Plants and the Conquest of Land 588
26.4 Molecular Clocks 538
26.5 Horizontal Gene Transfer 540 29.1 Ancestry and Diversity of Modern Plants 588
Genomes & Proteomes Connection: Due to Horizontal Gene Genomes & Proteomes Connection: Comparison of Plant
Transfer, the “Tree of Life” Is Really a “Web of Life” 541 Genomes Reveals Genetic Changes That Occurred During Plant
Evolution 597
29.2 An Evolutionary History of Land Plants 599
29.3 The Origin and Evolutionary Importance of the Plant
UNIT V  Diversity Embryo 602
Feature Investigation: Browning and Gunning Demonstrated
That Placental Transfer Tissues Facilitate the Movement of
Organic Molecules from Gametophytes to Sporophytes 603
29.4 The Origin and Evolutionary Importance of Leaves
and Seeds 605

Chapter 30

The Evolution and Diversity of Modern

Gymnosperms and Angiosperms 611
30.1 The Evolution and Diversity of Modern
Chapter 27 Gymnosperms 611
30.2 The Evolution and Diversity of Modern
Archaea and Bacteria 545 Angiosperms 617
Genomes & Proteomes Connection: Whole-Genome
27.1 Diversity and Evolution 545 Duplications Influenced the Evolution of Flowering Plants 622
27.2 Structure and Movement 550 Feature Investigation: Hillig and Mahlberg Analyzed Secondary
27.3 Reproduction 554 Metabolites to Explore Species Diversification in the Genus
27.4 Nutrition and Metabolism 556 Cannabis 626

CONTENTS xxv
 30.3 The Role of Coevolution in Angiosperm Chapter 34
Diversification 628
30.4 Human Influences on Angiosperm The Vertebrates 702
CONTENTS

Diversification 630
34.1 Vertebrates: Chordates with a Backbone 702
34.2 Gnathostomes: Jawed Vertebrates 705
Chapter 31
34.3 Tetrapods: Gnathostomes with Four Limbs 710
Fungi 633 Feature Investigation: Davis and Colleagues Provided
a Genetic-Developmental Explanation for Limb Length in
31.1 Evolution and Distinctive Features of Fungi 633 Tetrapods 711
31.2 Fungal Asexual and Sexual Reproduction 637 34.4 Amniotes: Tetrapods with a Desiccation-Resistant
31.3 Diversity of Fungi 639 Egg 714
31.4 Fungal Ecology and Biotechnology 644 34.5 Mammals: Milk-Producing Amniotes 720
Feature Investigation: Márquez and Associates Discovered That Genomes & Proteomes Connection: Comparing the Human
a Three-Partner Symbiosis Allows Plants to Cope with Heat and Chimpanzee Genetic Codes 727
Stress 648
Genomes & Proteomes Connection: Genomic Comparisons of
Forest Fungi Reveal Coevolution with Plants 650 UNIT VI   Flowering Plants
Chapter 32

An Introduction to Animal Diversity 654

32.1 Characteristics of Animals 655
32.2 Animal Classification 656
Genomes & Proteomes Connection: Changes in Hox
Gene Expression Control Body Segment
Specialization 662
32.3 Molecular Views of Animal Diversity 662
Feature Investigation: Aguinaldo and Colleagues Analyzed Chapter 35
SSU rRNA Sequences to Determine the Taxonomic
Relationships of Arthropods to Other Taxa 665
An Introduction to Flowering Plant Form
and Function 734
Chapter 33
35.1 From Seed to Seed—the Life of a Flowering
The Invertebrates 668 Plant 734
35.2 How Plants Grow and Develop 738
33.1 Parazoa: Sponges, the First Multicellular
35.3 The Shoot System: Stem and Leaf
Animals 669
Adaptations 743
33.2 Radiata: Jellyfish and Other Radially Symmetrical
Feature Investigation: Sack and Colleagues Showed
Animals 670 That Palmate Venation Confers Tolerance of Leaf Vein
33.3 Lophotrochozoa: The Flatworms, Rotifers, Bryozoans, Breakage 744
Brachiopods, Mollusks, and Annelids 672 Genomes & Proteomes Connection: Genetic Control of Guard-
Feature Investigation: Fiorito and Scotto’s Experiments Showed Cell Development 747
That Invertebrates Can Exhibit Sophisticated Observational 35.4 Root System Adaptations 751
Learning Behavior 680
33.4 Ecdysozoa: The Nematodes and
Arthropods 683 Chapter 36
Genomes & Proteomes Connection: DNA Barcoding: A New
Tool for Classification 693 Flowering Plants: Behavior 755
33.5 Deuterostomia: The Echinoderms 36.1 Overview of Plant Behavioral Responses 755
and Chordates 693 36.2 Plant Hormones 758

xxvi CONTENTS
 Feature Investigation: An Experiment Performed
by Briggs Revealed the Role of Auxin UNIT VII  Animals
in Phototropism 760

CONTENTS
Genomes & Proteomes Connection: Gibberellin Function Arose
in a Series of Stages During Plant Evolution 762
36.3 Plant Responses to Environmental Stimuli 765

Chapter 37

Flowering Plants: Nutrition 774

37.1 Plant Nutritional Requirements 774
37.2 The Role of Soil in Plant Nutrition 778
Feature Investigation: Hammond and Colleagues
Engineered Smart Plants That Can Communicate Their
Phosphate Needs 782
37.3 Biological Sources of Plant Nutrients 784
Genomes & Proteomes Connection: Development of Legume-
Rhizobia Symbioses 786
Chapter 38
Flowering Plants: Transport 790
38.1 Overview of Plant Transport 790
38.2 Uptake and Movement of Materials at the Cellular
Level 791
38.3 Tissue-Level Transport 794
38.4 Long-Distance Transport 796
Feature Investigation: The Park Team Genetically Engineered
an ABA Receptor Protein to Foster Crop Survival During
Droughts 802
Genomes & Proteomes Connection: Stomatal Responses
to Abscisic Acid Arose Relatively Late in Plant
Evolution 806
Chapter 39
Flowering Plants: Reproduction 811
39.1 An Overview of Flowering Plant Reproduction 811
39.2 Flower Production, Structure, and Development 815
Chapter 40
Animal Bodies and Homeostasis 831
40.1 Organization of Animal Bodies 831
Genomes & Proteomes Connection: Organ Development and
Function Are Controlled by Hox Genes 835
40.2 The Relationship Between Structure and Function 838
40.3 Homeostasis 840
Feature Investigation: Pavlov Demonstrated the Relationship
Between Learning and Feedforward Processes 845
Chapter 41
Neuroscience I:
Cells of the Nervous System 849
41.1 Cellular Components of Nervous Systems 850
41.2 Electrical Properties of Neurons and the Resting
Membrane Potential 852
41.3 Generation and Transmission of Electrical Signals Along
Neurons 856
41.4 Neurons Communicate Electrically or Chemically
at Synapses 860
Feature Investigation: Otto Loewi Discovered
Acetylcholine 865
Genomes & Proteomes Connection: Varied Subunit
Compositions of Neurotransmitter Receptors Allow Precise
Control of Neuronal Regulation 867

Feature Investigation: Liang and Mahadevan Used
Time-Lapse Video and Mathematical Modeling to
Explain How Flowers Bloom 818
39.3 Male and Female Gametophytes and Double
Fertilization 820
39.4 Embryo, Seed, Fruit, and Seedling Development 823
39.5 Asexual Reproduction in Flowering
Plants 827
Genomes & Proteomes Connection: The Evolution of Plantlet
Production in Kalanchoë 827
41.5 Impact on Public Health 868
Chapter 42
Neuroscience II: Evolution, Structure,
and Function of the Nervous System 872
42.1 The Evolution and Development of Nervous
Systems 872
42.2 Structure and Function of the Nervous Systems of
Humans and Other Vertebrates 875

CONTENTS xxvii
 Genomes & Proteomes Connection: Many Genes Have Been Genomes & Proteomes Connection: Evolution and Genetics
Important in the Evolution and Development of the Cerebral Explains Lactose Intolerance 949
Cortex 884 45.5 Neural and Endocrine Control of Digestion 952
CONTENTS

42.3 Cellular Basis of Learning and Memory 884 45.6 Impact on Public Health 953
Feature Investigation: Gaser and Schlaug Showed That the Feature Investigation: Marshall and Warren and Coworkers
Sizes of Certain Brain Structures Differ Between Musicians and Demonstrated a Link Between Bacterial Infection and Ulcers 955
Nonmusicians 888
42.4 Impact on Public Health 889
Chapter 46
Chapter 43 Control of Energy Balance, Metabolic Rate, and
Body Temperature 959
Neuroscience III:
Sensory Systems 893 46.1 Use and Storage of Energy 959
46.2 Regulation of the Absorptive and Postabsorptive
43.1 An Introduction to Sensation 893 States 962
43.2 Mechanoreception 895 Genomes & Proteomes Connection: A Family of GLUT
43.3 Thermoreception and Nociception 901 Proteins Transports Glucose in All Animal Cells 963
43.4 Electromagnetic Reception 902 46.3 Energy Balance and Metabolic Rate 965
43.5 Photoreception 902 Feature Investigation: Coleman Revealed a Satiety Factor in
Genomes & Proteomes Connection: Color Vision Is an Ancient Mammals 968
Adaptation in Animals 906 46.4 Regulation of Body Temperature 970
43.6 Chemoreception 910 46.5 Impact on Public Health 974
Feature Investigation: Buck and Axel Discovered a Family
of Olfactory Receptor Proteins That Bind Specific Odor
Molecules 912
Chapter 47
43.7 Impact on Public Health 915
Circulatory Systems 977
Chapter 44 47.1 Types of Circulatory Systems 977
Genomes & Proteomes Connection: A Four-Chambered Heart
Muscular-Skeletal Systems and Locomotion 919 Evolved from Simple Contractile Tubes 980
47.2 The Composition of Blood 981
44.1 Types of Animal Skeletons 919 47.3 The Vertebrate Heart and Its Function 983
44.2 Skeletal Muscle Structure and the Mechanism 47.4 Blood Vessels 987
of Force Generation 921
47.5 Relationship Among Blood Pressure, Blood Flow,
Genomes & Proteomes Connection: Did an Ancient Mutation in and Resistance 989
Myosin Play a Role in the Development of the Human Brain? 924
Feature Investigation: Furchgott Discovered a Vasodilatory
44.3 Types of Skeletal Muscle Fibers and Their Factor Produced by Endothelial Cells 991
Functions 928
47.6 Adaptive Functions of Closed Circulatory Systems 994
Feature Investigation: Evans and Colleagues Activated a Gene
to Produce “Marathon Mice” 930
47.7 Impact on Public Health 995
44.4 Animal Locomotion 932
44.5 Impact on Public Health 934 Chapter 48

Respiratory Systems 999

Chapter 45
48.1 Physical Properties of Gases 999
Nutrition and Animal Digestive Systems 938 48.2 Types of Respiratory Systems 1002
45.1 Animal Nutrition 938 48.3 Structure and Function of the Mammalian Respiratory
System 1006
45.2 General Principles of Digestion and Absorption
of Food 940 Feature Investigation: Fujiwara and Colleagues Demonstrated
the Effectiveness of Administering Surfactant to Newborns with
45.3 Overview of Vertebrate Digestive Systems 943
RDS 1010
45.4 Mechanisms of Digestion and Absorption in
48.4 Mechanisms of Oxygen Transport in Blood 1013
Vertebrates 948

xxviii CONTENTS
 Genomes & Proteomes Connection: Hemoglobin First Evolved Chapter 52
Over 500 mya 1016
48.5 Control of Ventilation 1017 Animal Development 1089

CONTENTS
48.6 Adaptations to Extreme Conditions 1018
52.1 Principles of Embryonic Development 1089
48.7 Impact on Public Health 1019
52.2 General Events of Embryonic Development 1090
52.3 Control of Cell Differentiation and Morphogenesis
Chapter 49 During Animal Development 1099
Genomes & Proteomes Connection: Groups of Embryonic Cells
Excretory Systems and the Homeostasis Can Produce Specific Body Structures Even When Transplanted
of Internal Fluids 1023 into Different Animals 1100
49.1 Principles of Homeostasis of Internal Fluids 1023 Feature Investigation: Harland and Coworkers Identified Genes
Expressed Specifically in the Organizer 1101
Feature Investigation: Cade and Colleagues Discovered Why
Athletes’ Performances Wane on Hot Days 1027 52.4 Impact on Public Health 1103
49.2 Excretory Systems in Different Animal Groups 1030
49.3 Structure and Function of the Mammalian Kidney 1033 Chapter 53
Genomes & Proteomes Connection: Aquaporins Comprise a
Large Family of Proteins That Are Found in All Animals 1039 Immune Systems 1107
49.4 Impact on Public Health 1040 53.1 Types of Pathogens 1108
53.2 Innate Immunity 1108
Chapter 50 Feature Investigation: Lemaitre and Colleagues Identify an
Immune Function for Toll Protein in Drosophila 1111
Endocrine Systems 1044 53.3 Acquired Immunity 1113
50.1 Types of Hormones and Their Mechanisms of Genomes & Proteomes Connection: Recombination and
Hypermutation Produce an Enormous Number of Different
Action 1045
Immunoglobulin Proteins 1118
50.2 Links Between the Endocrine and Nervous Systems 1048
53.4 Impact on Public Health 1125
50.3 Hormonal Control of Metabolism and Energy
Balance 1050
Feature Investigation: Banting, Best, MacLeod, and Collip Were
the First to Isolate Active Insulin 1056 UNIT VIII  Ecology
50.4 Hormonal Control of Mineral Balance 1058
Genomes & Proteomes Connection: Hormones and Receptors
Evolved as Tightly Integrated Molecular Systems 1061
50.5 Hormonal Control of Growth and Development 1062
50.6 Hormonal Control of Reproduction 1064
50.7 Impact on Public Health 1065

Chapter 51

Animal Reproduction 1069

51.1 Overview of Asexual and Sexual Reproduction 1069 Chapter 54
Feature Investigation: Paland and Lynch Provided Evidence
That Sexual Reproduction May Promote the Elimination of An Introduction to Ecology and Biomes 1131
Harmful Mutations in Populations 1071
54.1 The Scale of Ecology 1132
51.2 Gametogenesis and Fertilization 1072
Feature Investigation: Callaway and Aschehoug’s Experiments
51.3 Mammalian Reproductive Structure and Function 1075
Showed That the Secretion of Chemicals Gives Invasive Plants
51.4 Pregnancy and Birth in Mammals 1081 a Competitive Edge Over Native Species 1132
Genomes & Proteomes Connection: The Evolution of the 54.2 Ecological Methods 1134
Globin Gene Family Has Been Important for Internal Gestation
in Mammals 1083
54.3 The Environment’s Effect on the Distribution of
Organisms 1137
51.5 Impact on Public Health 1085

CONTENTS xxix
 Genomes & Proteomes Connection: Temperature Tolerance Chapter 58
May Be Manipulated by Genetic Engineering 1140
54.4 Climate and Its Relationship to Biological Community Ecology 1223
CONTENTS

Communities 1144
54.5 Major Biomes 1147 58.1 Differing Views of Communities 1223
54.6 Continental Drift and Biogeography 1157 58.2 Patterns of Species Richness 1225
58.3 Calculating Species Diversity 1227
Genomes & Proteomes Connection: Metagenomics May Be
Chapter 55
Used to Measure Species Diversity 1229

Behavioral Ecology 1162 58.4 Species Diversity and Community Stability 1230
58.5 Succession: Community Change 1231
55.1 The Influence of Genetics and Learning on 58.6 Island Biogeography 1234
Behavior 1163
Feature Investigation: Simberloff and Wilson’s Experiments
Genomes & Proteomes Connection: Some Behavior Results Tested the Predictions of the Equilibrium Model of Island
from Simple Genetic Influences 1163 Biogeography 1237
55.2 Local Movement and Long-Range Migration 1167
Feature Investigation: Tinbergen’s Experiments Showed That Chapter 59
Digger Wasps Use Landmarks to Find Their Nests 1167
55.3 Foraging Behavior 1171 Ecosystem Ecology 1241
55.4 Communication 1173
59.1 Food Webs and Energy Flow 1242
55.5 Living in Groups 1175
Genomes & Proteomes Connection: Using Genetically
55.6 Altruism 1176 Engineered Plants to Remove Pollutants 1248
55.7 Mating Systems 1179 59.2 Biomass Production in Ecosystems 1248
59.3 Biogeochemical Cycles 1251
Chapter 56 Feature Investigation: Stiling and Drake’s Experiments with
Elevated CO2 Showed an Increase in Plant Growth but a
Ecology 1183 Decrease in Herbivory 1254
56.1 Understanding Populations 1183
56.2 Demography 1188 Chapter 60
Feature Investigation: Murie’s Collections of Dall Mountain
Sheep Skulls Permitted the Accurate Construction of Life Biodiversity and Conservation Biology 1260
Tables 1189
60.1 What Is Biodiversity? 1261
56.3 How Populations Grow 1192
60.2 Why Conserve Biodiversity? 1261
Genomes & Proteomes Connection: Hexaploidy Increases the
Feature Investigation: Ecotron Experiments Showed
Growth of Coast Redwood Trees 1196
the Relationship Between Biodiversity and Ecosystem
56.4 Human Population Growth 1197 Function 1263
60.3 The Causes of Extinction and Loss of Biodiversity 1266
Chapter 57 60.4 Conservation Strategies 1270
Genomes & Proteomes Connection: Can Cloning Save
Species Interactions 1202 Endangered Species? 1277
57.1 Competition 1203 Appendix A: Periodic Table of the Elements A-1
Feature Investigation: Connell’s Experiments with Barnacle Appendix B: Answer Key A-2
Species Revealed Each Species’ Fundamental and Realized Glossary G-1
Niches 1206
Credits C-1
57.2 Predation, Herbivory, and Parasitism 1209
Index I-1
Genomes & Proteomes Connection: Transgenic Plants May Be
Used in the Fight Against Plant Diseases 1216
57.3 Mutualism and Commensalism 1217
57.4 Bottom-Up and Top-Down Control 1219

xxx CONTENTS
 CHAPTER OUTLINE
1.1  rinciples of Biology and the Levels of Biological
P An Introduction
Organization
1.2 Unity and Diversity of Life
to Biology
1.3 Biology as a Scientific Discipline

1
Summary of Key Concepts
Assess and Discuss

B iology is the study of life. The diverse forms of life

found on Earth provide biologists with an amazing array
of organisms to study. In many cases, the investigation of living
things leads to discoveries that no one would have imagined. For
example, researchers determined that the venom from certain
poisonous snakes contains a chemical that lowers blood pressure
in humans. By analyzing that chemical, scientists developed drugs
to treat high blood pressure (Figure 1.1).
Biologists have discovered that plants can communicate
with each other. For example, the beautiful umbrella thorn aca-
cia (Vachellia tortillis), shown in Figure 1.2, emits volatile organic
molecules when it is attacked by herbivores. These molecules
warn other nearby acacia trees that herbivores are in the area,
and those trees release toxins to protect themselves.
In the 20th century, biologists studied soil bacteria that natu- Gypsy moth (Lymantria dispar). The gypsy moth progresses
rally produce “chemical weapons” to kill competing bacteria in through different stages, including a caterpillar, to become an
their native environment. These chemicals have been character- adult moth. As discussed later in this chapter, the caterpillar can
ized and used to develop antibiotics such as streptomycin to treat be infected with a virus, which alters its behavior into that of
bacterial infections (Figure 1.3). a “zombie.”

C
N
O O OCH2CH3
CH2COOH
ACE inhibitor (Lotensin)

Figure 1.1 The Brazilian arrowhead viper and inhibitors of high

blood pressure. Derivatives of a chemical, called an angiotensin-
converting enzyme (ACE) inhibitor, are found in the venom of the Figure 1.2 Plant communication. If attacked by herbivores, this
Brazilian arrowhead viper and are commonly used to treat high acacia tree will emit molecules that will warn other acacia trees in
blood pressure. the area.
 To make new discoveries, biologists view life from many differ-
ent perspectives. What is the composition of living things? How is
NH2
H
life organized? How do organisms reproduce? Sometimes the ques-
C
N NH tions posed by biologists are fundamental and even philosophical in
HO OH
NH nature. How did living organisms originate? Can we live forever? What
OH
C H is the physical basis for memory? Can we save endangered species?
N N
O O H H
Future biologists will continue to make important advances.
H3C Biologists are scientific explorers looking for answers to some of
H C
OH
O H life’s most enduring mysteries. Unraveling these mysteries ­presents
N CH3
O O an exciting challenge to the best and brightest minds. The rewards of
a career in biology include the excitement of forging into uncharted
CH2OH OH
OH territory, the thrill of making discoveries that can improve the health
Streptomycin and lives of people, and the impact of trying to preserve the environ-
ment and protect endangered species. For these and many other
Figure 1.3 The soil bacterium Streptomyces griseus, which compelling reasons, students seeking challenging and rewarding
naturally produces the antibiotic streptomycin, kills competing careers may wish to choose biology as a lifelong pursuit.
bacteria in the soil. Doctors administer streptomycin to treat In this chapter, we will begin our survey of biology by examin-
bacterial infections.
ing the basic principles that underlie the characteristics of all living
organisms and the fields of biology. We will then take a deeper look
These are but a few of the many discoveries that make biol- at the process of evolution and how it has led to the development
ogy an intriguing discipline. The study of life not only reveals the of genomes—the entire genetic compositions of living organisms—
fascinating characteristics of living species but also leads to the which explains the unity and diversity that we observe among living
development of medicines and research tools that benefit the lives species. Finally, we will explore the general approaches that scien-
of people. tists follow when making new discoveries.

1.1 
Principles of Biology and
the Levels of Biological
Organization
Learning Outcomes:
1. Describe the principles of biology.
2. Explain how life can be viewed at different levels of biological
complexity.
Because biology is the study of life, a good way to begin a biology
textbook is to distinguish living organisms from nonliving objects. At
first, the distinction might seem obvious. A person is alive, but a rock
is not. However, the distinction between living and nonliving may
seem less obvious when we consider microscopic entities. Is a bac-
terium alive? What about a virus or a chromosome? In this section,
we will examine the principles that underlie the characteristics of all
forms of life and explore other broad principles in biology. We will
then consider the levels of organization that biologists study, ranging
from atoms and small molecules to very large geographical areas.
The Study of Life Has Revealed a Set
of Unifying Principles
Biologists have studied many different species and learned that a set of
principles applies to all fields of biology. Twelve broad principles are
described in Figure 1.4. The first eight principles are often used as crite-
ria to define the basic features of life. You will see these twelve principles
at many points as you progress through this textbook, and they are indi-
cated with the icon. In particular, we will draw your attention to them
at the beginning of each unit, and we will refer to them within particular
figures in Chapters 3 through 60. It should be noted that the principles of
biology are also governed by the laws of chemistry and physics, which
are discussed in Chapters 2, 3, and 6.
Principle 1: Cells are the simplest units of life. The term
organism can be applied to all living things. Organisms maintain an
internal order that is separated from the environment (Figure 1.4a).
The simplest unit of such organization is the cell, which we will exam-
ine in Unit II. One of the foundations of biology is the cell theory,
which states that (1) all organisms are composed of cells, (2) cells are
the smallest units of life, and (3) new cells come from pre-existing
cells via cell division. Unicellular organisms are composed of one cell,
whereas multicellular organisms such as plants and animals contain
many cells. In plants and animals, each cell has an internal order, and
the cells within the organism have specific arrangements and functions.
Principle 2: Living organisms use energy. The maintenance of
organization requires energy. Therefore, all living organisms acquire
energy from the environment and use that energy to maintain their
internal order. Cells carry out a variety of chemical reactions that
are responsible for the breakdown of nutrients. Such reactions often
release energy in a process called respiration. The energy may be
used to synthesize the components that make up individual cells
and living organisms. Chemical reactions involved with the break-
down and synthesis of cellular molecules are collectively known
as metabolism. Plants, algae, and certain bacteria can directly
 (a) Cells are the simplest (g) Populations of organisms
units of life: evolve from one generation
Organisms maintain an to the next:
internal order. The simplest Populations of organisms
unit of organization is the cell. change over the course of
Yeast cells are shown here. many generations. Evolution
results in traits that promote
13.7 m
survival and reproductive
success.

(b) Living organisms use energy: (h) All species (past and
Organisms need energy to present) are related by an
maintain internal order. evolutionary history:
These algae harness light The three mammal species
energy via photosynthesis. shown here share a common
Energy is used in chemical ancestor, which was also a
reactions collectively known mammal.
as metabolism.

(c) Living organisms interact (i) Structure determines

with their environment: function:
Organisms respond to In the example seen here,
environmental changes. webbed feet (on ducks)
These plants are growing function as paddles for
toward the light. swimming. Nonwebbed feet
(on chickens) function better
for walking on the ground.

(d) Living organisms maintain ( j) New properties of life

homeostasis: emerge from complex
Organisms regulate their cells interactions:
and bodies, maintaining relatively Our ability to see is an
stable internal conditions, a process emergent property due to
called homeostasis. For example, interactions among many
this bird maintains its internal body types of cells in the eye and
temperature on a cold day. neurons that send signals to
the brain.

(e) Living organisms grow and (k) Biology is an

develop: experimental science:
Growth produces more The discoveries of
or larger cells, whereas biology are made via
development produces experimentation, which
organisms with a defined leads to theories and
set of characteristics. biological principles.

(f) The genetic material provides (l) Biology affects our society:
a blueprint for reproduction: Many discoveries in biology
To sustain life over many have had major effects on
generations, organisms must our society. For example,
reproduce. Due to the biologists developed Bt-corn,
transmission of genetic which is resistant to insect
material, offspring tend to pests and widely planted by
have traits like their parents. farmers.

Figure 1.4 Twelve principles of biology. The first eight principles are often used as criteria for defining the basic features of life. Note: The
twelve principles described here were modeled after the themes and core competencies described in Vision and Change in Undergraduate
Biology, a report that was published in 2009 and organized by the American Association for the Advancement of Science. Vision and Change
proposed five themes. We have divided them into ten principles to make them more accessible to beginning biology students. The five Vision and
Change themes are related to our principles in the following manner: 1. Evolution (principles g and h); 2. Structure and Function (principle i);
3. Information Flow, Exchange, and Storage (principles e and f); 4. Pathways and Transformations of Energy and Matter (principles b, c, and d);
5. Systems (principles a and j). The last two principles are modeled after two core competencies described in Vision and Change: Ability to Apply
the Process of Science (principle k) and Ability to Understand the Relationship between Science and Society (principle l).
BioConnections:   Look ahead to Figure 52.11. Which of these principles is this figure emphasizing?

harness light energy to produce their own nutrients in the process of Principle 3: Living organisms interact with their environment.
photosynthesis (Figure 1.4b). They are the primary producers of food To survive, living organisms must be able to interact with their envi-
on Earth. In contrast, some organisms, such as animals and fungi, are ronment, which includes other organisms they may encounter. All
consumers—they must use other organisms as food to obtain energy. organisms must respond to environmental changes. In the winter,
4 CHAPTER 1
many species of mammals develop a thicker coat of fur that protects
them from the cold temperatures. Plants respond to changes in the
angle of the sun. If you place a plant in a window, it will grow toward
the light (Figure 1.4c).
Principle 4: Living organisms maintain homeostasis. Although
life is a dynamic process, living cells and organisms regulate their
cells and bodies to maintain relatively stable internal conditions, a
process called homeostasis (from the Greek, meaning to stay the
same). The degree to which homeostasis is achieved varies among
different organisms. For example, most mammals and birds maintain
a relatively stable body temperature in spite of changing environ-
mental temperatures (Figure 1.4d), whereas reptiles and amphibians
tolerate a wider fluctuation in body temperature. By comparison, all
organisms continually regulate their cellular metabolism so nutrient
molecules are used at an appropriate rate and new cellular compo-
nents are synthesized when they are needed.
Principle 5: Living organisms grow and develop. All living
organisms grow and develop. Growth produces more or larger cells.
In plants and animals, a fertilized egg undergoes multiple cell divi-
sions to develop into a mature organism with many cells. Among uni-
cellular organisms such as bacteria, new cells are relatively small, and
they increase in volume by the synthesis of additional cellular compo-
nents. Development is a series of changes in the state of a cell, tissue,
organ, or organism, eventually resulting in organisms with a defined
set of characteristics (Figure 1.4e).
Principle 6: The genetic material provides a blueprint for
reproduction. All living organisms have a finite life span. To
sustain life, organisms must reproduce, or generate offspring
(Figure 1.4f  ). A key feature of reproduction is that offspring tend
to have characteristics that greatly resemble those of their parent(s).
How is this possible? All living organisms contain genetic material
composed of DNA (deoxyribonucleic acid), which provides a
blueprint for the organization, development, and function of living
things. During reproduction, a copy of this blueprint is transmitted
from parent to offspring. DNA is heritable, which means that
offspring inherit DNA from their parents.
As discussed in Unit III, genes, which are segments of DNA,
govern the characteristics, or traits, of organisms. Most genes are
transcribed into a type of RNA (ribonucleic acid) molecule called
messenger RNA (mRNA) that is then translated into a polypeptide
with a specific amino acid sequence. A protein is composed of one
or more polypeptides. The structures and functions of proteins are
largely responsible for the traits of living organisms.
Principle 7: Populations of organisms evolve from one gener-
ation to the next. The first six characteristics of life, which we
have just considered, apply to individual organisms over the short
run. Over the long run, another universal characteristic of life is
biological evolution, or simply evolution, which refers to a herita-
ble change in a population of organisms from generation to genera-
tion. As a result of evolution, populations become better adapted to
the environment in which they live. For example, the long snout of
an anteater is an adaptation that enhances its ability to obtain food,
namely ants, from hard-to-reach places (Figure 1.4g). Over the course
of many generations, the fossil record indicates that the long snout
occurred via biological evolution in which modern anteaters evolved
from populations of organisms with shorter snouts.
Principle 8: All species (past and present) are related by an
evolutionary history. Principle 7 considers evolution as an ongo-
ing process that happens from one generation to the next. Evidence
from a variety of sources, including the fossil record and DNA
sequences, also indicates that all organisms on Earth share a common
ancestry. For example, the three species of mammals shown in Figure
1.4h shared a common ancestor in the past, which was also a mam-
mal. We will discuss evolutionary relationships later in Section 1.2
and more thoroughly in Units IV and V.
As described later in this chapter, biologists often view evolution
within the context of genomes and proteomes. The term genome refers
to the complete genetic composition of an organism or species. Because
most genes encode proteins, these genetic changes are often associated
with changes in the proteome, which is the complete protein composi-
tion of a cell or organism. By studying how evolution affects genomes
and proteomes, biologists can better understand how the changes that
occur during evolution affect the characteristics of species. Because
evolution is a core unifying principle in biology, we will draw your
attention to it in Chapters 4 through 60 by including a brief topic that
we call “Genomes & Proteomes Connection.” This topic connects the
principle of evolution to the subject matter in each chapter.
Principle 9: Structure determines function. In addition to the
preceding eight characteristics of life, biologists have identified other
principles that are important in all fields of biology. The principle
“structure determines function” pertains to very tiny biological mol-
ecules and to very large biological structures. For example, at the
microscopic level, a cellular protein called actin naturally assembles
into structures that are long filaments. The function of these filaments
is to provide support and shape to cells. At the macroscopic level,
let’s consider the feet of different birds (Figure 1.4i). Aquatic birds
have webbed feet that function as paddles for swimming. By com-
parison, the feet of nonaquatic birds are not webbed and are better
adapted for grasping food, perching on branches, and running along
the ground. In this case, the structure of a bird’s feet, webbed versus
nonwebbed, is a critical feature that affects their function.
Principle 10: New properties of life emerge from complex
interactions. In biology, when individual components in an organ-
ism interact with each other or with the external environment to create
novel structures and functions, the resulting characteristics are called
emergent properties. For example, the human eye is composed of
many different types of cells that are organized to sense incoming
light and transmit signals to the brain (Figure 1.4j). Our ability to
see is an emergent property of this complex arrangement of different
cell types. As discussed later in this chapter, biologists use the term
systems biology to describe the study of how new properties of life
arise by complex interactions of its components.
Principle 11: Biology is an experimental science. Biology is an
inquiry process. Biologists are curious about the characteristics of living
organisms and ask questions about those characteristics. For example, a
cell biologist may wonder why a cell produces a specific protein when it
is confronted with high temperature. An ecologist may ask herself why
Another random document with
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 DANCE ON STILTS AT THE GIRLS’ UNYAGO, NIUCHI

Newala, too, suffers from the distance of its water-supply—at least

the Newala of to-day does; there was once another Newala in a lovely
valley at the foot of the plateau. I visited it and found scarcely a trace
of houses, only a Christian cemetery, with the graves of several
missionaries and their converts, remaining as a monument of its
former glories. But the surroundings are wonderfully beautiful. A
thick grove of splendid mango-trees closes in the weather-worn
crosses and headstones; behind them, combining the useful and the
agreeable, is a whole plantation of lemon-trees covered with ripe
fruit; not the small African kind, but a much larger and also juicier
imported variety, which drops into the hands of the passing traveller,
without calling for any exertion on his part. Old Newala is now under
the jurisdiction of the native pastor, Daudi, at Chingulungulu, who,
as I am on very friendly terms with him, allows me, as a matter of
course, the use of this lemon-grove during my stay at Newala.
 FEET MUTILATED BY THE RAVAGES OF THE “JIGGER”
(Sarcopsylla penetrans)

The water-supply of New Newala is in the bottom of the valley,

some 1,600 feet lower down. The way is not only long and fatiguing,
but the water, when we get it, is thoroughly bad. We are suffering not
only from this, but from the fact that the arrangements at Newala are
nothing short of luxurious. We have a separate kitchen—a hut built
against the boma palisade on the right of the baraza, the interior of
which is not visible from our usual position. Our two cooks were not
long in finding this out, and they consequently do—or rather neglect
to do—what they please. In any case they do not seem to be very
particular about the boiling of our drinking-water—at least I can
attribute to no other cause certain attacks of a dysenteric nature,
from which both Knudsen and I have suffered for some time. If a
man like Omari has to be left unwatched for a moment, he is capable
of anything. Besides this complaint, we are inconvenienced by the
state of our nails, which have become as hard as glass, and crack on
the slightest provocation, and I have the additional infliction of
pimples all over me. As if all this were not enough, we have also, for
the last week been waging war against the jigger, who has found his
Eldorado in the hot sand of the Makonde plateau. Our men are seen
all day long—whenever their chronic colds and the dysentery likewise
raging among them permit—occupied in removing this scourge of
Africa from their feet and trying to prevent the disastrous
consequences of its presence. It is quite common to see natives of
this place with one or two toes missing; many have lost all their toes,
or even the whole front part of the foot, so that a well-formed leg
ends in a shapeless stump. These ravages are caused by the female of
Sarcopsylla penetrans, which bores its way under the skin and there
develops an egg-sac the size of a pea. In all books on the subject, it is
stated that one’s attention is called to the presence of this parasite by
an intolerable itching. This agrees very well with my experience, so
far as the softer parts of the sole, the spaces between and under the
toes, and the side of the foot are concerned, but if the creature
penetrates through the harder parts of the heel or ball of the foot, it
may escape even the most careful search till it has reached maturity.
Then there is no time to be lost, if the horrible ulceration, of which
we see cases by the dozen every day, is to be prevented. It is much
easier, by the way, to discover the insect on the white skin of a
European than on that of a native, on which the dark speck scarcely
shows. The four or five jiggers which, in spite of the fact that I
constantly wore high laced boots, chose my feet to settle in, were
taken out for me by the all-accomplished Knudsen, after which I
thought it advisable to wash out the cavities with corrosive
sublimate. The natives have a different sort of disinfectant—they fill
the hole with scraped roots. In a tiny Makua village on the slope of
the plateau south of Newala, we saw an old woman who had filled all
the spaces under her toe-nails with powdered roots by way of
prophylactic treatment. What will be the result, if any, who can say?
The rest of the many trifling ills which trouble our existence are
really more comic than serious. In the absence of anything else to
smoke, Knudsen and I at last opened a box of cigars procured from
the Indian store-keeper at Lindi, and tried them, with the most
distressing results. Whether they contain opium or some other
narcotic, neither of us can say, but after the tenth puff we were both
“off,” three-quarters stupefied and unspeakably wretched. Slowly we
recovered—and what happened next? Half-an-hour later we were
once more smoking these poisonous concoctions—so insatiable is the
craving for tobacco in the tropics.
Even my present attacks of fever scarcely deserve to be taken
seriously. I have had no less than three here at Newala, all of which
have run their course in an incredibly short time. In the early
afternoon, I am busy with my old natives, asking questions and
making notes. The strong midday coffee has stimulated my spirits to
an extraordinary degree, the brain is active and vigorous, and work
progresses rapidly, while a pleasant warmth pervades the whole
body. Suddenly this gives place to a violent chill, forcing me to put on
my overcoat, though it is only half-past three and the afternoon sun
is at its hottest. Now the brain no longer works with such acuteness
and logical precision; more especially does it fail me in trying to
establish the syntax of the difficult Makua language on which I have
ventured, as if I had not enough to do without it. Under the
circumstances it seems advisable to take my temperature, and I do
so, to save trouble, without leaving my seat, and while going on with
my work. On examination, I find it to be 101·48°. My tutors are
abruptly dismissed and my bed set up in the baraza; a few minutes
later I am in it and treating myself internally with hot water and
lemon-juice.
Three hours later, the thermometer marks nearly 104°, and I make
them carry me back into the tent, bed and all, as I am now perspiring
heavily, and exposure to the cold wind just beginning to blow might
mean a fatal chill. I lie still for a little while, and then find, to my
great relief, that the temperature is not rising, but rather falling. This
is about 7.30 p.m. At 8 p.m. I find, to my unbounded astonishment,
that it has fallen below 98·6°, and I feel perfectly well. I read for an
hour or two, and could very well enjoy a smoke, if I had the
wherewithal—Indian cigars being out of the question.
Having no medical training, I am at a loss to account for this state
of things. It is impossible that these transitory attacks of high fever
should be malarial; it seems more probable that they are due to a
kind of sunstroke. On consulting my note-book, I become more and
more inclined to think this is the case, for these attacks regularly
follow extreme fatigue and long exposure to strong sunshine. They at
least have the advantage of being only short interruptions to my
work, as on the following morning I am always quite fresh and fit.
My treasure of a cook is suffering from an enormous hydrocele which
makes it difficult for him to get up, and Moritz is obliged to keep in
the dark on account of his inflamed eyes. Knudsen’s cook, a raw boy
from somewhere in the bush, knows still less of cooking than Omari;
consequently Nils Knudsen himself has been promoted to the vacant
post. Finding that we had come to the end of our supplies, he began
by sending to Chingulungulu for the four sucking-pigs which we had
bought from Matola and temporarily left in his charge; and when
they came up, neatly packed in a large crate, he callously slaughtered
the biggest of them. The first joint we were thoughtless enough to
entrust for roasting to Knudsen’s mshenzi cook, and it was
consequently uneatable; but we made the rest of the animal into a
jelly which we ate with great relish after weeks of underfeeding,
consuming incredible helpings of it at both midday and evening
meals. The only drawback is a certain want of variety in the tinned
vegetables. Dr. Jäger, to whom the Geographical Commission
entrusted the provisioning of the expeditions—mine as well as his
own—because he had more time on his hands than the rest of us,
seems to have laid in a huge stock of Teltow turnips,[46] an article of
food which is all very well for occasional use, but which quickly palls
when set before one every day; and we seem to have no other tins
left. There is no help for it—we must put up with the turnips; but I
am certain that, once I am home again, I shall not touch them for ten
years to come.
Amid all these minor evils, which, after all, go to make up the
genuine flavour of Africa, there is at least one cheering touch:
Knudsen has, with the dexterity of a skilled mechanic, repaired my 9
× 12 cm. camera, at least so far that I can use it with a little care.
How, in the absence of finger-nails, he was able to accomplish such a
ticklish piece of work, having no tool but a clumsy screw-driver for
taking to pieces and putting together again the complicated
mechanism of the instantaneous shutter, is still a mystery to me; but
he did it successfully. The loss of his finger-nails shows him in a light
contrasting curiously enough with the intelligence evinced by the
above operation; though, after all, it is scarcely surprising after his
ten years’ residence in the bush. One day, at Lindi, he had occasion
to wash a dog, which must have been in need of very thorough
cleansing, for the bottle handed to our friend for the purpose had an
extremely strong smell. Having performed his task in the most
conscientious manner, he perceived with some surprise that the dog
did not appear much the better for it, and was further surprised by
finding his own nails ulcerating away in the course of the next few
days. “How was I to know that carbolic acid has to be diluted?” he
mutters indignantly, from time to time, with a troubled gaze at his
mutilated finger-tips.
 Since we came to Newala we have been making excursions in all
directions through the surrounding country, in accordance with old
habit, and also because the akida Sefu did not get together the tribal
elders from whom I wanted information so speedily as he had
promised. There is, however, no harm done, as, even if seen only
from the outside, the country and people are interesting enough.
The Makonde plateau is like a large rectangular table rounded off
at the corners. Measured from the Indian Ocean to Newala, it is
about seventy-five miles long, and between the Rovuma and the
Lukuledi it averages fifty miles in breadth, so that its superficial area
is about two-thirds of that of the kingdom of Saxony. The surface,
however, is not level, but uniformly inclined from its south-western
edge to the ocean. From the upper edge, on which Newala lies, the
eye ranges for many miles east and north-east, without encountering
any obstacle, over the Makonde bush. It is a green sea, from which
here and there thick clouds of smoke rise, to show that it, too, is
inhabited by men who carry on their tillage like so many other
primitive peoples, by cutting down and burning the bush, and
manuring with the ashes. Even in the radiant light of a tropical day
such a fire is a grand sight.
Much less effective is the impression produced just now by the
great western plain as seen from the edge of the plateau. As often as
time permits, I stroll along this edge, sometimes in one direction,
sometimes in another, in the hope of finding the air clear enough to
let me enjoy the view; but I have always been disappointed.
Wherever one looks, clouds of smoke rise from the burning bush,
and the air is full of smoke and vapour. It is a pity, for under more
favourable circumstances the panorama of the whole country up to
the distant Majeje hills must be truly magnificent. It is of little use
taking photographs now, and an outline sketch gives a very poor idea
of the scenery. In one of these excursions I went out of my way to
make a personal attempt on the Makonde bush. The present edge of
the plateau is the result of a far-reaching process of destruction
through erosion and denudation. The Makonde strata are
everywhere cut into by ravines, which, though short, are hundreds of
yards in depth. In consequence of the loose stratification of these
beds, not only are the walls of these ravines nearly vertical, but their
upper end is closed by an equally steep escarpment, so that the
western edge of the Makonde plateau is hemmed in by a series of
deep, basin-like valleys. In order to get from one side of such a ravine
to the other, I cut my way through the bush with a dozen of my men.
It was a very open part, with more grass than scrub, but even so the
short stretch of less than two hundred yards was very hard work; at
the end of it the men’s calicoes were in rags and they themselves
bleeding from hundreds of scratches, while even our strong khaki
suits had not escaped scatheless.

NATIVE PATH THROUGH THE MAKONDE BUSH, NEAR

MAHUTA

I see increasing reason to believe that the view formed some time
back as to the origin of the Makonde bush is the correct one. I have
no doubt that it is not a natural product, but the result of human
occupation. Those parts of the high country where man—as a very
slight amount of practice enables the eye to perceive at once—has not
yet penetrated with axe and hoe, are still occupied by a splendid
timber forest quite able to sustain a comparison with our mixed
forests in Germany. But wherever man has once built his hut or tilled
his field, this horrible bush springs up. Every phase of this process
may be seen in the course of a couple of hours’ walk along the main
road. From the bush to right or left, one hears the sound of the axe—
not from one spot only, but from several directions at once. A few
steps further on, we can see what is taking place. The brush has been
cut down and piled up in heaps to the height of a yard or more,
between which the trunks of the large trees stand up like the last
pillars of a magnificent ruined building. These, too, present a
melancholy spectacle: the destructive Makonde have ringed them—
cut a broad strip of bark all round to ensure their dying off—and also
piled up pyramids of brush round them. Father and son, mother and
son-in-law, are chopping away perseveringly in the background—too
busy, almost, to look round at the white stranger, who usually excites
so much interest. If you pass by the same place a week later, the piles
of brushwood have disappeared and a thick layer of ashes has taken
the place of the green forest. The large trees stretch their
smouldering trunks and branches in dumb accusation to heaven—if
they have not already fallen and been more or less reduced to ashes,
perhaps only showing as a white stripe on the dark ground.
This work of destruction is carried out by the Makonde alike on the
virgin forest and on the bush which has sprung up on sites already
cultivated and deserted. In the second case they are saved the trouble
of burning the large trees, these being entirely absent in the
secondary bush.
After burning this piece of forest ground and loosening it with the
hoe, the native sows his corn and plants his vegetables. All over the
country, he goes in for bed-culture, which requires, and, in fact,
receives, the most careful attention. Weeds are nowhere tolerated in
the south of German East Africa. The crops may fail on the plains,
where droughts are frequent, but never on the plateau with its
abundant rains and heavy dews. Its fortunate inhabitants even have
the satisfaction of seeing the proud Wayao and Wamakua working
for them as labourers, driven by hunger to serve where they were
accustomed to rule.
But the light, sandy soil is soon exhausted, and would yield no
harvest the second year if cultivated twice running. This fact has
been familiar to the native for ages; consequently he provides in
time, and, while his crop is growing, prepares the next plot with axe
and firebrand. Next year he plants this with his various crops and
lets the first piece lie fallow. For a short time it remains waste and
desolate; then nature steps in to repair the destruction wrought by
man; a thousand new growths spring out of the exhausted soil, and
even the old stumps put forth fresh shoots. Next year the new growth
is up to one’s knees, and in a few years more it is that terrible,
impenetrable bush, which maintains its position till the black
occupier of the land has made the round of all the available sites and
come back to his starting point.
The Makonde are, body and soul, so to speak, one with this bush.
According to my Yao informants, indeed, their name means nothing
else but “bush people.” Their own tradition says that they have been
settled up here for a very long time, but to my surprise they laid great
stress on an original immigration. Their old homes were in the
south-east, near Mikindani and the mouth of the Rovuma, whence
their peaceful forefathers were driven by the continual raids of the
Sakalavas from Madagascar and the warlike Shirazis[47] of the coast,
to take refuge on the almost inaccessible plateau. I have studied
African ethnology for twenty years, but the fact that changes of
population in this apparently quiet and peaceable corner of the earth
could have been occasioned by outside enterprises taking place on
the high seas, was completely new to me. It is, no doubt, however,
correct.
The charming tribal legend of the Makonde—besides informing us
of other interesting matters—explains why they have to live in the
thickest of the bush and a long way from the edge of the plateau,
instead of making their permanent homes beside the purling brooks
and springs of the low country.
“The place where the tribe originated is Mahuta, on the southern
side of the plateau towards the Rovuma, where of old time there was
nothing but thick bush. Out of this bush came a man who never
washed himself or shaved his head, and who ate and drank but little.
He went out and made a human figure from the wood of a tree
growing in the open country, which he took home to his abode in the
bush and there set it upright. In the night this image came to life and
was a woman. The man and woman went down together to the
Rovuma to wash themselves. Here the woman gave birth to a still-
born child. They left that place and passed over the high land into the
valley of the Mbemkuru, where the woman had another child, which
was also born dead. Then they returned to the high bush country of
Mahuta, where the third child was born, which lived and grew up. In
course of time, the couple had many more children, and called
themselves Wamatanda. These were the ancestral stock of the
Makonde, also called Wamakonde,[48] i.e., aborigines. Their
forefather, the man from the bush, gave his children the command to
bury their dead upright, in memory of the mother of their race who
was cut out of wood and awoke to life when standing upright. He also
warned them against settling in the valleys and near large streams,
for sickness and death dwelt there. They were to make it a rule to
have their huts at least an hour’s walk from the nearest watering-
place; then their children would thrive and escape illness.”
The explanation of the name Makonde given by my informants is
somewhat different from that contained in the above legend, which I
extract from a little book (small, but packed with information), by
Pater Adams, entitled Lindi und sein Hinterland. Otherwise, my
results agree exactly with the statements of the legend. Washing?
Hapana—there is no such thing. Why should they do so? As it is, the
supply of water scarcely suffices for cooking and drinking; other
people do not wash, so why should the Makonde distinguish himself
by such needless eccentricity? As for shaving the head, the short,
woolly crop scarcely needs it,[49] so the second ancestral precept is
likewise easy enough to follow. Beyond this, however, there is
nothing ridiculous in the ancestor’s advice. I have obtained from
various local artists a fairly large number of figures carved in wood,
ranging from fifteen to twenty-three inches in height, and
representing women belonging to the great group of the Mavia,
Makonde, and Matambwe tribes. The carving is remarkably well
done and renders the female type with great accuracy, especially the
keloid ornamentation, to be described later on. As to the object and
meaning of their works the sculptors either could or (more probably)
would tell me nothing, and I was forced to content myself with the
scanty information vouchsafed by one man, who said that the figures
were merely intended to represent the nembo—the artificial
deformations of pelele, ear-discs, and keloids. The legend recorded
by Pater Adams places these figures in a new light. They must surely
be more than mere dolls; and we may even venture to assume that
they are—though the majority of present-day Makonde are probably
unaware of the fact—representations of the tribal ancestress.
 The references in the legend to the descent from Mahuta to the
Rovuma, and to a journey across the highlands into the Mbekuru
valley, undoubtedly indicate the previous history of the tribe, the
travels of the ancestral pair typifying the migrations of their
descendants. The descent to the neighbouring Rovuma valley, with
its extraordinary fertility and great abundance of game, is intelligible
at a glance—but the crossing of the Lukuledi depression, the ascent
to the Rondo Plateau and the descent to the Mbemkuru, also lie
within the bounds of probability, for all these districts have exactly
the same character as the extreme south. Now, however, comes a
point of especial interest for our bacteriological age. The primitive
Makonde did not enjoy their lives in the marshy river-valleys.
Disease raged among them, and many died. It was only after they
had returned to their original home near Mahuta, that the health
conditions of these people improved. We are very apt to think of the
African as a stupid person whose ignorance of nature is only equalled
by his fear of it, and who looks on all mishaps as caused by evil
spirits and malignant natural powers. It is much more correct to
assume in this case that the people very early learnt to distinguish
districts infested with malaria from those where it is absent.
This knowledge is crystallized in the
ancestral warning against settling in the
valleys and near the great waters, the
dwelling-places of disease and death. At the
same time, for security against the hostile
Mavia south of the Rovuma, it was enacted
that every settlement must be not less than a
certain distance from the southern edge of the
plateau. Such in fact is their mode of life at the
present day. It is not such a bad one, and
certainly they are both safer and more
comfortable than the Makua, the recent
intruders from the south, who have made USUAL METHOD OF
good their footing on the western edge of the CLOSING HUT-DOOR
plateau, extending over a fairly wide belt of
country. Neither Makua nor Makonde show in their dwellings
anything of the size and comeliness of the Yao houses in the plain,
especially at Masasi, Chingulungulu and Zuza’s. Jumbe Chauro, a
Makonde hamlet not far from Newala, on the road to Mahuta, is the
most important settlement of the tribe I have yet seen, and has fairly
spacious huts. But how slovenly is their construction compared with
the palatial residences of the elephant-hunters living in the plain.
The roofs are still more untidy than in the general run of huts during
the dry season, the walls show here and there the scanty beginnings
or the lamentable remains of the mud plastering, and the interior is a
veritable dog-kennel; dirt, dust and disorder everywhere. A few huts
only show any attempt at division into rooms, and this consists
merely of very roughly-made bamboo partitions. In one point alone
have I noticed any indication of progress—in the method of fastening
the door. Houses all over the south are secured in a simple but
ingenious manner. The door consists of a set of stout pieces of wood
or bamboo, tied with bark-string to two cross-pieces, and moving in
two grooves round one of the door-posts, so as to open inwards. If
the owner wishes to leave home, he takes two logs as thick as a man’s
upper arm and about a yard long. One of these is placed obliquely
against the middle of the door from the inside, so as to form an angle
of from 60° to 75° with the ground. He then places the second piece
horizontally across the first, pressing it downward with all his might.
It is kept in place by two strong posts planted in the ground a few
inches inside the door. This fastening is absolutely safe, but of course
cannot be applied to both doors at once, otherwise how could the
owner leave or enter his house? I have not yet succeeded in finding
out how the back door is fastened.

MAKONDE LOCK AND KEY AT JUMBE CHAURO

 This is the general way of closing a house. The Makonde at Jumbe
Chauro, however, have a much more complicated, solid and original
one. Here, too, the door is as already described, except that there is
only one post on the inside, standing by itself about six inches from
one side of the doorway. Opposite this post is a hole in the wall just
large enough to admit a man’s arm. The door is closed inside by a
large wooden bolt passing through a hole in this post and pressing
with its free end against the door. The other end has three holes into
which fit three pegs running in vertical grooves inside the post. The
door is opened with a wooden key about a foot long, somewhat
curved and sloped off at the butt; the other end has three pegs
corresponding to the holes, in the bolt, so that, when it is thrust
through the hole in the wall and inserted into the rectangular
opening in the post, the pegs can be lifted and the bolt drawn out.[50]

MODE OF INSERTING THE KEY

With no small pride first one householder and then a second

showed me on the spot the action of this greatest invention of the
Makonde Highlands. To both with an admiring exclamation of
“Vizuri sana!” (“Very fine!”). I expressed the wish to take back these
marvels with me to Ulaya, to show the Wazungu what clever fellows
the Makonde are. Scarcely five minutes after my return to camp at
Newala, the two men came up sweating under the weight of two
heavy logs which they laid down at my feet, handing over at the same
time the keys of the fallen fortress. Arguing, logically enough, that if
the key was wanted, the lock would be wanted with it, they had taken
their axes and chopped down the posts—as it never occurred to them
to dig them out of the ground and so bring them intact. Thus I have
two badly damaged specimens, and the owners, instead of praise,
come in for a blowing-up.
The Makua huts in the environs of Newala are especially
miserable; their more than slovenly construction reminds one of the
temporary erections of the Makua at Hatia’s, though the people here
have not been concerned in a war. It must therefore be due to
congenital idleness, or else to the absence of a powerful chief. Even
the baraza at Mlipa’s, a short hour’s walk south-east of Newala,
shares in this general neglect. While public buildings in this country
are usually looked after more or less carefully, this is in evident
danger of being blown over by the first strong easterly gale. The only
attractive object in this whole district is the grave of the late chief
Mlipa. I visited it in the morning, while the sun was still trying with
partial success to break through the rolling mists, and the circular
grove of tall euphorbias, which, with a broken pot, is all that marks
the old king’s resting-place, impressed one with a touch of pathos.
Even my very materially-minded carriers seemed to feel something
of the sort, for instead of their usual ribald songs, they chanted
solemnly, as we marched on through the dense green of the Makonde
bush:—
 “We shall arrive with the great master; we stand in a row and have
no fear about getting our food and our money from the Serkali (the
Government). We are not afraid; we are going along with the great
master, the lion; we are going down to the coast and back.”
With regard to the characteristic features of the various tribes here
on the western edge of the plateau, I can arrive at no other
conclusion than the one already come to in the plain, viz., that it is
impossible for anyone but a trained anthropologist to assign any
given individual at once to his proper tribe. In fact, I think that even
an anthropological specialist, after the most careful examination,
might find it a difficult task to decide. The whole congeries of peoples
collected in the region bounded on the west by the great Central
African rift, Tanganyika and Nyasa, and on the east by the Indian
Ocean, are closely related to each other—some of their languages are
only distinguished from one another as dialects of the same speech,
and no doubt all the tribes present the same shape of skull and
structure of skeleton. Thus, surely, there can be no very striking
differences in outward appearance.
 Even did such exist, I should have no time
to concern myself with them, for day after day,
I have to see or hear, as the case may be—in
any case to grasp and record—an
extraordinary number of ethnographic
phenomena. I am almost disposed to think it
fortunate that some departments of inquiry, at
least, are barred by external circumstances.
Chief among these is the subject of iron-
working. We are apt to think of Africa as a
country where iron ore is everywhere, so to
speak, to be picked up by the roadside, and
where it would be quite surprising if the
inhabitants had not learnt to smelt the
material ready to their hand. In fact, the
knowledge of this art ranges all over the
continent, from the Kabyles in the north to the
Kafirs in the south. Here between the Rovuma
and the Lukuledi the conditions are not so
favourable. According to the statements of the
Makonde, neither ironstone nor any other
form of iron ore is known to them. They have
not therefore advanced to the art of smelting
the metal, but have hitherto bought all their
THE ANCESTRESS OF
THE MAKONDE
iron implements from neighbouring tribes.
Even in the plain the inhabitants are not much
better off. Only one man now living is said to
understand the art of smelting iron. This old fundi lives close to
Huwe, that isolated, steep-sided block of granite which rises out of
the green solitude between Masasi and Chingulungulu, and whose
jagged and splintered top meets the traveller’s eye everywhere. While
still at Masasi I wished to see this man at work, but was told that,
frightened by the rising, he had retired across the Rovuma, though
he would soon return. All subsequent inquiries as to whether the
fundi had come back met with the genuine African answer, “Bado”
(“Not yet”).
 BRAZIER

Some consolation was afforded me by a brassfounder, whom I

came across in the bush near Akundonde’s. This man is the favourite
of women, and therefore no doubt of the gods; he welds the glittering
brass rods purchased at the coast into those massive, heavy rings
which, on the wrists and ankles of the local fair ones, continually give
me fresh food for admiration. Like every decent master-craftsman he
had all his tools with him, consisting of a pair of bellows, three
crucibles and a hammer—nothing more, apparently. He was quite
willing to show his skill, and in a twinkling had fixed his bellows on
the ground. They are simply two goat-skins, taken off whole, the four
legs being closed by knots, while the upper opening, intended to
admit the air, is kept stretched by two pieces of wood. At the lower
end of the skin a smaller opening is left into which a wooden tube is
stuck. The fundi has quickly borrowed a heap of wood-embers from
the nearest hut; he then fixes the free ends of the two tubes into an
earthen pipe, and clamps them to the ground by means of a bent
piece of wood. Now he fills one of his small clay crucibles, the dross
on which shows that they have been long in use, with the yellow
material, places it in the midst of the embers, which, at present are
only faintly glimmering, and begins his work. In quick alternation
the smith’s two hands move up and down with the open ends of the
bellows; as he raises his hand he holds the slit wide open, so as to let
the air enter the skin bag unhindered. In pressing it down he closes
the bag, and the air puffs through the bamboo tube and clay pipe into
the fire, which quickly burns up. The smith, however, does not keep
on with this work, but beckons to another man, who relieves him at
the bellows, while he takes some more tools out of a large skin pouch
carried on his back. I look on in wonder as, with a smooth round
stick about the thickness of a finger, he bores a few vertical holes into
the clean sand of the soil. This should not be difficult, yet the man
seems to be taking great pains over it. Then he fastens down to the
ground, with a couple of wooden clamps, a neat little trough made by
splitting a joint of bamboo in half, so that the ends are closed by the
two knots. At last the yellow metal has attained the right consistency,
and the fundi lifts the crucible from the fire by means of two sticks
split at the end to serve as tongs. A short swift turn to the left—a
tilting of the crucible—and the molten brass, hissing and giving forth
clouds of smoke, flows first into the bamboo mould and then into the
holes in the ground.
The technique of this backwoods craftsman may not be very far
advanced, but it cannot be denied that he knows how to obtain an
adequate result by the simplest means. The ladies of highest rank in
this country—that is to say, those who can afford it, wear two kinds
of these massive brass rings, one cylindrical, the other semicircular
in section. The latter are cast in the most ingenious way in the
bamboo mould, the former in the circular hole in the sand. It is quite
a simple matter for the fundi to fit these bars to the limbs of his fair
customers; with a few light strokes of his hammer he bends the
pliable brass round arm or ankle without further inconvenience to
the wearer.
SHAPING THE POT

SMOOTHING WITH MAIZE-COB

CUTTING THE EDGE

 FINISHING THE BOTTOM

LAST SMOOTHING BEFORE

BURNING

FIRING THE BRUSH-PILE

 LIGHTING THE FARTHER SIDE OF
THE PILE

TURNING THE RED-HOT VESSEL

NYASA WOMAN MAKING POTS AT MASASI

Pottery is an art which must always and everywhere excite the
interest of the student, just because it is so intimately connected with
the development of human culture, and because its relics are one of
the principal factors in the reconstruction of our own condition in
prehistoric times. I shall always remember with pleasure the two or
three afternoons at Masasi when Salim Matola’s mother, a slightly-
built, graceful, pleasant-looking woman, explained to me with
touching patience, by means of concrete illustrations, the ceramic art
of her people. The only implements for this primitive process were a
lump of clay in her left hand, and in the right a calabash containing
the following valuables: the fragment of a maize-cob stripped of all
its grains, a smooth, oval pebble, about the size of a pigeon’s egg, a
few chips of gourd-shell, a bamboo splinter about the length of one’s
hand, a small shell, and a bunch of some herb resembling spinach.
 Nothing more. The woman scraped with the
shell a round, shallow hole in the soft, fine
sand of the soil, and, when an active young
girl had filled the calabash with water for her,
she began to knead the clay. As if by magic it
gradually assumed the shape of a rough but
already well-shaped vessel, which only wanted
a little touching up with the instruments
before mentioned. I looked out with the
MAKUA WOMAN closest attention for any indication of the use
MAKING A POT. of the potter’s wheel, in however rudimentary
SHOWS THE a form, but no—hapana (there is none). The
BEGINNINGS OF THE embryo pot stood firmly in its little
POTTER’S WHEEL
depression, and the woman walked round it in
a stooping posture, whether she was removing
small stones or similar foreign bodies with the maize-cob, smoothing
the inner or outer surface with the splinter of bamboo, or later, after
letting it dry for a day, pricking in the ornamentation with a pointed
bit of gourd-shell, or working out the bottom, or cutting the edge
with a sharp bamboo knife, or giving the last touches to the finished
vessel. This occupation of the women is infinitely toilsome, but it is
without doubt an accurate reproduction of the process in use among
our ancestors of the Neolithic and Bronze ages.
There is no doubt that the invention of pottery, an item in human
progress whose importance cannot be over-estimated, is due to
women. Rough, coarse and unfeeling, the men of the horde range
over the countryside. When the united cunning of the hunters has
succeeded in killing the game; not one of them thinks of carrying
home the spoil. A bright fire, kindled by a vigorous wielding of the
drill, is crackling beside them; the animal has been cleaned and cut
up secundum artem, and, after a slight singeing, will soon disappear
under their sharp teeth; no one all this time giving a single thought
to wife or child.
To what shifts, on the other hand, the primitive wife, and still more
the primitive mother, was put! Not even prehistoric stomachs could
endure an unvarying diet of raw food. Something or other suggested
the beneficial effect of hot water on the majority of approved but
indigestible dishes. Perhaps a neighbour had tried holding the hard
roots or tubers over the fire in a calabash filled with water—or maybe
an ostrich-egg-shell, or a hastily improvised vessel of bark. They
became much softer and more palatable than they had previously
been; but, unfortunately, the vessel could not stand the fire and got
charred on the outside. That can be remedied, thought our
ancestress, and plastered a layer of wet clay round a similar vessel.
This is an improvement; the cooking utensil remains uninjured, but
the heat of the fire has shrunk it, so that it is loose in its shell. The
next step is to detach it, so, with a firm grip and a jerk, shell and
kernel are separated, and pottery is invented. Perhaps, however, the
discovery which led to an intelligent use of the burnt-clay shell, was
made in a slightly different way. Ostrich-eggs and calabashes are not
to be found in every part of the world, but everywhere mankind has
arrived at the art of making baskets out of pliant materials, such as
bark, bast, strips of palm-leaf, supple twigs, etc. Our inventor has no
water-tight vessel provided by nature. “Never mind, let us line the
basket with clay.” This answers the purpose, but alas! the basket gets
burnt over the blazing fire, the woman watches the process of
cooking with increasing uneasiness, fearing a leak, but no leak
appears. The food, done to a turn, is eaten with peculiar relish; and
the cooking-vessel is examined, half in curiosity, half in satisfaction
at the result. The plastic clay is now hard as stone, and at the same
time looks exceedingly well, for the neat plaiting of the burnt basket
is traced all over it in a pretty pattern. Thus, simultaneously with
pottery, its ornamentation was invented.
Primitive woman has another claim to respect. It was the man,
roving abroad, who invented the art of producing fire at will, but the
woman, unable to imitate him in this, has been a Vestal from the
earliest times. Nothing gives so much trouble as the keeping alight of
the smouldering brand, and, above all, when all the men are absent
from the camp. Heavy rain-clouds gather, already the first large
drops are falling, the first gusts of the storm rage over the plain. The
little flame, a greater anxiety to the woman than her own children,
flickers unsteadily in the blast. What is to be done? A sudden thought
occurs to her, and in an instant she has constructed a primitive hut
out of strips of bark, to protect the flame against rain and wind.
This, or something very like it, was the way in which the principle
of the house was discovered; and even the most hardened misogynist
cannot fairly refuse a woman the credit of it. The protection of the
hearth-fire from the weather is the germ from which the human
dwelling was evolved. Men had little, if any share, in this forward
step, and that only at a late stage. Even at the present day, the
plastering of the housewall with clay and the manufacture of pottery
are exclusively the women’s business. These are two very significant
survivals. Our European kitchen-garden, too, is originally a woman’s
invention, and the hoe, the primitive instrument of agriculture, is,
characteristically enough, still used in this department. But the
noblest achievement which we owe to the other sex is unquestionably
the art of cookery. Roasting alone—the oldest process—is one for
which men took the hint (a very obvious one) from nature. It must
have been suggested by the scorched carcase of some animal
overtaken by the destructive forest-fires. But boiling—the process of
improving organic substances by the help of water heated to boiling-
point—is a much later discovery. It is so recent that it has not even
yet penetrated to all parts of the world. The Polynesians understand
how to steam food, that is, to cook it, neatly wrapped in leaves, in a
hole in the earth between hot stones, the air being excluded, and
(sometimes) a few drops of water sprinkled on the stones; but they
do not understand boiling.
To come back from this digression, we find that the slender Nyasa
woman has, after once more carefully examining the finished pot,
put it aside in the shade to dry. On the following day she sends me
word by her son, Salim Matola, who is always on hand, that she is
going to do the burning, and, on coming out of my house, I find her
already hard at work. She has spread on the ground a layer of very
dry sticks, about as thick as one’s thumb, has laid the pot (now of a
yellowish-grey colour) on them, and is piling brushwood round it.
My faithful Pesa mbili, the mnyampara, who has been standing by,
most obligingly, with a lighted stick, now hands it to her. Both of
them, blowing steadily, light the pile on the lee side, and, when the
flame begins to catch, on the weather side also. Soon the whole is in a
blaze, but the dry fuel is quickly consumed and the fire dies down, so
that we see the red-hot vessel rising from the ashes. The woman
turns it continually with a long stick, sometimes one way and
sometimes another, so that it may be evenly heated all over. In
twenty minutes she rolls it out of the ash-heap, takes up the bundle
of spinach, which has been lying for two days in a jar of water, and
sprinkles the red-hot clay with it. The places where the drops fall are
marked by black spots on the uniform reddish-brown surface. With a
sigh of relief, and with visible satisfaction, the woman rises to an
erect position; she is standing just in a line between me and the fire,
from which a cloud of smoke is just rising: I press the ball of my
camera, the shutter clicks—the apotheosis is achieved! Like a
priestess, representative of her inventive sex, the graceful woman
stands: at her feet the hearth-fire she has given us beside her the
invention she has devised for us, in the background the home she has
built for us.
At Newala, also, I have had the manufacture of pottery carried on
in my presence. Technically the process is better than that already
described, for here we find the beginnings of the potter’s wheel,
which does not seem to exist in the plains; at least I have seen
nothing of the sort. The artist, a frightfully stupid Makua woman, did
not make a depression in the ground to receive the pot she was about
to shape, but used instead a large potsherd. Otherwise, she went to
work in much the same way as Salim’s mother, except that she saved
herself the trouble of walking round and round her work by squatting
at her ease and letting the pot and potsherd rotate round her; this is
surely the first step towards a machine. But it does not follow that
the pot was improved by the process. It is true that it was beautifully
rounded and presented a very creditable appearance when finished,
but the numerous large and small vessels which I have seen, and, in
part, collected, in the “less advanced” districts, are no less so. We
moderns imagine that instruments of precision are necessary to
produce excellent results. Go to the prehistoric collections of our
museums and look at the pots, urns and bowls of our ancestors in the
dim ages of the past, and you will at once perceive your error.
MAKING LONGITUDINAL CUT IN
BARK

DRAWING THE BARK OFF THE LOG

REMOVING THE OUTER BARK

 BEATING THE BARK

WORKING THE BARK-CLOTH AFTER BEATING, TO MAKE IT

SOFT

MANUFACTURE OF BARK-CLOTH AT NEWALA

To-day, nearly the whole population of German East Africa is
clothed in imported calico. This was not always the case; even now in
some parts of the north dressed skins are still the prevailing wear,
and in the north-western districts—east and north of Lake
Tanganyika—lies a zone where bark-cloth has not yet been
superseded. Probably not many generations have passed since such
bark fabrics and kilts of skins were the only clothing even in the
south. Even to-day, large quantities of this bright-red or drab
material are still to be found; but if we wish to see it, we must look in
the granaries and on the drying stages inside the native huts, where
it serves less ambitious uses as wrappings for those seeds and fruits
which require to be packed with special care. The salt produced at
Masasi, too, is packed for transport to a distance in large sheets of
bark-cloth. Wherever I found it in any degree possible, I studied the
process of making this cloth. The native requisitioned for the
purpose arrived, carrying a log between two and three yards long and
as thick as his thigh, and nothing else except a curiously-shaped
mallet and the usual long, sharp and pointed knife which all men and
boys wear in a belt at their backs without a sheath—horribile dictu!
[51]
Silently he squats down before me, and with two rapid cuts has
drawn a couple of circles round the log some two yards apart, and
slits the bark lengthwise between them with the point of his knife.
With evident care, he then scrapes off the outer rind all round the
log, so that in a quarter of an hour the inner red layer of the bark
shows up brightly-coloured between the two untouched ends. With
some trouble and much caution, he now loosens the bark at one end,
and opens the cylinder. He then stands up, takes hold of the free
edge with both hands, and turning it inside out, slowly but steadily
pulls it off in one piece. Now comes the troublesome work of
scraping all superfluous particles of outer bark from the outside of
the long, narrow piece of material, while the inner side is carefully
scrutinised for defective spots. At last it is ready for beating. Having
signalled to a friend, who immediately places a bowl of water beside
him, the artificer damps his sheet of bark all over, seizes his mallet,
lays one end of the stuff on the smoothest spot of the log, and
hammers away slowly but continuously. “Very simple!” I think to
myself. “Why, I could do that, too!”—but I am forced to change my
opinions a little later on; for the beating is quite an art, if the fabric is
not to be beaten to pieces. To prevent the breaking of the fibres, the
stuff is several times folded across, so as to interpose several
thicknesses between the mallet and the block. At last the required
state is reached, and the fundi seizes the sheet, still folded, by both
ends, and wrings it out, or calls an assistant to take one end while he
holds the other. The cloth produced in this way is not nearly so fine
and uniform in texture as the famous Uganda bark-cloth, but it is
quite soft, and, above all, cheap.
Now, too, I examine the mallet. My craftsman has been using the
simpler but better form of this implement, a conical block of some
hard wood, its base—the striking surface—being scored across and
across with more or less deeply-cut grooves, and the handle stuck
into a hole in the middle. The other and earlier form of mallet is
shaped in the same way, but the head is fastened by an ingenious
network of bark strips into the split bamboo serving as a handle. The
observation so often made, that ancient customs persist longest in
connection with religious ceremonies and in the life of children, here
finds confirmation. As we shall soon see, bark-cloth is still worn
during the unyago,[52] having been prepared with special solemn
ceremonies; and many a mother, if she has no other garment handy,
will still put her little one into a kilt of bark-cloth, which, after all,
looks better, besides being more in keeping with its African
surroundings, than the ridiculous bit of print from Ulaya.
MAKUA WOMEN