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 Contents vii

Viruses 315
KEY THEME : molecular Plant Viruses 316 20 Introduction to the Plant
Viroids and Prions 320 Kingdom: Bryophytes 376
SUMMARY 320
REVIEW QUESTIONS 321 Overview 377
DISCUSSION QUESTIONS 321 Learning Outcomes 377
ADDITIONAL READING 321 Introduction to the Bryophytes 378
LEARNING ONLINE 321
Phylum Hepaticophyta—Liverworts 379
Phylum Anthocerophyta—Hornworts 383
18 Kingdom Protista 322 Phylum Bryophyta—Mosses 383
KEY THEME : ecology Hibernating Mosses 387
Overview 323 Human and Ecological Relevance of
Learning Outcomes 323 Bryophytes 388
Features of Kingdom Protista 324 SUMMARY 388
Algae 324 REVIEW QUESTIONS 389
Phylum Chlorophyta—the Green DISCUSSION QUESTIONS 389
ADDITIONAL READING 390
Algae 324
LEARNING ONLINE 390
Phylum Chromophyta—the Yellow-Green
Algae, Golden-Brown Algae, Diatoms,
and Brown Algae 331
Phylum Rhodophyta—the Red Algae 336
21 The Seedless Vascular Plants:
Ferns and Their Relatives 391
Phylum Euglenophyta—the Euglenoids 337
Phylum Dinophyta—the Dinoflagellates 338 Overview 392
Phylum Cryptophyta—the Learning Outcomes 392
Cryptomonads 340 Phylum Psilotophyta—the Whisk Ferns 392
Phylum Prymnesiophyta (Haptophyta)—the Phylum Lycophyta—the Ground Pines, Spike
Haptophytes 340 Mosses, and Quillworts 394
Phylum Charophyta—the Stoneworts 340 Phylum Equisetophyta—the Horsetails
Human and Ecological Relevance of the and Scouring Rushes 400
Algae 341 Phylum Polypodiophyta—the Ferns 404
Other Members of Kingdom Protista 344 Fossils 411
Phylum Myxomycota—the Plasmodial KEY THEME : ecology Ferns and Fossil
Slime Molds 345 Fuels 412
SUMMARY 413
Phylum Dictyosteliomycota—the Cellular
REVIEW QUESTIONS 414
Slime Molds 346 DISCUSSION QUESTIONS 414
Phylum Oomycota—the Water Molds 346 ADDITIONAL READING 414
SUMMARY 348 LEARNING ONLINE 415
REVIEW QUESTIONS 349
DISCUSSION QUESTIONS 350
ADDITIONAL READING 350
LEARNING ONLINE 350
22 Introduction to Seed Plants:
Gymnosperms 416
19 Kingdom Fungi 351
Overview 417
Learning Outcomes 417
Overview 352 Phylum Pinophyta—the Conifers 418
Learning Outcomes 352 KEY THEME : ecology Resilient and Useful
Distinctions Between Kingdoms Protista Gymnosperms 419
and Fungi 352 Other Gymnosperms 422
Kingdom Fungi 353 Human Relevance of Gymnosperms 427
Lichens 371 KEY THEME : evolution A Living Fossil? 432
SUMMARY 373 SUMMARY 433
REVIEW QUESTIONS 374 REVIEW QUESTIONS 434
DISCUSSION QUESTIONS 375 DISCUSSION QUESTIONS 434
ADDITIONAL READING 375 ADDITIONAL READING 434
LEARNING ONLINE 375 LEARNING ONLINE 434
viii Contents

SUMMARY 503
23 Seed Plants: Angiosperms 435 REVIEW QUESTIONS 504
DISCUSSION QUESTIONS 504
Overview 436 ADDITIONAL READING 504
Learning Outcomes 436 LEARNING ONLINE 505

Phylum Magnoliophyta—the Flowering

Plants 437
KEY THEME : molecular The Difference between
26 Biomes 506
“n” and “x” in Plant Life Cycles 444 Overview 507
Pollination Ecology 446 Learning Outcome 507
Herbaria and Plant Preservation 450 Major Biomes of the World 507
SUMMARY 453
KEY THEME : ecology Alpine Flora as an Indication
REVIEW QUESTIONS 453
of Climate Change: The Gloria Project 512
DISCUSSION QUESTIONS 454
SUMMARY 516
ADDITIONAL READING 454
REVIEW QUESTIONS 517
LEARNING ONLINE 454
DISCUSSION QUESTIONS 517
ADDITIONAL READING 517

24 Flowering Plants and

LEARNING ONLINE 517

Civilization 455 Appendix 1 Scientific Names of Organisms

Mentioned in the Text A1
Overview 456
Learning Outcomes 456 Appendix 2 Biological Controls A20
Origin of Cultivated Plants 456 General Controls A20
Selected Families of Flowering Plants 457 Specific Controls A22
Dicots (Now Recognized in Two Groups) 459 Companion Planting A22
Monocots 474 Additional Reading A26
KEY THEME : ecology Wild Rice—More Than Appendix 3 Useful and Poisonous Plants,
Just Food 476 Fungi, and Algae A27
KEY THEME : ecology Coffee Wild Edible Plants, Fungi, and Algae A27
and Caffeine 478 Poisonous Plants and Fungi A27
SUMMARY 479 Medicinal Plants, Fungi, and Algae A27
REVIEW QUESTIONS 479
Hallucinogenic Plants A45
DISCUSSION QUESTIONS 480
ADDITIONAL READING 480
Spice Plants A45
LEARNING ONLINE 480 Dye Plants A45
Additional Reading A51
Appendix 4 House Plants and Home
25 Ecology 481 Gardening A52
Overview 482 Growing House Plants A52
Learning Outcomes 482 Common House Plants A53
Plants and the Environment 482 Growing Vegetables A62
Life Histories 487 Common Vegetables and their Nutritional
Natural Cycles 488 Values A63
Succession 491 Pruning A68
KEY THEME : ecology Plant Population Major Types of Grafting A69
Ecology 491 Additional Reading A75
Impact of Humans on Plant Appendix 5 Metric Equivalents and
Communities 496
Conversion Tables A76
Loss of Biodiversity 499
Restoration of the Land 501 Glossary G1
KEY THEME : ecology John Muir, Father of Photo Credits C1
America’s National Park System 502 Index I1
 About the Authors
Plant Propogation and Biotechnology ix

Introductory Plant Biology was originally written by Kingsley

R. Stern (1927–2006), who spent over 40 years as a devoted botanist
and teacher. It is estimated that Dr. Stern educated 15,000 students
through classroom/laboratory teaching and inspired thousands of
botanists as the author of this book, which has sold over 250,000
copies since the first edition was published in 1979. Kingsley’s
enthusiasm for the botanical world captivated those around him
for many decades. He will long be remembered for his attention
to detail and dedication to high standards, along with a refreshing
sense of humor. It has always been Kingsley’s aspiration that those
who read Stern’s Introductory Plant Biology will share his lifelong
love of botany.

Jim Bidlack, Kingsley Stern, and Shelley Jansky at Kingsley’s office

residence in Paradise, California.

In late 1999/early 2000, Drs. Jim Bidlack and Shelley Jansky joined Kingsley Stern in editing and updating this textbook. They
worked with Dr. Stern on several editions and have since carried his legacy forward to educate and inspire young botanists.
This thirteenth edition reflects the same accuracy, content, and enthusiasm of the Stern writing style, along with revisions and
updates to make it an up-to-date and enjoyable resource and textbook for plant biology.

James E. Bidlack Shelley H. Jansky

Jim Bidlack received a B.S. Degree
in Agronomy, with a Soil & Crop
Option, from Purdue University in
1984 and continued his education with
a Master’s Degree in Crop Physiology
at the University of Arkansas in 1986.
Upon completing a Ph.D. in Plant Physiology at Iowa State
University in 1990, Jim joined the teaching faculty at the
University of Central Oklahoma (UCO) where he is a Professor
of Biology. His first paper was published from undergraduate
research at Purdue University on the use of synthetic growth
regulators to stimulate seed germination. Subsequent work at
Arkansas, Iowa, and Oklahoma focused on soybean physiol-
ogy, cell wall chemistry, and alternative crops, as well as teach-
ing responsibilities in plant biology. Equipment and student
salaries for Jim’s research projects have been funded by grants
from the National Science Foundation (NSF) and the United
States Department of Agriculture (USDA). Nearly 20 refer-
eed publications, as well as 50 abstracts and popular articles,
have resulted from this work. Jim has been recognized with
UCO’s Presidential Partner’s Excellence in Teaching Award;
University Merit Awards in Service, Research, and Teaching; as
well as the Biology Club and the Pre-Med Teaching Awards.
Some of Jim’s additional responsibilities have included par-
ticipation on NSF and USDA Review Panels, Editor of the
Biology Discipline for the Multimedia Educational Resource
for Learning and Online Teaching (MERLOT), and President
of Metabolism Foundation.
Shelley Jansky received a Bachelor’s
Degree in Biology from the University
of Wisconsin–Stevens Point in 1982,
and M.S. and Ph.D. degrees in Plant
Breeding and Plant Genetics from the
University of Wisconsin–Madison in
1984 and 1986, respectively. Her graduate work focused
on developing methods to incorporate genes from wild
relatives of potato into the cultivated potato. Then, she
spent four years as an Assistant Professor at North Dakota
State University, teaching courses in plant breeding and
plant propagation, and performing research in the potato
breeding program. She taught courses in botany, genetics,
and horticulture, and continued to perform potato genet-
ics research at the University of Wisconsin–Stevens Point
from 1990 until 2004. She was the Chair of the Department
of Biology and was promoted to Associate Professor in
1992 and Professor in 1995. In 2004, she accepted a posi-
tion as a Research Geneticist with the U.S. Department of
Agriculture and an Assistant Professor in the Department
of Horticulture at the University of Wisconsin–Madison.
Her potato research program focuses on the utilization of
disease resistance and nutritional quality genes from wild
potato relatives for the improvement of cultivated potato
varieties. She received the University of Wisconsin–
Stevens Point Excellence in Teaching Award in 1992 and
the University Scholar Award in 2000. She has published
65 refereed research articles and five book chapters.
ix
Preface
Plants and algae are essential for life on earth as it exists today.
They provide our world with oxygen and food, contribute an
essential part of water and nutrient cycling in ecosystems, pro-
vide clothing and shelter, and add beauty to our environment.
Some scientists believe that if photosynthetic organisms exist
on planets beyond our solar system, it would be possible to
sustain other forms of life that depend upon them to survive.
Botany today plays a special role in many interests of
both major and nonmajor students. For example, in this
text, topics such as global warming, ozone layer depletion,
acid rain, genetic engineering, organic gardening, Native
American and pioneer uses of plants, pollution and recy-
cling, house plants, backyard vegetable gardening, natural
dye plants, poisonous and hallucinogenic plants, nutritional
values of edible plants, and many other topics are discussed.
To intelligently pursue such topics, one needs to understand
how plants are constructed, and how they function. To this
end, the text assumes little prior knowledge of the sciences
on the part of the student, but covers basic botany, without
excessively resorting to technical terms. The coverage, how-
ever, includes sufficient depth to prepare students to go fur-
ther in the field, should they choose to do so.
The text is arranged so that certain sections can be omit-
ted in shorter courses. Such sections may include topics
such as soils, molecular genetics, and phylum Bryophyta.
Because botany instructors vary greatly in their opinions
about the depth of coverage needed for photosynthesis and
respiration in an introductory botany course open to both
majors and nonmajors, these topics are presented at three
different levels. Some instructors will find one or two levels
sufficient, whereas others will want to include all three.
Both majors in botany and nonmajors who may initially
be disinterested in the subject matter of a required course fre-
quently become engrossed if the material is related repeatedly
to their popular interests. This is reflected, as intimated above,
in the considerable amount of ecology and ethnobotany
included with traditional botany throughout the book.
Organization of the Text
A relatively conventional sequence of botanical subjects is
followed. Chapters 1 and 2 cover introductory and background
information; Chapters 3 through 11 deal with structure and
function; Chapters 12 and 13 introduce meiosis, genetics, and
molecular biology. Chapter 14 discusses plant propagation
and biotechnology; Chapter 15 introduces evolution; Chapter
16 deals with classification; Chapters 17 through 23 stress, in
phylogenetic sequence, the diversity of organisms traditionally
regarded as plants; and Chapter 24 deals with ethnobotanical
x
aspects
aspect
ts andd oth
other
ther in
iinformation
f rmatio
fo i n of general interest pertaining to
16 major plant families or groups of families. Chapters 25 and
26 present an overview of the vast topic of ecology, although
ecological topics and applied botany are included in the pre-
ceding chapters as well. Some of these topics are broached in
anecdotes that introduce the chapters, while others are men-
tioned in text boxes as well as the appendices.
Learning Aids
A chapter outline, review questions, discussion questions, and
additional reading lists are provided for each chapter. New
terms are defined as they are introduced, and those that are
boldfaced are included, with their pronunciation, in a glos-
sary. A list of the scientific names of all organisms mentioned
throughout the text is given in Appendix 1. Appendix 2 deals
with biological controls and companion planting. Appendix
3 includes wild edible plants, poisonous plants, medicinal
plants, hallucinogenic plants, spices, tropical fruits, and natu-
ral dye plants. Appendix 4 gives horticultural information on
house plants, along with brief discussions on how to cultivate
vegetables. Nutritional values of the vegetables are included.
Appendix 5 covers metric equivalents and conversion tables.
New to this Edition
The thirteenth edition has retained the hallmark style and
pedagogy that have made it one of the most enduring and
popular introductory plant biology books on the market. At
the same time, this edition has undergone many changes to
expand upon, revise, and update topics in plant biology. All
chapters now incorporate measurable learning outcomes
and updated additional readings. For those instructors using
Connect Botany, all learning outcomes (LOs) are directly
tied to assessment within the question and test banks. In
addition, most of the chapters include revisions and updates
as suggested by reviewers. Some of the major changes
include an introduction to the new classification of plants
and plant-related species, integration of biotechnology into
several chapters, and inclusion of new featured text boxes
addressing key themes of ecology, evolution, and molecular
biology throughout the text. These new text boxes provide
instructors and students with information that helps them to
understand how modern botany affects our everyday lives.
Many new photographs have replaced some of the older pic-
tures or have been added within individual chapters. Some of
the more interesting components that make this 13th Edition
more accurate and up-to-date with our current understanding
of plant biology include:

Chapter 1 (What is Plant Biology?): New information on
climate change, population dynamics, and alternative energy
has been added to many parts of the chapter to make it more
engaging and up-to-date. Text boxes on “Who Needs Plants?”
as well as “Biology and the Internet” have been extensively
re-written and several photographs have been replaced.
Chapter 2 (The Nature of Life): Figures on energy conver-
sion and enzyme action have been replaced to improve accuracy,
and a new text box entitled “The Skinny on Fats” has been added.
Chapter 3 (Cells): Many of the figures have been
revised; particularly the cell and its components to more
accurately represent how these are seen under a microscope.
New, modern photographs of microscopes have been added
along with a better explanation of confocal electron micros-
copy. Updated text information now includes discussion of
endosymbiotic theory as well as functions of vacuoles.
Chapter 4 (Tissues): The text box on “Chimeras and
Variegated Leaves” has been modified to provide more
detail. New information has been added on discussion of
cell wall thickening in collenchyma cells and tracheary ele-
ments, and an improved depiction of lenticels is now shown
in the section on periderm.
Chapter 5 (Roots and Soils): Improved photographs of
sweet potatoes, cassava, and a soil profile have been incor-
porated into the chapter. The text has been modified to pro-
vide a more accurate description of Casparian strips, lateral
root formation, and soil horizons. The sections on parasitic
roots and mycorrhizae have been extensively re-written to
incorporate new terminology for these topics.
Chapter 6 (Stems): Artwork showing heartwood and
softwood has been replaced to show a better representation
of the characteristics and location of these parts of woody
stems. The section on wood and its uses has been exten-
sively modified so that the figures are better organized and
figure legends are easier to read.
Chapter 7 (Leaves): Photographs of thorns and prickles,
as well as some modified leaves, have been replaced to show
better depiction of these structures.
Chapter 8 (Flowers, Fruits, and Seeds): A photograph of
a rare Australian orchid, as well as a more clearly-displayed
specimen of Rafflesia have been added to the chapter. A new
text box on “Goober Peas” has been incorporated into text,
along with a more accurate description of germination.
Chapter 9 (Water in Plants): A sharper opening pho-
tograph for the chapter has been added along with more
refined artwork to show diffusion and the pressure-flow
hypothesis. The section on water potential has been exten-
sively rewritten and a new text box on “Measuring Water
Potential and Psychrometry” has been added to the chapter.
Chapter 10 (Plant Metabolism): Artwork has been revised
to show a better summary of photosynthesis and respiration,
and more details in the Calvin cycle. A new text box has been
added on “Photosynthesis, Global Warming, and Tropical Rain
Forests” to show applications of metabolic concepts.
Chapter 11 (Growth and Development): New figures
have been incorporated to show structures of plant hormones
as well as signal transduction and a new text box entitled,
Preface xi
“Plant Hormones—Beyond the Classic Five,” has been
added. Updated information has also been added about auxin-
tryptophan interactions, the role of ethylene in fruit ripening,
photoreceptor proteins, and the flowering locus T gene.
Chapter 12 (Meiosis and Alternation of Generations):
Descriptions of meiotic phases have been revised to be more
accurate, along with new discussions of chiasma and poly-
ploidy. Two new text boxes on “Why Plants Have a Sex Life”
and “FISH and GISH Molecular Techniques” have been added.
Chapter 13 (Genetics and Molecular Biology): A new
textbox has been added on “Massive DNA Sequencing,”
along with an improved image of Gregor Mendel.
Chapter 14 (Plant Breeding, Propagation, and
Biotechnology): Updated images have been incorporated to
provide better explanations of transgenics, plant propaga-
tion, and tissue culture. New text has been added to describe
the development of golden rice.
Chapter 15 (Evolution): Several parts of this chapter have
been revised and updated to include discussions on epigen-
esis and polyploidy. A new text box on “Our Daily Bread” has
been added to describe the domestication of wheat.
Chapter 16 (Plant Names and Classification): This chap-
ter has been extensively rewritten to describe modern clas-
sification techniques. Sections on the International Code of
Nomenclature of Cultivated Plants and the Phylocode have
been added, along with a newly-proposed classification key.
Chapter 17 (Domain (Kingdom) Bacteria, Domain
(Kingdom) Archaea, and Viruses): New, three-dimensional
depictions of bacteria have been incorporated into the chap-
ter along with improved artistic representations of viruses.
A discussion of Clostridium difficile, has been added, as well
as a new text box entitled “The Social Life of Prokaryotes.”
Chapter 18 (Kingdom Protista): Improved images of
red algae, Euglena, and dinoflagellates have been incorpo-
rated to show more accurate depictions of these specimens.
Chapter 19 (Kingdom Fungi): The use of DNA sequence
data for classification of fungi is described. New information is
also introduced, including use of fungi in cleaning oil spills and
treatment of certain tree species to prevent the spread of disease.
Chapter 20 (Introduction to the Plant Kingdom:
Bryophytes): New introductory information has been added
on the evolutionary origin of plants, along with a summary
of bryophyte classification. Artwork for hornworts has been
improved and new pictures of hairy cap moss and peat moss
have been added to the chapter.
Chapter 21 (The Seedless Vascular Plants: Ferns and
Their Relatives): Improved photographs of ferns have been
incorporated into the text, along with new evolutionary
information about the origin of seedless vascular plants. In
addition, a new text box on “Ferns and Fossil Fuels” has
been added to the chapter.
Chapter 22 (Introduction to Seed Plants: Gymnosperms):
A new photograph of the rare Wollemi pine, along with a more
modern photograph of a newsprint factory have been added
to the text. The introductory paragraph has been updated with
new information about molecular analysis of gymnosperms
as well as a summary of their current classification. A text box
xii Preface
on “Resilient and Useful Gymnosperms” has been added to
show modern uses of pines and related species.
Chapter 23 (Seed Plants: Angiosperms): Improved
photographs of snowplant flowers, as well as pollination of
other flowers, have been included in the text. A more exten-
sive explanation of angiosperm origin has also been added
to the introduction.
Chapter 24 (Flowering Plants and Civilization): The
photograph of a poppy capsule has been replaced and a new
Acknowledgements
Over 250 reviewers for the past few editions, along with
over 20 reviewers for this thirteenth edition, have helped
to revise and update Stern’s Introductory Plant Biology.
In particular, David Spooner, USDA-ARS Botanist and
Professor at the University of Wisconsin, provided excellent
input and rewrote much of Chapter 16 on Plant Names and
Classification. Additional contributions and encouragement
were provided by Jan Monelo; colleagues at the University
of Central Oklahoma and University of Wisconsin; and the
design, editorial, and production staffs of McGraw-Hill
Publishers. The authors extend thanks to the following
reviewers who provided recent feedback on the text and
the illustrations. Their help has been very useful in shaping
the thirteenth edition of Stern’s Introductory Plant Biology.
These reviewers include the following:
Donovan Bailey, New Mexico State University
Terese Barta, University of Wisconsin-Stevens Point
Lisa Boucher, University of Nebraska-Omaha
Roger del Moral, University of Washington
Stephen W. Fuller, University of Mary Washington
Ed Gasque, University of Wisconsin-Stevens Point
Rick Hammer, Hardin-Simmons University
Terry Jordan, College of Southern Maryland
John Kiss, Miami University
Fengshan Ma, Wilfrid Laurier University
Karen McGinnis, Florida State University
James Mickle, North Carolina State University
Carrie Monlux, California State University-Chico
Clark L. Ovrebo, University of Central Oklahoma
Bob Peregoy, Spokane Community College
Flona Redway, Barry University
Traesha R. Robertson, Texas Tech University
Linda Brooke Stabler, University of Central Oklahoma
Leslie Towill, Arizona State University
Carol Wake, South Dakota State University
Upon reaching this milestone thirteenth edition, we would
also like to once again extend gratitude to the reviewers of
earlier editions, who have provided considerable comments
and suggestions. Although too numerous to include here,
their contributions have been much appreciated. The follow-
ing is a list of reviewers for recent editions:
Ligia Arango, Stone Child College
Joseph Arditti, University of California–Irvine
text box on “Wild Rice—More than Just Food” has been
added to the chapter.
Chapter 25 (Ecology): Topics that address the impact of
humans on the biosphere have been reorganized to follow a
more logical approach to headings and sub-headings.
Chapter 26 (Biomes): This chapter has been re-
organized and rewritten to incorporate world biomes in
addition to those found in North America.
Mark H. Armitage, Azusa Pacific University
Janice Asel, Mitchell Community College
Tasneem K. Ashraf, Cochise College–Sierra Vista
Ralph A. Backhaus, Arizona State University
Nina L. Baghai-Riding, Delta State University
Randy G. Balice, New Mexico Highlands University
Susan C. Barber, Oklahoma City University
Paul W. Barnes, Southwest Texas State University
Sharon Bartholomew-Began, West Chester University
Robert W. Bauman, Jr., Amarillo College
Dorothea Bedigian, Washington University
Patricia Bedinger, Colorado State University
Maria Begonia, Jackson State University
Robert A. Bell, University of Wisconsin–Stevens Point
Cynthia A. Bottrell, Scott Community College
Richard R. Bounds, Mount Olive College
Richard G. Bowmer, Idaho State University
Rebecca D. Bray, Old Dominion University
James A. Brenneman, University of Evansville
George M. Briggs, State University of New York
Michelle Briggs, Lycoming College
George M. Brooks, Ohio University
Suzanne Butler, Miami-Dade College
William J. Campbell, Louisiana Technical University
Ajoy G. Chakrabarti, South Carolina State University
Brad S. Chandler, Palo Alto College
Gregory Chandler, University of North Carolina–
Wilmington
James A. Christian, Louisiana Technical University
Richard Churchill, Southern Maine Technical College
Jerry A. Clonts, Anderson College
William Cook, Midwestern State University
John Cruzan, Geneva College
Kenneth J. Curry, University of Southern Mississippi
David B. Czarnecki, Loras College
Stephen S. Daggett, Avila College
Raviprakash G. Dani, Texas Tech University
Roy Darville, East Texas Baptist University
Cynthia Dassler, Ohio State University
Bill D. Davis, Rutgers University
Jerry D. Davis, University of Wisconsin–LaCrosse
John W. Davis, Benedictine College
Roger del Moral, University of Washington
Semma Dhir, Fort Valley State University
Rebecca M. DiLiddo, Mount Ida College
Susan C. Dixon, Walla Walla College
Ben L. Dolbeare, Lincoln Land Community College

Patricia M. Dooris, Saint Leo College
Donald Drake, University of Hawaii
Tom Dudley, Angelina College
Jan Federic Dudt, Bartlesville Wesleyan College
Diane Dudzinski, Washington State Community College
Kerry B. Dunbar, Dalton State College
Carolyn S. Dunn, University of North Carolina–
Wilmington
Robert Ediger, California State University–Chico
H. Herbert Edwards, Western Illinois University
William Eisinger, Santa Clara University
Inge Eley, Hudson Valley Community College
Thomas E. Elthon, University of Nebraska–Lincoln
Frederick B. Essig, University of South Florida
G. F. Estabrook, The University of Michigan
James Ethridge, Joliet Junior College
Paul G. Fader, Freed-Hardeman University
Bruce Felgenhauer, University of Louisiana–Lafayette
Jorge F. S. Ferreira, Southern Illinois University–
Carbondale
David G. Fisher, Maharishi University of Management
Rosemary H. Ford, Washington College
Stephen W. Fuller, Mary Washington College
James Garner, Horry-Georgetown Technical College
Sibdas Ghosh, University of Wisconsin–Whitewater
Mike Gipson, Oklahoma Christian University
Katherine Glew, University of Washington
Richard Glick, Winston-Salem State University
Charles Good, Ohio State University
David L. Gorchov, Miami University of Ohio
Scott A. Gordon, University of Southern Illinois
Govindjee, University of Illinois
Steve Greenwald, Gordon College
Sharon Gusky, Northwestern Connecticut Community
Technical College
Timothy C. Hall, Texas A & M University
Mark Hammer, Wayne State College
Susan Han, University of Massachusetts
Laszlo Hanzely, Northern Illinois University
Joyce Phillips Hardy, Chadron State College
Nancy E. Harris, Elon College
David Hartsell, Phillips Community College
Jill F. Haukos, South Plains College
David L. Herrin, University of Texas–Austin
Peter Heywood, Brown University
Jeffrey P. Hill, Idaho State University
L. Michael Hill, Bridgewater College
H. H. Ho, State University of New York–New Paltz
A. Scott Holaday, Texas Tech University
Elisabeth A. Hooper, Truman State University
Susan Houseman, Southeastern Community College
Lauren D. Howard, Norwich University
Vernon R. Huebschwerlen, Reedley Community College
Patricia L. Ireland, San Jacinto College, South
William A. Jensen, Ohio State University
Cindy Johnson-Groh, Gustavus Adolphus College
Chad Jordan, North Carolina State University
Toney Keeney, Southwest Texas State
Preface xiii
Sekender A. Khan, Elizabeth City State University
Joanne M. Kilpatrick, Auburn University–Montgomery
Helen G. Kiss, Miami University
John Z. Kiss, Miami University of Ohio
Kaoru Kitajima, University of Florida
Sharon Klavins, University of Wisconsin–Platteville
Roger C. Klockziem, Martin Luther College
Robert L. Koenig, Southwest Texas Junior College
David W. Kramer, Ohio State University–Mansfield
Robert N. Kruger, Mayville State University
Martin LaBar, Southern Wesleyan University
Vic Landrum, Washburn University
James M. Lang, Greenville College
Brenda Price Latham, Merced College
Cheryl R. Laursen, Eastern Illinois University
Peter J. Lemay, College of the Holy Cross
Donald C. Leynaud, Wabash Valley College
Barbara E. Liedl, Central College
John F. Logue, University of South Carolina–Sumter
Elizabeth L. Lucyszyn, Medaille College
Karen Lustig, Harper College
Erin D. MacKenzie, Weatherford College
Paul Mangum, Midland College
Steve Manning, Arkansas State University–Beebe
Michael H. Marcovitz, Midland Lutheran College
Bernard A. Marcus, Genesee Community College
David Martin, Centralia College
Margaret Massey, Mississippi University for Women
William J. Mathena, Kaskaskia College
Alicia Mazari-Andersen, Kwantlen University College
Andrew McCubbin, Washington State University
Joseph H. McCulloch, Normandale Community College
Julie A. Medlin, Northwestern Michigan College
Larry Mellichamp, University of North Carolina at
Charlotte
Richard G. Merritt, Houston Community College
Andrew S. Methven, Eastern Illinois University
Timothy Metz, Campbell University
David H. Miller, Oberlin College
David W. Miller, Clark State Community College
Lillian W. Miller, Florida Community College–
Jacksonville
Subhash C. Minocha, University of New Hampshire
L. Maynard Moe, California State University–
Bakersfield
Beth Morgan, University of Illinois, Urbana-Champaign
Dale M. J. Mueller, Texas A & M University
Lytton John Musselman, Old Dominion University
Nusrat H. Naqvi, Southern University
Joanna H. Norris, University of Rhode Island
Chuks A. Ogbonnaya, Mountain Empire College
Jeanette C. Oliver, Flathead Valley Community College
Sebastine O. Onwuka, Lesley College
Clark L. Ovrebo, University of Central Oklahoma
A. D. Owings, Southeastern Louisiana University
Julie M. Palmer, University of Texas–Austin
Richard A. Palmer, Fresno City College
Carolyn Peters, Spoon River College
xiv Preface
Martha M. Phillips, The College of St. Catherine
Jerry L. Pickering, Indiana University of Pennsylvania
Wayne S. Pierce, California State University–Stanislaus
Indiren Pillay, Southwestern Tennessee Community
College
Mary Ann Polasek, Cardinal Stritch University
Dr. Robert J. Porra, CSIRO
Kumkum Prabhakar, Nassau Community College
Tyre J. Proffer, Kent State University
Francis Putz, University of Florida
V. Raghaven, The Ohio State University
Mohammad A. Rana, St. Joseph College
Margene M. Ranieri, Bob Jones University
W. T. Rankin, University of Montevallo
Dennis T. Ray, University of Arizona
Linda Mary Reeves, San Juan College
Maralyn A. Renner, College of the Redwoods
Penelope ReVelle, Community College of Baltimore
County–Essex
Tom Reynolds, University of North Carolina–Charlotte
Stanley A. Rice, Southeastern Oklahoma State
University
Dennis F. Ringling, Pennsylvania College of Technology
Daryl Ritter, Okaloosa-Walton Community College
Suzanne M. D. Rogers, Salem International University
Wayne C. Rosing, Middle Tennessee State University
Robert G. Ross, University of Puerto Rico
Jimmy Rozell, Tyler Junior College
Manfred Ruddat, University of Chicago
Patricia Rugaber, Coastal Georgia Community College
Frances Rundlett, Georgia State University
Robert M. Rupp, Ohio State University, Agricultural
Technical Institute
Thomas H. Russ, Charles County Community College
Dennis J. Russell, University of Alaska Southeast
Connie Rye, Bevill State Community College
C. L. Sagers, University of Arkansas
A. Edwards Salgado, Christian Brothers University
Thomas Sasek, Northeast Louisiana University
Michael A. Savka, University of West Florida
Neil W. Sawyer, University of Wisconsin–Whitewater
Neil Schanker, College of the Siskiyous
Renee M. Schloupt, Delaware Valley College
Bruce S. Serlin, DePauw University
Wilbur J. Settle, State University of New York–
Oneonta
Barbara Greene Shipes, Hampton University
Richard H. Shippee, Vincennes University
Brian R. Shmaefsky, Kingwood College
Shaukat M. Siddiqi, Virginia State University
Dilbagh Singh, Blackburn College
Del William Smith, Modesto Junior College
James Smith, Boise State University
Joanna M. K. Smith
Steven Smith, University of Arizona
Nancy Smith-Huerta, Miami University
F. Lee St. John, Ohio State University–Newark
Spencer S. Stober, Alvernia College
Marshall D. Sundberg, Emporia State University
Eric Sundell, University of Arkansas–Monticello
Donald D. Sutton, California State University–
Fullerton
Stan R. Szarek, Arizona State University
Mesfin Tadesse, Ohio State University
Max R. Terman, Tabor College
R. Dale Thomas, Northeast Louisiana University
Stephen L. Timme, Pittsburgh State University
Leslie R. Towill, Arizona State University
Richard E. Trout, Oklahoma City Community College
Jun Tsuji, Sienna Heights College
Claudia Uhde-Stone, California State University–
East Bay
Gordon E. Uno, University of Oklahoma
Rani Vajravelu, University of Central Florida
John Vanderploeg, Ferris State University
Staria Vanderpool, Arkansas State University
Delmar Vander Zee, Dordt College
C. Gerald Van Dyke, North Carolina State University
Carol Wake, South Dakota State University
Leon Walker, University of Findlay
Ami Lea Wangeline, Colorado State University
Betty J. Washington, Albany State University
Edgar E. Webber, Keuka College
Christopher R. Wenzel, Eastern Wyoming College
Cherie Wetzel, City College of San Francisco
Ingelia White, Windward Community College
Garrison Wilkes, University of Massachusetts–
Boston
Donald L. Williams, Sterling College
Justin K. Williams, Sam Houston State University
Marvin Williams, University of Nebraska–Kearney
Dwina W. Willis, Freed-Hardeman University
James A. Winsor, The Pennsylvania State University
Clarence C. Wolfe, Northern Virginia Community
College
Chris Wolverton, Ohio Wesleyan University
Kathleen Wood, University of Mary Hardin-Baylor
Richard J. Wright, Valencia Community College
Todd Christian Yetter, Cumberland College
Brenda Young, Daemen College
Rebecca Zamora, South Plains College
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Preface xv
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xvi Preface
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 C H A P T E R

What Is Plant Biology?

Overview
Learning Outcomes
The Relationship of Humans to Their
Environment
ECOLOGY: Who Needs Plants?
Human and Animal Dependence on Plants
Botany as a Science
Hypotheses
Diversification of Plant Study
GENERAL: Plant Biology and the Internet
SUMMARY
REVIEW QUESTIONS
DISCUSSION QUESTIONS
ADDITIONAL READING
LEARNING ONLINE

A mountain iris (Iris missouriensis) growing along a

slope near the roadside in the Carson National Forest,
New Mexico.
OVERVIEW

This chapter introduces you to botany: what it is, how it developed, how it relates to our everyday lives, and what its potential is for the future.
The discussion includes a brief introduction to some common questions about plants and their functions, an examination of the scientific
method, and a brief look at botany after the invention of the microscope. It concludes with a brief survey of the major disciplines within the
field of botany.

Learning Outcomes
1. Explain how humans have impacted their environment, 3. Explain how and why all life is dependent on green
particularly during the past century. organisms.
2. Describe how hypotheses are formulated and used in the 4. List the aspects of botany with which each of the major
scientific method. botanical disciplines is concerned.

W hile in high school in southern Africa, I was once

invited to a friend’s farm during spring break.
One day as I was returning to the farmhouse from
a walk around the farm, I heard groaning coming from inside.
I learned that my friend’s father had been clearing cactus-
like Euphorbia plants from some land. The plants produce
a poisonous, milky latex, which the father had taken great
care to wash thoroughly from his hands. Absentmindedly,
however, he had splashed some of the water in his face, and
traces of the poison had gotten into his eyes, causing great
pain. Another family member immediately ran to the nearby
barn and obtained some colostrum milk from a cow that had
just given birth. The eyes were bathed in the milk, which con-
tains an antidote for that poison, and the pain subsided. I was
told that if the milk had not been quickly available, the man
would have been blind within half an hour. In Venezuela and
Brazil, however, cow trees (e.g., Brosimum utile; Mimusops
huberi) produce a sweet, nutritive latex that is relished by the
natives of the region. Still other plants such as opium pop-
pies produce latex that contains narcotic and medicinal drugs
(Fig. 1.1). Why do plants such as Euphorbia species produce
poisons, while parts of so many other plants are perfectly
edible, and some produce spices, medicines, and a myriad of
products useful to humans?
In the popular story “Little Shop of Horrors,” Audrey II
is a plant that thrives on human blood. While this scenario Figure 1.1 Immature opium poppy capsules that were gashed with
a razor blade. Note the opium-containing latex oozing from the gashes.
makes for an interesting piece of dark comedy, there are
plants that feed on insects and occasionally frogs, rodents,
and birds (Fig. 1.2). What do these plants gain from their car- evidence of pumps of any kind within them. How, then, does
nivorous diet? water get from the roots below ground to the tops of these and
Occasionally we hear or read of experiments—often other trees? Do they have to expend large amounts of energy
associated with school science fairs—that suggest plants to carry water such long distances (Fig. 1.3)?
respond in some positive way to music or soothing talk; con- Our tropical rain forests, which once covered 14% of
versely, some plants are said to grow poorly when they are the earth’s land surface, now occupy only 6% of land area.
harshly yelled at. Do plants really respond to their surround- They are disappearing at the rate of several acres a minute as
ings, and, if so, how and to what extent? the plant life is cleared for agriculture, wood supplies (pri-
California’s huge coastal redwoods and Tasmania’s giant marily for fuel), cattle ranching, and other human activities
gum trees can grow to heights of 90 or more meters (300 or such as mining for gold. Experts estimate that all rain forests
more feet). When these giant trees are cut down, there is no may be destroyed within 40 years. Rain forests are home to

2

Figure 1.2 Pitcher plants. Pitcher plants are carnivorous plants
that grow in nutrient-poor environments. Small insects trapped in their
pitchers are digested and used as a source of minerals.
50% of all the species of living organisms; it is estimated
that 137 species are destroyed every day due to rain-forest
deforestation. What will be the long-term impacts of these
activities, and can they be reversed?
There is no doubt that our climate is changing. There is
much debate, though, about the effects that global warming
will have on life as we know it. Are those who proclaim that
global warming will eventually have disastrous effects on mod-
ern civilization and living organisms simply exaggerating, or
is there a scientific basis for the claims? Plant life constitutes
more than 98% of the total biomass (collective dry weight of
living organisms) of the earth. Plants and other green organisms
have the exclusive capacity to produce oxygen while convert-
ing the sun’s energy into forms vital to the existence of both
plant and animal life. At the same time, plants remove the large
amounts of carbon dioxide given off by all living organisms
as they respire. In other words, virtually all living organisms
are totally dependent on green organisms for their existence.
If some major disease were to kill off all or most of the green
organisms on land and in the oceans and lakes, all the animals
on land, in the sea, and in the air would soon starve. Even if
some alternative source of energy were available, animal life
would suffocate within 11 years—the time estimated for all the
earth’s oxygen to be completely used up if it were not replaced.
Just how do green plants capture the sun’s energy, use carbon
dioxide, and give off oxygen?
This book tries to answer these and other questions about
living organisms—particularly those pertaining to plants,
algae, fungi, and bacteria. Moreover, additional information
about plant biology related to future societies, conservation,
and human benefits is discussed.
What Is Plant Biology? 3
Figure 1.3 California coastal redwoods (Sequoia sempervirens).
Coastal redwoods may grow for thousands of years and some reach
heights of nearly 100 meters (330 feet).
The Relationship of Humans
to Their Environment
It has been estimated that the total human population of the
world was less than 20 million in 6000 b.c. During the next
7,750 years, it rose to 500 million; by 1850, it had doubled
to 1 billion; and 70 years later, it had doubled again to 2 bil-
lion. The 4.48-billion mark was reached in 1980, and within
5 years it had grown to 4.89 billion. In 2011, the human pop-
ulation size reached a milestone, exceeding 7 billion people.
The earth remains constant in size, but the human popula-
tion continues to grow.
In feeding, clothing, and housing ourselves, we have
had a major impact on our environment. We have drained
wetlands and cleared natural vegetation from vast areas of
land. California, for example, now has less than 5% of the
wetland it had 100 years ago. We have dumped wastes and
other pollutants into our waters and added pollutants to the
atmosphere. We have killed pests and plant disease organ-
isms with poisons. These poisons have also killed natural
4 Chapter 1
KEY THEME : ecology
predators and other useful organisms and, in general, have
disrupted the delicate balance of nature that existed before
humans began degrading their natural surroundings.
If we are to survive on this planet beyond the 21st cen-
tury, there is little question that humans have to stop increas-
ing in numbers, and the many unwise agricultural and
industrial practices that have accompanied the mushrooming
of human populations must be replaced with practices more
in tune with restoring some ecological balance. Agricultural
practices of the future will have to include the return of
organic material to the soil after each harvest, instead of add-
ing only inorganic fertilizers. Harvesting of timber and other
crops will have to be done in a manner that prevents topsoil
erosion, and the practice of clearing brush with chemicals will
have to be abolished. Industrial pollutants will have to be ren-
dered harmless and recycled whenever possible.
Many products that now are still largely discarded
(e.g., garbage, paper products, glass, metal cans) will also
have to be recycled on a much larger scale. Biological pest
controls (discussed in Appendix 2) will have to replace
the use of poisonous controls whenever possible. Water
and energy conservation will have to be universally prac-
ticed, and rare plant species, with their largely unknown
gene potential for future crop plants, will need to be saved
from extinction by preservation of their habitats and by
other means. The general public will have to be made even
more aware of the urgency for wise land management and
conservation—which will be especially needed when pres-
sures are exerted by influential forces promoting unwise
measures in the name of “progress”—before additional
large segments of our natural resources are irreparably
damaged or lost forever. Alternatives appear to be nothing
less than death from starvation, respiratory diseases, poi-
soning of our food and drink, and other catastrophic events
that could ensure the premature demise of large segments
of the world’s population.
Scientists and, increasingly, the general public have
become alarmed about the effects of human carelessness
on our environment. Damage to forests and lakes caused
by acid rain, contamination of ground water by nitrates and
pesticides, reduction of the ozone shield, major global cli-
matic changes, depletion of aquifers, and loss of biodiversity
have gained widespread publicity.
Human and Animal Dependence
on Plants
Our dependence on green organisms to produce the oxy-
gen in the air we breathe and to remove the carbon dioxide
we give off doesn’t stop there. Plants are also the sources
of products that are so much a part of human society that
we largely take them for granted. We know, of course, that
wheat, rice, corn, potatoes, and other vegetables are plants
(Fig. 1.4); but all foods, including meat, fish, poultry, eggs,
cheese, and milk, to mention just a few, owe their existence
to plants. Condiments such as spices (Fig. 1.5) and luxuries
such as perfumes are produced by plants, as are some dyes,
adhesives, digestible surgical stitching fiber, food stabiliz-
ers, beverages (Fig. 1.6), and emulsifiers.
Our houses are constructed with lumber from trees,
which also furnish the cellulose for paper, cardboard, and syn-
thetic fibers. Some of our clothing, camping equipment, bed-
ding, draperies, and other textile goods are made from fibers
of many different plant families (Fig. 1.7). Coal is fossilized
plant material, and oil came from microscopic green organ-
isms or animals that either directly or indirectly were plant
consumers. All medicines and drugs at one time came from
plants, fungi, or bacteria, and many important ones, includ-
ing most of the antibiotics, still do (Fig. 1.8). Microscopic
organisms play a vital role in recycling both plant and ani-
mal wastes and aid in the building of healthy soils. Others are
responsible for human diseases and allergies.
Although shortages of oil and other fossil fuels may
sometimes be politically or economically manipulated,
there is no question that these fuels are finite and eventually
will disappear. Accordingly, the development of alternative
energy sources is receiving increased attention.
Methane gas, which can be used as a substitute for natu-
ral gas, has been produced from animal manures and decom-
posed plants in villages in India and elsewhere for many
years, and after several years of trial on a small scale in the
United States, the production of methane on a larger scale
from human sewage is being investigated.
The United States is the largest producer of ethanol in
the world. Corn, switchgrass, and other sources of carbohy-
drates are currently used in the manufacture of ethanol, which

(a)
(b)
Figure 1.4 Rice cakes being manufactured. (a) Unprocessed rice is
poured into small ovens, where the kernels are expanded. The kernels are
then compressed into cakes, which are conveyed by belt to a packaging area.
(b) Part of a produce section in a grocery store.
What Is Plant Biology? 5
Figure 1.5 Some of the spices derived from plants.
is blended with gasoline. Most cars in the United States can run
on fuel containing up to 10% ethanol. Flexible fuel vehicles
have been designed to use fuel blends containing up to
85% ethanol. In 2010, over 200 ethanol plants in the United
States produced 13.2 billion gallons of ethanol, up from
1.6 billion gallons in 2000. The Energy Independence and
Security Act of 2007 calls for 39 billion gallons of renew-
able fuel to be used annually in the United States. Currently,
ethanol fuel in the United States is mainly produced from
corn, but there are concerns about losing food crop land to
produce fuel. In addition, the energy and pollution balance
of ethanol production is under debate. Cellulosic ethanol,
which is derived from inedible plant fiber such as wood chips
or switchgrass may overcome some of these concerns.
What about plants and the future? As you read this,
the population of the earth already has exceeded 7 billion
persons, every one of whom needs food, clothing, and shel-
ter in order to survive. To ensure survival, we may need to
learn not only how to cultivate food plants but also how to
use plants to remove pollutants from the water, air, and soil
(Fig. 1.9), to make land productive again, and to renew
urban areas. In addition, we need to minimize the destruc-
tion of plant habitats caused primarily by the huge increase
in the number of earth’s inhabitants. This subject and related
matters are further discussed in Chapter 25.
At present the idea that humanity may not be able to save
itself may seem radical, but there are a few who have sug-
gested that it might become necessary in the future to emi-
grate to other planets. Regardless of humanity’s future, it is
essential that our understanding of plants be used to sustain
life on this and maybe even other planets. Experiments with
portable oxygen generators have been in progress for many
years. Tanks of water teeming with tiny green algae are taken
aboard a spacecraft and installed so that they are exposed to
light for at least part of the time. The algae not only produce
oxygen, which the spacecraft inhabitants can breathe, but
also utilize the waste carbon dioxide produced by respiration.
As the algae multiply, they can be fed to a special kind of
shrimp, which in turn multiply and become food for the space
6 Chapter 1
(a)
(b)
Figure 1.6 Ripening coffee berries. (a) Berries are picked by hand
when they are red. The seeds are extracted for roasting after the berries
are fermented. (b) Coffee beans cooling after being roasted.
Figure 1.7 Cotton plants. The white fibers, in which seeds are
embedded, are the source of textiles and fabrics. The seeds are the
source of vegetable oils used in margarine and shortening. After
the oils have been extracted, the remaining “cotton cake” is used for
cattle feed.
Figure 1.8 A Penicillium colony. The tiny beads of fluid on the
surface contain penicillin, widely used as an antibiotic.
travelers. Other wastes are recycled by different microscopic
organisms. When this self-supporting arrangement, called
a closed system, is perfected, the range of spacecraft should
greatly increase because heavy oxygen tanks will not be nec-
essary, and the amount of food reserves will be reduced.
Today, teams of botanists, anthropologists, and medi-
cal doctors are interviewing medical practitioners and
herbal healers in remote tropical regions and taking notes
on various uses of plants by the local inhabitants. These
scientists are doing so in the hope of preserving at least
some plants with potential for contributions to modern civi-
lization before disruption of their habitats results in their
extinction.

Figure 1.9 A polluted waterway in an urban area.
Botany as a Science
The study of plants, called botany—from three Greek words,
botanikos (botanical), botane (plant or herb), and boskein (to
feed), and the French word botanique (botanical)—appears
to have had its origins with Stone Age peoples who tried to
modify their surroundings and feed themselves. At first, their
interest in plants was mostly practical and centered around
how plants might provide food, fibers, fuel, and medicine.
Eventually, however, an intellectual interest arose. Individuals
became curious about how plants reproduced and how they
were put together. This inquisitiveness led to plant study
becoming a science, which, broadly defined, is simply “a
search for knowledge of the natural world.” Botanists are
scientists who study plants.
A science is distinguished from other fields of study by
several features. It involves the observation, recording, orga-
nization, and classification of facts, and more important, it
involves what is done with the facts. Scientific procedure
involves the process of experimentation, observation, and
the verifying or discarding of information, chiefly through
inductive reasoning from known samples. There is no uni-
versal agreement on the precise details of the process. A few
decades ago, scientific procedure was considered to involve
a routine series of steps that involved first asking a ques-
tion, then formulating a hypothesis, followed by conducting
experiments, and finally developing a theory. This series of
steps came to be known as the scientific method, and there are
still instances where such a structured approach works well.
In general, however, the scientific method now describes the
procedures of developing and testing hypotheses.
Hypotheses
A hypothesis is simply a tentative, unproven explanation for
something that has been observed. It may not be the correct
explanation—testing will determine whether it is correct
What Is Plant Biology? 7
or incorrect. To be accepted by scientists, the results of any
experiments designed to test the hypothesis must be repeat-
able and capable of being duplicated by others.
The nature of the testing will vary according to the cir-
cumstances and materials, but good experiments are run in
two forms, the second form being called a control. In the
first form, a specific aspect, or variable, is changed. The
control is run in precisely the same way but without chang-
ing the specific aspect, or variable. The scientist then can be
sure that any differences in the results of the parallel experi-
ments are due to the change in the variable.
For example, we may observe that a ripe orange we
have eaten tastes sweet. We may then make the hypothesis
that all ripe citrus fruits taste sweet. We may test the hypoth-
esis by tasting oranges and other citrus fruits such as tan-
gerines and lemons. As a result of our testing (since lemons
taste sour), we may modify the hypothesis to state that only
some ripe citrus fruits are sweet. In such an experiment, the
variable involves more than one kind of ripe citrus fruit; the
control, on the other hand, involves only ripe oranges.
When a hypothesis is tested, data (bits of information)
are accumulated and may lead to the formulation of a useful
generalization called a principle. Several related principles
may lend themselves to grouping into a theory, which is
not simply a guess. A theory is a group of generalizations
(principles) that help us understand something. We reject
or modify theories only when new principles increase our
understanding of a phenomenon.
While the testing of hypotheses is a rigorous, well-
defined process, the development of hypotheses is not. This
aspect of science is creative because it involves finding new
ways to look at our natural world. Often, the most successful
scientists are those who can think “outside the box,” inte-
grating observations across fields or organisms to create a
hypothesis that explains a complex phenomenon.
Diversification of Plant Study
Plant anatomy, which is concerned chiefly with the inter-
nal structure of plants, was established through the efforts
of several scientific pioneers. Early plant anatomists of note
included Marcello Malpighi (1628–1694) of Italy, who dis-
covered various tissues in stems and roots, and Nehemiah
Grew (1628–1711) of England, who described the structure of
wood more precisely than any of his predecessors (Fig. 1.10).
Today, knowledge of plant anatomy is used to help us
find clues to the past, as well as for many practical purposes.
For example, the related discipline of dendrochronology
deals with determining past climates by examining the width
and other features of tree rings. We can also learn much from
archaeological sites by matching tree rings found in the
wood of ancient buildings to the rings of wood of known age.
Plant anatomy is also used to solve crimes. Forensic labo-
ratories may use fragments of plant tissues found on cloth-
ing or under fingernails to determine where a crime took
place or if certain persons could have been present where the
8 Chapter 1
Figure 1.10 A thin section of Magnolia wood as seen through
a light microscope. ⫻40.
crime was committed. The anatomy of leaves, stems, and
other plant parts is used to unravel and sort out relationships
among plants. A form of plant anatomy, known as paleobot-
any, involves the study of plant fossils.
Plant physiology, which is concerned with plant func-
tion, was established by J. B. van Helmont (1577–1644),
a Flemish physician and chemist, who was the first to
demonstrate that plants do not have the same nutritional
needs as animals. In a classic experiment, van Helmont
planted a willow branch weighing 5 pounds in an earth-
enware tub filled with 90.7 kilograms (200 pounds) of dry
soil. He covered the soil to prevent dust from settling on it
from the air. The willow produced roots and grew, and after
5 years, he reweighed the willow and the soil. He found that
the soil weighed only 56.7 grams (2 ounces), less than it
had at the beginning of the experiment, but that the willow
had gained 76.7 kilograms (169 pounds). He concluded that
the tree had added to its bulk and size from the water it had
absorbed. We know now that most of the weight came as a
result of photosynthetic activity (discussed in Chapter 10),
but van Helmont deserves credit for landmark experimenta-
tion in plant physiology.
Modern plant physiologists use cloned genes (units of
heredity that are found mostly within the nuclei of cells)
to learn in precise detail much more about plant functions,
including how plants conduct materials internally; how tem-
perature, light, and water are involved in growth; why plants
flower; and how plant growth regulatory substances are pro-
duced, to mention just a few.
During past centuries, Europeans who explored other
continents took large numbers of plants back home with
them, and it soon became clear to those working with the
plants that some sort of formalized system was necessary
just to keep the collections straight. Several plant taxono-
mists (botanists who specialize in the identifying, naming,
and classifying of plants) proposed ways of accomplishing
this, but we owe much of our present system of naming and
classifying plants to the Swedish botanist Carolus Linnaeus
(1707–1778) (see Fig. 16.2).
Plant taxonomy involves describing, naming, and
classifying organisms. Plant systematics is a related field
but is broader than taxonomy. It is the science of devel-
oping methods for grouping organisms. Plant taxonomy
is the oldest branch of plant study, begun in antiquity, but
Linnaeus did more for the field than any other person in
history. Thousands of plant names in use today are those
originally recorded in Linnaeus’s book Species Plantarum,
published in 1753. An expanded account of Linnaeus and
his system of classification is given in Chapter 16.
There are still thousands of plants, fungi, and other
organisms that have not yet been described or even discov-
ered. Although it obviously is already too late to identify
species that were not described before they became extinct,
plant taxonomists around the world have united to try to
identify and describe as many new organisms as possible—
many with food, medicinal, and other useful potential—
before much more of their natural habitat disappears. Other
plant taxonomists, through the use of cladistics (analysis
of shared features) and molecular techniques, are refining
our knowledge of plant relationships. By the year 2000 we
had acquired so much new information about natural rela-
tionships that some major reclassification took place (see
Chapter 16). The molecular knowledge and techniques are
also contributing to the improvement of many of our food
crops, although some of the changes are controversial.
Many plant taxonomists specialize in certain groups of
plants. For example, pteridologists specialize in the study of
ferns; bryologists study mosses and plants with similar life
cycles.
The discipline of plant geography, the study of how
and why plants are distributed where they are, did not
develop until the 19th century (Fig. 1.11). The allied field of
plant ecology, which is the study of the interaction of plants
with one another and with their environment, also developed
in the 19th century.
After the publication in 1962 of a best-seller entitled
Silent Spring (authored by Rachel Carson), public aware-
ness of the field of ecology as a whole increased consider-
ably. In this book, based on more than 4 years of literature
research, Carson noted that more than 500 new toxic chemi-
cals are put to use annually as pesticides in the United States
alone, and she detailed how these chemicals and other pol-
lutants are having a negative impact on all facets of human
life and the environment.
The study of the form and structure of plants, plant mor-
phology, was developed during the 19th century, and during
the 20th century much of our basic knowledge about the form
and life cycles of plants was incorporated into the plant sci-
ences as we know them today. During this time, the number of
scientists engaged in investigating plants also greatly increased.
Genetics, the science of heredity, was founded by the
Austrian monk Gregor Mendel (1822–1884), who performed
 What Is Plant Biology? 9

classic experiments with pea plants. Today, various bran-

Figure 1.11 Plant ecologists, plant geographers, and other
biologists recognize large communities of plants and animals that
occur in areas with distinctive combinations of environmental
features. These areas, called biomes, are represented here by the tropical
rain forest, which, although occupying less than 5% of the earth’s surface,
is home to more than half of the world’s species of organisms.
ches of genetics include plant breeding, which has greatly
improved the yields and quality of crop plants, and genetic
engineering. Genetic engineering involves the transfer
of genes from one organism to another and has already
improved the pest, frost, and disease resistance of some
crop plants. Genetic engineering is being used to develop
better agricultural, medicinal, and other useful plants. It is
also being used to control human, animal, and plant dis-
eases. In recent years, the field of genomics, which focuses
on genes and their function, has burst onto the genetics
scene and now impacts nearly all genetics research.
Cell biology (previously called cytology), the sci-
ence of cell structure and function, received a boost from
the discovery of how cells multiply and how their various
components perform and integrate a variety of functions,
including that of sexual reproduction. The mid-20th-century
development of electron microscopes (see Chapter 3) further
spurred cell research and led to vast new insights into cells
and new forms of cell research that continues to the present.
Economic botany and ethnobotany, which focus on
practical uses of plants and plant products, had their origin in
antiquity as humans discovered, used, and eventually cultivated
plants for food, fiber, medicines, and other purposes. Today
there is increased interest in herbal medicines (see Appendix 3)
and many other uses of plants by the general public. Research
is being conducted in collaboration with indigenous peoples
with an eye to discovering new medicines and other useful
plant products previously unknown in developed countries.
There is still a vast amount of botanical information to
be discovered. Dozens of research journals publish thousands
of botanical scientific papers every year. In recent years, open
source journals have allowed free online access to research
findings for scientists all over the world. While scientists
have identified about 2 million species of organisms, the
earth likely contains between 5 and 100 million species.

Plant Biology
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10 Chapter 1
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Box Figure 1.1 A botany student uses the Internet to identify and
learn about a botanical specimen in the field.
A variety of communication tools are available through
the Internet, including e-mail, texting, and video streaming, as
well as various software applications such as Facebook, MySpace,
and Twitter. Additional applications that have become popular
include those that provide instant messaging, such as Windows
Live Messenger, Yahoo Messenger, and Google Talk, and those that
allow downloading videos on a variety of topics such as YouTube.
Among the most widely used aspects of the Internet are those that
access the World Wide Web (also referred to as the Web), which can
be tapped into through Internet browsers such as Internet Explorer,
Firefox, and Netscape. With these programs, a user can find informa-
tion on almost any topic through the use of a specific web address
called a universal resource locator (URL), and then start exploring.
Client software interprets the URL address and initiates communi-
cation with a specified server. For instance, the following URL is for
an image collection sponsored by the Botanical Society of America:
http;//www.botany.org/plantimages/PlantAnatomy.php
1. http is the acronym for hyper text transfer protocol and is used
by the client and server to communicate with each other.
2. www.botany.org is the address of the server (and the domain
name for the Botanical Society of America).
3. plantimages is a directory containing the files, and
PlantAnatomy.php is the file name of the home page where
images for plant anatomy can be found.
What if a user wants to search the Internet for a specific
topic? There are many search engines available that allow users
to search by key word(s). The search software scans numerous
Web servers for key words and returns any number of hits, or
positive matches. A user can then navigate to any of the matches
by clicking the mouse pointer on the hyperlinked search results.
One of the most widely used search engines is called Google
(http://www.google.com), although there are many others,
including Yahoo (http://www.yahoo.com), Bing (http://www.bing
.com), Ask (http://www.ask.com) and AOL Search (http://search
.aol.com/aol/webhome).
What botanical information is available on the Internet? Just
about anything a user can imagine about plant biology is available
on the Internet and, if it isn’t, it can be added. Sites that include
materials on botany education, botanical gardens, plant databases,
and videos about plants are a few of the many resources available
on the Web. The following are some interesting websites that can
be explored. Try them out sometime! Maybe you’ll find a good idea
for a research paper.
1. The Multimedia Educational Resource for Learning and Online
Teaching (MERLOT) includes many learning objects for plant
biology and other disciplines. This website includes a search
engine as well as peer-reviewed learning materials for many
topics in higher education. http://www.merlot.org/
2. The Missouri Botanical Garden is one of the oldest botanical
institutions in the United States. It is a center for botanical
research and science education. http://www.mobot.org/
3. The New York Botanical Garden is situated on 250 acres in
the Bronx and includes 27 outdoor gardens and plant collec-
tions, the nation’s most beautiful Victorian conservatory, and a
40-acre presettlement forest. http://www.nybg.org/
4. The Arnold Arboretum of Harvard University is the nation’s
oldest arboretum. The site includes a catalog of over 5,000
kinds of woody plants cultivated in the arboretum as well as
educational and visitor information. http://www.arboretum
.harvard.edu/
5. Australian National Botanic Gardens provides a wealth of
botanical and biological information about Australia. http://
www.anbg.gov.au/gardens/
6. The United States Department of Agriculture contains news
and information about the nation’s agricultural economy.
There is an excellent section on the history of American agri-
culture from 1776 to 2000. http://www.usda.gov/
7. GardenWeb is an information center for gardening enthusi-
asts. http://www.gardenweb.com/
8. California Flora Database contains geographic and ecological
distribution information for 7,975 California vascular plant taxa,
as well as additional habitat information for rare taxa and spe-
cies of the Sierra Nevada. http://www.calflora.org/
9. Common Conifers of the Pacific Northwest provides informa-
tion about the conifers of Oregon, including a dichotomous
key for their identification. http://www.oregonstate.edu/trees/
10. Carnivorous Plants Database includes over 3,000 entries, giv-
ing an exhaustive nomenclatural synopsis of all carnivorous
plants. http://www.omnisterra.com/bot/cp_home.cgi
11. Plant in Motion provides short video clips of different types of
plant movement, including germination, flowering, and tropic
responses. http://plantsinmotion.bio.indiana.edu/

SUMMARY
1. Why do some plants produce poisons, while others are
edible and useful? Why are some plants carnivorous?
How and why do plants respond to their environment?
What is the future of tropical rain forests? What can be
done about global warming and other environmental
problems? This book addresses these questions and
more about plants.
2. Human populations have increased dramatically in the
past few centuries, and the disruption of the balance of
nature by the activities directly or indirectly associated
with the feeding, clothing, and housing of billions of
people threatens the survival of not only humans but
other organisms as well.
3. We are totally dependent on green organisms because
they alone can convert the sun’s energy into forms
that are usable by, and vital to the very existence of,
animal life.
4. We largely take plants and plant products for granted.
Animals, animal products, many luxuries and
condiments, and other useful substances such as fibers,
lumber, coal, medicines, and drugs either depend on
plants or are produced by them.
5. To ensure human survival, all persons soon may need
to acquire some knowledge of plants and how to use
them. Plants will undoubtedly play a vital role in space
exploration as portable oxygen generators.
6. Teams of scientists are interviewing medical
practitioners and herbal healers in the tropics to locate
little-known plants used by local inhabitants before the
plants become extinct.
7. Botany, the study of plants, apparently began with
Stone Age peoples’ practical uses of plants. Eventually,
botany became a science as intellectual curiosity about
plants arose.
8. A science involves the observation, recording,
organization, and classification of facts. The verifying
or discarding of facts is done chiefly from known
samples. The scientific method involves following
a routine series of steps and generally assuming and
testing hypotheses.
9. The microscope has had a profound effect on studies
in the biological sciences and led to the discovery
of cells.
10. Plant anatomy and plant physiology developed during
the 17th century. J. B. van Helmont was the first to
demonstrate that plants have nutritional needs different
from those of animals. During the 17th century,
Europeans engaged in botanical exploration on other
continents and took plants back to Europe.
What Is Plant Biology? 11
11. During the 18th century, Linnaeus produced the
elements of a system of naming and classifying
plants. In recent years, molecular and cladistical
investigations have resulted in modifications of
Linnaeus’s system.
12. During the 19th century, plant ecology, plant
geography, and plant morphology developed, and by
the beginning of the 20th century, genetics and cell
biology became established. Much remains yet to be
discovered and investigated.
REVIEW QUESTIONS
1. How and to what extent have humans affected their
natural environment?
2. What is meant by the scientific method?
3. To what extent is animal life dependent on green
organisms for its existence?
4. In terms of biological experiments, what are hypotheses
and controls?
5. What is the oldest branch of botany, and why did it
precede other branches?
6. What are the basic features of each of the other
branches of botany?
DISCUSSION QUESTIONS
1. Since humans survived on wild plants for thousands of
years, might it be desirable to return to that practice?
2. What factors are involved in determining if and when
humans might not be able to sustain themselves on this
planet?
3. How do you suppose that Stone Age peoples discovered
medicinal uses for plants?
4. Why do you suppose that many of the early botanists
were also medical doctors?
5. Consider the following hypothesis: “The majority
of mushrooms that grow in grassy areas are not
poisonous.” How could you go about testing this
hypothesis scientifically?
ADDITIONAL READING
Capon, B. 2010. Botany for gardeners. 3d ed. Portland, OR:
Timber Press.
Carson, R. L. 2002. Silent spring. Boston: Houghton Mifflin
Harcourt.
Coyle, H. M. 2004. Forensic botany. New York: Taylor and Francis.
12 Chapter 1
Jacobs, P. F., and J. Brett. 2004. Guide to information sources
in the botanical sciences. Englewood, CO: Libraries
Unlimited.
Johnson, T. 1998. CRC ethnobotany desk reference. Boca Raton,
FL: CRC Press.
Minnus, P. E. 2000. Ethnobotany. Norman: University of
Oklahoma Press.
Pollan, M. 2002. The botany of desire. New York: Random House.
Pollan, M. 2007. The omnivore’s dilemma: A natural history of
four meals. New York: Penguin Group.
Sumner, J. 2004. American household botany: A history of useful
plants, 1620–1900. Portland, OR: Timber Press.
van Wyk, B. 2005. Food plants of the world: An illustrated guide.
Portland, OR: Timber Press.
LEARNING ONLINE
Visit our website at http://www.mhhe.com/stern13e for
additional information and learning tools.
 C H A P T E R

The Nature of Life

Overview
Learning Outcomes
Attributes of Living Organisms
Composition and Structure
Growth
Reproduction
Response to Stimuli
Metabolism
Movement
Complexity of Organization
Adaptation to the Environment
Chemical and Physical Bases of Life
The Elements: Units of Matter
Molecules: Combinations of Elements
Valence
Bonds and Ions
Acids, Bases, and Salts
The pH Scale
Energy
Chemical Components of Cells
Monomers and Polymers
MOLECULAR: The Skinny on Fats
SUMMARY
REVIEW QUESTIONS
DISCUSSION QUESTIONS
ADDITIONAL READING
LEARNING ONLINE

Lavender (Lavandula augustifolia) field located

in southern France. Lavender is used in soaps, in
shampoos, and as an aromatic plant because of its
soothing fragrance.
OVERVIEW

This chapter begins with a discussion of the attributes of living organisms. These include growth, reproduction, response to stimuli, metabo-
lism, movement, complexity of organization, and adaptation to the environment. Then it examines the chemical and physical bases of life. A
brief look at the elements and their atoms is followed by a discussion of compounds, molecules, valence, bonds, ions, acids, bases, and salts.
Forms of energy and the chemical components of cells are examined next. The chapter concludes with an introduction to macromolecules:
carbohydrates, lipids, proteins, and nucleic acids.

Learning Outcomes
1. List the attributes of living organisms.
2. Define matter and describe its basic state.
3. Describe the features of compounds, acids, bases, and
salts.
4. Describe the various forms of energy.
5. List the chemical elements found in cells.
6. Describe the main properties and functions of carbohydrates,
lipids, proteins, and nucleic acids.

H Growth
ave you ever dropped a pellet of dry ice (frozen car-
bon dioxide) into a pan of water and watched what
happens? The solid pellet darts randomly on the sur-
face, looking like a highly energetic water bug, as the warm-
er water rapidly converts it to a gas. Does all that motion
make the dry ice alive? No, although one of the attributes
of living things is the capacity to move. But if living things
can move, what about plants? If a tree is not able to crawl
down the sidewalk, does that mean it isn’t alive? Again the
answer is no, but these questions do point out some of the
difficulties encountered in defining life. In fact, some argue
that there is no such thing as life—only living organisms—
and that life is a concept based on the collective attributes
of living organisms.
Attributes of Living
Organisms
Some have described growth as simply an increase in mass
(a body of matter—the basic “stuff” of the universe), usu-
ally accompanied by an increase in volume. Most growth
results from the production of new cells and includes varia-
tions in form—some the result of inheritance, some the result
of response to the environment. If you plant two varieties of
tulips near each other and grow them under identical condi-
tions, they are likely to differ in size, color, and other charac-
teristics due to differences in genetic makeup. On the other
hand, if you plant bulbs of the same variety next to each other,
they may also look different from each other, especially if
you treat them differently. That is, they are exposed to differ-
ent environments. If you water one just enough to allow it to
grow, while you water the other one freely and work fertiliz-
ers and conditioners into the soil around it, you might expect
the second one to grow larger and produce more flowers than
the first. Growth pattern, therefore, is controlled by both a
plant’s genetic makeup and the environment in which it is
grown. Various aspects of growth are discussed in Chapter 11.

Composition and Structure

The activities of living organisms originate in tiny structural
units called cells, which consist of cytoplasm (a souplike
fluid) bounded by a very thin membrane. All living cells
contain genetic material that controls their development and
activities. In the cells of most organisms, this genetic mate-
rial, known as DNA (deoxyribonucleic acid), is housed in a
spherical structure called the nucleus, which is suspended in
the cytoplasm. In bacteria, however, the DNA is distributed
directly in the cytoplasm. The cells of plants, algae, fungi,
and many simpler organisms have a cell wall outside of the
membrane that bounds the cytoplasm. The cell wall pro-
vides support and rigidity. Cells are discussed in more detail
in Chapter 3.
Reproduction
Dinosaurs were abundant 160 million years ago, but none
exist today. Hundreds of mammals, birds, reptiles, plants,
and other organisms are now listed as endangered or threat-
ened species, and many of them will become extinct within
the next decade or two. All of these once-living or currently
threatened organisms have one feature in common: it became
impossible or it has become difficult for them to reproduce.
Reproduction is such an obvious feature of living organisms
that we take it for granted—until it no longer takes place.
When organisms reproduce, the offspring always
resemble the parents: guppies never have puppies—just
more guppies—and a petunia seed, when planted, will

14

not develop into a pineapple plant. Also, offspring tend to
resemble their parents more than they do other individuals
of the same kind. The laws governing these aspects of inher-
itance are discussed in Chapter 13.
Response to Stimuli
If you stick a pin into a pillow, you certainly don’t expect any
reaction from the pillow, but if you stick the same pin into a
friend, you know your friend will react immediately, because
responding to stimuli is a major characteristic of all living
things. You might argue, however, that when you stuck a pin
into your house plant, nothing happened, even though you
were fairly certain the plant was alive. You might not have been
aware that the house plant did indeed respond, but in a manner
very different from that of a human. Plant responses to stimuli
are generally of a different nature than those of animals. If the
house plant’s food-conducting tissue was pierced, it probably
responded by producing a plugging substance called callose in
the affected cells. Some studies have shown that callose may
form within as little as 5 seconds after wounding. Also, an unor-
ganized tissue called callus, which forms much more slowly,
may be produced at the site of the wound. Responses of plants
to injury and to other stimuli, such as light, temperature, and
gravity, are discussed in Chapters 9 through 11.
Metabolism
Metabolism is the collective product of all the biochemical
reactions taking place within an organism. All living organ-
isms undergo metabolic activities, which include the produc-
tion of new cytoplasm, the repair of damage, and normal cell
maintenance. The most important activities include respira-
tion, an energy-releasing process that takes place in all liv-
ing things; photosynthesis, an energy-harnessing process in
green cells that is, in turn, associated with energy storage;
digestion, the conversion of large or insoluble food molecules
to smaller soluble ones; and assimilation, the conversion of
raw materials into cytoplasm and other cell substances. These
topics are discussed in Chapters 9 through 11.
Movement
At the beginning of this chapter, we mentioned that plants gen-
erally don’t move from one place to another (although their
reproductive cells may do so). This does not mean, however,
that plants do not exhibit movement, a universal characteristic
of living things. The leaves of sensitive plants (Mimosa pudica)
fold within seconds after being disturbed or subjected to sudden
environmental changes, and the tiny underwater traps of blad-
derworts (Utricularia) snap shut in less than one-hundredth
of a second. But most plant movements, when compared with
those of animals, are slow and imperceptible and are mostly
related to growth phenomena. They become obvious only when
demonstrated experimentally or when shown by time-lapse
photography. Time-lapse photography often reveals many
The Nature of Life 15
types and directions of motion, particularly in young organs.
Movement is not confined to the organism as a whole but
occurs at the cellular level. For example, the cytoplasm of living
cells constantly flows like a river within cells; this streaming
motion is called cyclosis, or cytoplasmic streaming. Cyclosis
usually appears to run clockwise or counterclockwise within
the boundaries of each cell, but movement is not limited to a
circular pattern.
Complexity of Organization
The cells of living organisms are composed of large numbers
of molecules (the smallest unit of an element or compound
retaining its own identity). Even the most complex nonliving
object has only a tiny fraction of the types of molecules of the
simplest living organism. Typically there are more than 1 tril-
lion molecules in a single cell. The molecules are not simply
mixed, like the ingredients of a cake or the concrete in a side-
walk, but are organized into compartments, membranes, and
other structures within cells. Furthermore, the arrangements
of these molecules in living organisms are highly structured
and complex. Bacteria, for example, are considered to have
the simplest cells known, yet each cell contains a minimum
of 600 different kinds of protein as well as hundreds of other
substances, with each component playing an important role
in the function of the cell. When flowering plants and other
larger living objects are examined, the complexity of organi-
zation is overwhelming, and the number of molecule types
can run into the millions.
Adaptation to the Environment
If you move a rock from a cold mountain to a warm desert,
the structure of that rock will not change in response to its
new environment. Living organisms, however, do respond
to the air, light, water, and soil of their environment, as will
be explained in later chapters. They are also, after countless
generations of natural selection (as discussed in Chapter 15),
genetically adapted to their environment in many subtle ways.
Some weeds (e.g., dandelions) can thrive in a wide variety of
soils and climates, whereas many species now threatened with
extinction have adaptations to their environment that are so
specific they cannot tolerate even relatively minor changes.
Chemical and Physical
Bases of Life
The Elements: Units of Matter
The basic “stuff of the universe,” called matter, occurs in
three states—solid, liquid, and gas. In simple terms, matter’s
characteristics are as follows:
1. It occupies space.
2. It has mass, which we commonly associate with weight.
16 Chapter 2
3. It is composed of elements. There are 92 elements
that occur naturally on our planet. At least 29 more
elements have been produced artificially. Only a few
of the natural elements (e.g., nitrogen, oxygen, gold,
silver, copper) occur in pure form; the others are found
combined together chemically in various ways. Each
element has a designated symbol, often derived from its
Latin name. The symbol for copper, for example, is Cu
(from the Latin cuprum); and for sodium, Na (from the
Latin natrium). The symbols for carbon, hydrogen, and
oxygen are C, H, and O, respectively.
The smallest stable subdivision of an element that can
exist is called an atom. Atoms are so minute that until the mid-
l980s, individual atoms were not directly visible to us with even
the most powerful electron microscopes. Atoms consist of sev-
eral kinds of subatomic particles. Each atom has a tiny nucleus
consisting of protons, which are particles with positive electri-
cal charges, and other particles called neutrons, which have
no electrical charges. Both protons and neutrons have a small
amount of mass and are composed of quarks. If the nucleus,
which contains nearly all of the atom’s mass, were enlarged
so that it was as big as a beach ball, the atom, which is mostly
space, would be larger than a professional football stadium
(Fig. 2.1). Because each atom is mostly space, solid objects are
not as “solid” as they appear. Objects that hit each other are not
actually contacting solid surfaces. Instead, negative charges on
the objects repel each other. Without these charges, the objects
would pass through each other.
Atoms are extremely long-lived. It is estimated that they
survive for about 1035 years. Accordingly, the atoms in every
living thing were once found in stars. Each tree you see out-
side your window probably contains a billion atoms, many of
which may well have been in the bodies of your ancestors.
Each atom of an element has a specific number of pro-
tons in its nucleus, ranging from one in hydrogen, the light-
est element, to 92 in uranium, the heaviest natural element.
This number is referred to as the atomic number. The atomic
number is often shown as a subscript to the left of the chemi-
cal symbol. For example, nitrogen, which has seven pro-
tons in its nucleus, has its atomic number of seven shown
as 7N. The combined number of protons and neutrons in
a single atom is referred to as its atomic mass (Table 2.1).
The atomic mass number is commonly shown as a super-
script to the left of the chemical symbol. For example, the
atomic mass of nitrogen, which has seven protons and seven
neutrons in its nucleus, is shown as 14N, and when both the
atomic number and the atomic mass are shown, the chemical
symbol appears as 147 N.
Electrons, which are little more than negative electric
charges, whirl around an atom’s nucleus. Electron masses
are about 1,840 times lighter than those of both protons
and neutrons and are so minute that they are generally dis-
regarded. Since opposite electric charges attract each other,
the positive electric charges of protons attract the negative
electric charges of electrons and determine the paths of the
electrons whirling around the nucleus.
– neutron
– + proton
–
–
electron
–
+ + +
+ ++ +
– +
– –
–
Figure 2.1 Model of an oxygen atom. The nucleus in the center
consists of eight electrically neutral neutrons and eight positively
charged protons. Eight negatively charged electrons whirl around the
nucleus. In a real atom the electrons would not be spaced or confined
as shown in this simple diagram. The nucleus is one-millionth of one-
billionth the diameter of the atom.
The region occupied by electrons around the nucleus is
called an orbital. Each orbital has an imaginary axis and is
somewhat cloudlike, but it doesn’t have a precise boundary,
and so we can’t be certain of an electron’s position within an
orbital at any time. This has led to an orbital being defined
as a volume of space in which a given electron occurs 90%
of the time. Electrons actually occupy all space in an orbital
simultaneously, so they do not circle around the nucleus like
planets. In addition, according to the quantum leap theory
of physics, an electron can move instantaneously from one
orbital to another without visiting the space between them!
Electrons may be located in one or more energy levels of
an atom, and their distance from the nucleus depends on their
energy level. Each energy level is usually referred to as an elec-
tron shell. The outermost electron shell determines how or if an
atom reacts with another atom. Only two electrons can occupy
the first and lowest energy level associated with the innermost
orbital; this orbital is more or less spherical and is so close to
the nucleus that it is often not shown on diagrams of atoms.
One to several additional orbitals, which are mostly spindle
shaped (like the tips of cotton swabs), generally occupy much
more space. Up to eight electrons can be held by the second
energy level, and although the third and fourth energy lev-
els can hold more than eight electrons each, they can become
unstable if more than eight electrons are present. If an electron
in one orbital is provided with more energy, it can jump to an
orbital farther away from the nucleus. Conversely, if an electron
releases energy, it drops to an energy level closer to the nucleus.
The electrons of each orbital tend to repel those of other orbit-
als, so that the axes of all the orbitals of an atom are oriented as
far apart from each other as possible; the outer parts of the orbit-
als, however, actually overlap more than shown in diagrams of
them. Orbitals usually have diameters thousands of times more
extensive than that of an atomic nucleus (Fig. 2.2).
Because each atom usually has as many electrons as it
does protons, the negative electric charges of the electrons
 The Nature of Life 17

Table 2.1 Atomic Numbers, Masses, and Functions of Some Elements Found in Plants

balance the positive charges of the protons,

making the atom electrically neutral. The num-
ber of neutrons in the atoms of an element
can vary slightly, so the element may occur
in forms having different weights but with all
forms behaving alike chemically. Such varia-
tions of an element are called isotopes. The
element oxygen has seven known isotopes. For
instance, the nucleus of one of these isotopes
contains eight protons and eight neutrons; the
(a) (b) nucleus of another isotope holds eight pro-
tons and 10 neutrons (Fig. 2.3). If the number
Figure 2.2 Models of orbitals. (a) The two electrons closest to the atom’s nucleus of neutrons in an isotope of a particular ele-
occupy a single spherical orbit. (b) Additional orbitals are dumbbell-shaped, with axes that
ment varies too greatly from the average num-
are perpendicular to one another. The atom’s nucleus is at the intersection of the axes.
ber of neutrons for its atoms, the isotope may
be unstable and split into smaller parts, with the release
of a great deal of energy. Such an isotope is said to be
radioactive.

Molecules: Combinations of Elements

8p 8p
8n 10n The atoms of most elements can combine with other
atoms of the same or different elements; in fact, most ele-
ments do not exist independently as single atoms. When
two or more elements are united in a definite ratio by
chemical bonds, the substance is called a compound.
Figure 2.3 Isotopes of oxygen portrayed two-dimensionally. Table salt (sodium chloride, NaCl), for example, is a com-
As mentioned in Figure 2.1, the nucleus is proportionally much smaller pound consisting of sodium and chlorine atoms combined
in an atom. in a 1:1 ratio.
18 Chapter 2
A molecule consists of two or more atoms bound
together and is the smallest independently existing particle
of a compound or an element. The molecules of the gases
oxygen and hydrogen, for example, exist in nature as com-
binations of two atoms of oxygen (O2) or two atoms of
hydrogen (H2), respectively. Water molecules (H2O) con-
sist of two atoms of hydrogen and one atom of oxygen
(Fig. 2.4). Molecules are in constant motion, with an
increase or decrease in temperature speeding up or slow-
ing down the motion. The more molecular movement there
is, the greater the chances are that some molecules will
collide with each other. Also, the chances of random colli-
sions increase in proportion to the density of the molecules
(i.e., the number of molecules present in a given space).
Random collisions between molecules capable of sharing
electrons are the basis for all chemical reactions. The reac-
tions often result in new molecules being formed. Each chem-
ical reaction in a cell usually takes place in a watery fluid and
is controlled by a specific enzyme. Enzymes are organic cata-
lysts (a catalyst speeds up a chemical reaction without being
used up in the reaction; enzymes are discussed on page 25).
When a water molecule is formed, two hydrogen atoms
become attached to an oxygen atom at an angle averaging
105° in liquid water (for ice, the angle is precisely 105°).
The electrons of the three atoms are shared and form an
electron cloud around the core, giving the molecule an
asymmetrical shape. Although the electron and proton
charges balance each other, the asymmetrical shape and
unequal sharing of the electrons in the bond between oxygen
and hydrogen cause one side of the water molecule to have a
slight positive charge and the other a slight negative charge.
Such molecules are said to be polar. Since negative charges
attract positive charges, polarity affects the way in which
molecules become aligned toward each other; polarity also
causes molecules other than water to be water soluble.
oxygen (O2)
water (H2O)
hydrogen (H2)
Figure 2.4 Models of oxygen, water, and hydrogen molecules.
A water molecule is 0.6 nanometer in diameter. Each sphere represents
the electron cloud of the outer orbital.
+ + + +
+ + – + + – + +
– + + – + + –
– + + –
–
Figure 2.5 The asymmetrical shape of water molecules and
the resulting unequal sharing of electrons in the bond between the
oxygen and hydrogen atoms cause one side of a water molecule to
have a slight positive charge and the other side a slight negative
charge. Such molecules are said to be polar. The polarity of water
molecules causes them to be attracted to one another in a cohesive
network. The cohesion of water molecules is partly responsible for their
capacity to be pulled in a continuous column through fine (capillary)
tubes, such as those of living wood.
Water molecules form a cohesive network as their
slightly positive hydrogen atoms are attracted to the slightly
negative oxygen atoms of other water molecules (Fig. 2.5).
The cohesion between water molecules is partly responsi-
ble for their movement through fine (capillary) tubes, such
as those present in the wood and other parts of plants. The
attraction between the hydrogen atoms of water and other,
negatively charged molecules, such as those of fibers, also
causes adhesion (attraction of charged molecules to each
other) and is the basis for water wetting substances. When
there is no attraction between water and other substances
(e.g., between water and the waxy surface of a cabbage
leaf ), the cohesion between the water molecules results in
droplets beading in the same way that raindrops bead on a
freshly waxed automobile.
Valence
The combining capacity of an atom or ion based on electron
number is called valence. For example, atoms of the element
calcium, an important element in cell walls and in transmit-
ting chemical “messages” in plant cells, have a valence of
two, while those of the element chlorine have a valence of
one. In order for the atoms of these two elements to combine,
there must be a balance between electrons lost or gained (i.e.,
the valences must balance); it takes two chlorine atoms, for
example, to combine with one calcium atom. The compound
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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