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Stern’s Introductory

Plant Fourteenth Edition

Biology

James E. Bidlack | Shelley H. Jansky

University of Central Oklahoma University of Wisconsin – Madison
STERN’S INTRODUCTORY PLANT BIOLOGY, FOURTEENTH EDITION

Published by McGraw-Hill, a business unit of The McGraw-Hill Companies, Inc., 1221 Avenue of the Americas,
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Names: Bidlack, James E., author. | Jansky, Shelley.

Title: Introductory plant biology / James E. Bidlack, University of Central
Oklahoma, Shelley H. Jansky, University of Wisconsin - Madison.
Other titles: Stern’s introductory plant biology
Description: 14th edition. | New York, NY : McGraw-Hill, 2017. | “Stern’s
introductory plant biology, 14th edition”—T.p. verso.
Identifiers: LCCN 2016044453 | ISBN 9781259682742 (alk. paper)
Subjects: LCSH: Botany.
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www.mhhe.com
 Contents in Brief

About the Authors ix

Preface  x

1 What Is Plant Biology? 1

2 The Nature of Life 13
3 Cells 29
4 Tissues 53
5 Roots and Soils 65
6 Stems 85
7 Leaves 104
8 Flowers, Fruits, and Seeds 124
9 Water in Plants 147
10 Plant Metabolism 164
11 Growth and Development 191
12 Meiosis and Alternation of Generations 216
13 Genetics and Molecular Biology 226
14 Plant Breeding, Propagation, and Biotechnology 249
15 Evolution 268
16 Plant Names and Classification 282
17 Domain (Kingdom) Bacteria, Domain (Kingdom) Archaea, and Viruses 298
18 Kingdom Protista 324
19 Kingdom Fungi 353
20 Introduction to the Plant Kingdom: Bryophytes 378
21 The Seedless Vascular Plants: Ferns and Their Relatives 393
22 Introduction to Seed Plants: Gymnosperms 418
23 Seed Plants: Angiosperms 437
24 Flowering Plants and Civilization 457
25 Ecology 483
26 Biomes 508

Appendix 1 Scientific Names of Organisms Mentioned in the Text A1

Appendix 2 Biological Controls A20
Appendix 3 Useful and Poisonous Plants, Fungi, and Algae A26
Appendix 4 House Plants and Home Gardening A51
Appendix 5 Metric Equivalents and Conversion Tables A75
Appendix 6 Periodic Table of the Elements A77

Glossary  G1
Index  I1
iii
Contents

About the Authors ix

Preface  x
4 Tissues 53
Overview 54
1 What Is Plant Biology? 1 Learning Outcomes 54
Meristematic Tissues 54
Overview 2 KEY THEME: molecular Chimeras and Variegated
Learning Outcomes 2 Leaves 55
KEY THEME: ecology Who Needs Plants? 4 Tissues Produced by Meristems 56
The Relationship of Humans to Their KEY THEME: ecology Plants and Environment 61
Environment 4 SUMMARY 63
REVIEW QUESTIONS 64
Botany as a Science 7
DISCUSSION QUESTIONS 64
Diversification of Plant Study 7 ADDITIONAL READING 64
Plant Biology on the Internet 10 LEARNING ONLINE 64
SUMMARY 11
REVIEW QUESTIONS 11
DISCUSSION QUESTIONS  11
ADDITIONAL READING 11
5 Roots and Soils 65
LEARNING ONLINE 12 Overview 66
Learning Outcomes 66
2 The Nature of Life 13 How Roots Develop 66
Root Structure 67
Overview 14 Specialized Roots 71
Learning Outcomes 14 KEY THEME: ecology Plants Need Roots 75
Attributes of Living Organisms 14 Mycorrhizae 76
Chemical and Physical Bases of Life 15 Root Nodules 77
KEY THEME: molecular The Skinny on Fats 23 Human Relevance of Roots 77
SUMMARY 27
Soils 78
REVIEW QUESTIONS 27
KEY THEME: ecology Metal-Munching
DISCUSSION QUESTIONS 28
ADDITIONAL READING 28 Plants 81
LEARNING ONLINE 28 SUMMARY 82
REVIEW QUESTIONS 83

3 Cells 29
DISCUSSION QUESTIONS  83
ADDITIONAL READING 84
LEARNING ONLINE 84
Overview 30
Learning Outcomes 30
Cells 30 6 Stems 85
Eukaryotic versus Prokaryotic Cells 33 Overview 86
Cell Structure and Communication 33 Learning Outcomes 86
Cellular Components 36 External Form of a Woody Twig 86
Cellular Reproduction 44 Origin and Development of Stems 87
Microscapes 48 KEY THEME: ecology Standing in Fields of
Higher Plant Cells versus Animal Cells 50 Stone 88
SUMMARY 51
Tissue Patterns in Stems 90
REVIEW QUESTIONS 52
DISCUSSION QUESTIONS  52
KEY THEME: ecology Dendroclimatology 92
ADDITIONAL READING 52 Specialized Stems 97
LEARNING ONLINE 52 Wood and Its Uses 99
iv
 Contents v

SUMMARY 102 Transport of Food Substances (Organic

REVIEW QUESTIONS 103
Solutes) in Solution 157
DISCUSSION QUESTIONS 103
ADDITIONAL READING 103
Mineral Requirements for Growth 159
SUMMARY 162
LEARNING ONLINE 103
REVIEW QUESTIONS 162
DISCUSSION QUESTIONS 162

7 Leaves 104
ADDITIONAL READING 163
LEARNING ONLINE 163

Overview 105
Learning Outcomes 105
Leaf Arrangements and Types 106 10 Plant Metabolism 164
Internal Structure of Leaves 107 Overview 165
Stomata 108 Learning Outcomes 165
Mesophyll and Veins 110 Enzymes and Energy Transfer 166
Specialized Leaves 111 Photosynthesis 166
KEY THEME: ecology More on Leaf Structure 112 Photosynthesis and Pizza 176
Autumnal Changes in Leaf Color 119 Respiration 180
Abscission 120 Additional Metabolic Pathways 185
Human and Ecological Relevance of Assimilation and Digestion 186
Leaves 121 SUMMARY 187

Glass Cuts from Grass? 122 KEY THEME: ecology Photosynthesis, Global
SUMMARY 122 Warming, and Tropical Rain Forests 188
REVIEW QUESTIONS 123 REVIEW QUESTIONS 190
DISCUSSION QUESTIONS 123 DISCUSSION QUESTIONS 190
ADDITIONAL READING 123 ADDITIONAL READING 190
LEARNING ONLINE 123 LEARNING ONLINE 190

8 Flowers, Fruits, and Seeds 124 11 Growth and Development 191

Overview 125 Overview 192
Learning Outcomes 125 Learning Outcomes 192
Differences between Dicots and Nutrients, Vitamins, and Hormones 192
Monocots 128 Plant Hormones beyond “The Classic
Structure of Flowers 128 Five” 200
Fruits 129 Hormonal Interactions 201
KEY THEME: ecology Goober Peas 131 Other Hormonal Interactions 201
Fruit and Seed Dispersal 137 Plant Movements 202
Seeds 141 Photoperiodism 209
The Seed That Slept for 1,200 Years 144 Phytochromes and Cryptochromes 210
SUMMARY 145 A Flowering Hormone? 211
REVIEW QUESTIONS 146 Temperature and Growth 212
DISCUSSION QUESTIONS 146
Dormancy and Quiescence 213
ADDITIONAL READING 146
SUMMARY 214
LEARNING ONLINE 146
REVIEW QUESTIONS 215
DISCUSSION QUESTIONS 215

9 Water in Plants 147

ADDITIONAL READING 215
LEARNING ONLINE 215

Overview 148
Learning Outcomes 148
Molecular Movement 149
12 Meiosis and Alternation
Measuring Water Potential and
of Generations 216
Psychrometry 151 Overview 217
Water and Its Movement through the Plant 153 Learning Outcomes 217
Regulation of Transpiration 156 The Phases of Meiosis 218
vi Contents

KEY THEME: evolution Why Plants Have Sex Charles Darwin 273
Lives 219 Evidence for Evolution 274
Alternation of Generations 221 Microevolution—Evolution within
KEY THEME: molecular FISH and GISH Molecular Species 275
Techniques 223 Rates of Evolution 276
SUMMARY 223 Macroevolution—How Species Evolve 276
REVIEW QUESTIONS 224
The Role of Polyploidy in Evolution 278
DISCUSSION QUESTIONS  224
ADDITIONAL READING 224
Discussion 279
LEARNING ONLINE 224 KEY THEME: evolution Our Daily Bread 280
SUMMARY 280

13 Genetics and Molecular

REVIEW QUESTIONS 281
DISCUSSION QUESTIONS  281

Biology 226 ADDITIONAL READING 281

LEARNING ONLINE 281
Overview 227
Learning Outcomes 227
Molecular Genetics 228
16 Plant Names and
KEY THEME: molecular Massive DNA
Classification 282
Sequencing 230 Overview 283
KEY THEME: molecular The Polymerase Chain Learning Outcomes 283
Reaction (PCR) 232 Development of the Binomial System of
Cytogenetics 237 Nomenclature 283
Mendelian Genetics 238 Development of the Kingdom Concept 286
Quantitative Traits 244 Classification of Major Groups 287
Extranuclear DNA 245 Species Concepts 292
Linkage and Mapping 245 A Key to Major Groups of Organisms
The Hardy-Weinberg Law 247 (Exclusive of Kingdom Animalia) 294
SUMMARY 247 The Future of Plant Classification 296
REVIEW QUESTIONS 248 SUMMARY 296
DISCUSSION QUESTIONS 248 REVIEW QUESTIONS 297
ADDITIONAL READING 248 DISCUSSION QUESTIONS  297
LEARNING ONLINE 248 ADDITIONAL READING 297
LEARNING ONLINE 297

14 Plant Breeding, Propagation,

17 Domain (Kingdom) Bacteria,
and Biotechnology 249
Domain (Kingdom) Archaea,
Overview 250
Learning Outcomes 250
and Viruses 298
Crop Plant Evolution 250 Overview 299
Plant Breeding 252 Learning Outcomes 299
KEY THEME: molecular Genome Editing 259 Features of Domains (Kingdoms) Bacteria
Plant Propagation 260 and Archaea 300
SUMMARY 266 Domain (Kingdom) Bacteria—the True
REVIEW QUESTIONS 267
Bacteria 304
DISCUSSION QUESTIONS  267
ADDITIONAL READING 267
Human Relevance of the Unpigmented,
LEARNING ONLINE 267 Purple, and Green Sulfur Bacteria 304
KEY THEME: ecology The Social Life of

15 Evolution 268
Prokaryotes 305
Class Cyanobacteriae—the Cyanobacteria
Overview 269 (Blue-Green Bacteria) 311
Learning Outcomes 269 KEY THEME: ecology Cyanobacteria and Algae
An Introduction to Evolution 269 Blooms 314
A Brief Overview of the Early Development Class Prochlorobacteriae—the
of Evolutionary Concepts 271 Prochlorobacteria 315
 Contents vii

Domain (Kingdom) Archaea—the REVIEW QUESTIONS 376

DISCUSSION QUESTIONS  377
Archaebacteria 315
ADDITIONAL READING 377
Viruses 317 LEARNING ONLINE 377
KEY THEME : molecular Plant Viruses 318
Viroids and Prions 321
SUMMARY 322 20 Introduction to the Plant
REVIEW QUESTIONS 323 Kingdom: Bryophytes 378
DISCUSSION QUESTIONS  323
ADDITIONAL READING 323 Overview 379
LEARNING ONLINE 323 Learning Outcomes 379
Introduction to the Bryophytes 380

18 Kingdom Protista 324

Phylum Hepaticophyta—Liverworts 381
Phylum Anthocerophyta—Hornworts 385
Overview 325 Phylum Bryophyta—Mosses 385
Learning Outcomes 325 KEY THEME: ecology Hibernating Mosses 389
KEY THEME: evolution Green Plant Human and Ecological Relevance of
Phylogeny 326 Bryophytes 390
SUMMARY 391
Features of Kingdom Protista 326
REVIEW QUESTIONS 391
Algae 326
DISCUSSION QUESTIONS 392
Phylum Chlorophyta—the Green Algae 327 ADDITIONAL READING 392
Phylum Chromophyta—the Yellow-Green LEARNING ONLINE 392
Algae, Golden-Brown Algae, Diatoms,
and Brown Algae 333
Phylum Rhodophyta—the Red Algae 337
21 The Seedless Vascular Plants:
Phylum Euglenophyta—the Euglenoids 339
Ferns and Their Relatives 393
Phylum Dinophyta—the Dinoflagellates 340 Overview 394
Phylum Cryptophyta—the Cryptomonads 341 Learning Outcomes 394
Phylum Prymnesiophyta (Haptophyta)—the Phylum Psilotophyta—the Whisk Ferns 394
Haptophytes 342 Phylum Lycophyta—the Ground Pines,
Phylum Charophyta—the Stoneworts 342 Spike Mosses, and Quillworts 396
KEY THEME: ecology Biofuels from Algae 343 Phylum Equisetophyta—the Horsetails and
Human and Ecological Relevance of the Scouring Rushes 402
Algae 343 Phylum Polypodiophyta—the Ferns 406
Other Members of Kingdom Protista 347 Fossils 413
Phylum Myxomycota—the Plasmodial KEY THEME: ecology Ferns and Fossil Fuels 414
Slime Molds 347 SUMMARY 415
REVIEW QUESTIONS 416
Phylum Dictyosteliomycota—the Cellular
DISCUSSION QUESTIONS 416
Slime Molds 348
ADDITIONAL READING 416
Phylum Oomycota—the Water Molds 350 LEARNING ONLINE 417
SUMMARY 351

22 Introduction to Seed Plants:

REVIEW QUESTIONS 352
DISCUSSION QUESTIONS 352
ADDITIONAL READING 352
LEARNING ONLINE 352 Gymnosperms 418
Overview 419
19 Kingdom Fungi 353 Learning Outcomes 419
Phylum Pinophyta—the Conifers 420
Overview 354 KEY THEME: ecology Resilient and Useful
Learning Outcomes 354 Gymnosperms 421
Distinctions between Kingdoms Protista Other Gymnosperms 424
and Fungi 354 Human Relevance of Gymnosperms 429
Kingdom Fungi 355 KEY THEME: evolution A Living Fossil? 434
Lichens 373 SUMMARY 435
SUMMARY 375 REVIEW QUESTIONS 436
viii Contents

DISCUSSION QUESTIONS 436

KEY THEME: ecology John Muir, Father of
ADDITIONAL READING 436
America’s National Park System 504
LEARNING ONLINE 436
SUMMARY 505
REVIEW QUESTIONS 506

23 Seed Plants: Angiosperms 437 DISCUSSION QUESTIONS 506

ADDITIONAL READING 506
LEARNING ONLINE 507
Overview 438
Learning Outcomes 438
Phylum Magnoliophyta—the Flowering
Plants 439 26 Biomes 508
KEY THEME: molecular The Difference between Overview 509
“n” and “x” in Plant Life Cycles 446 Learning Outcome 509
Pollination Ecology 448 Major Biomes of the World 509
Herbaria and Plant Preservation 452 KEY THEME: ecology Alpine Flora as an Indication
SUMMARY 455
of Climate Change: The GLORIA Project 514
REVIEW QUESTIONS 455
SUMMARY 518
DISCUSSION QUESTIONS 456
REVIEW QUESTIONS 519
ADDITIONAL READING 456
DISCUSSION QUESTIONS 519
LEARNING ONLINE 456
ADDITIONAL READING 519
LEARNING ONLINE 519

24 Flowering Plants Appendix 1 Scientific Names of Organisms

and Civilization 457 Mentioned in the Text A1
Overview 458 Appendix 2 Biological Controls A20
Learning Outcomes 458 General Controls A20
Origin of Cultivated Plants 458 Specific Controls A22
Selected Families of Flowering Companion Planting A22
Plants 459 Additional Reading A22
Dicots (Now Recognized in Two Appendix 3 Useful and Poisonous Plants, Fungi,
Groups) 461 and Algae A26
Monocots 476 Wild Edible Plants, Fungi, and Algae A26
KEY THEME: ecology Wild Rice—More Than Just Poisonous Plants and Fungi A26
Food 478 Medicinal Plants, Fungi, and Algae A26
KEY THEME: ecology Coffee and Caffeine 480 Hallucinogenic Plants A44
SUMMARY 480
Spice Plants A44
REVIEW QUESTIONS 481
DISCUSSION QUESTIONS 482
Dye Plants A44
ADDITIONAL READING 482 Additional Reading A50
LEARNING ONLINE 482 Appendix 4 House Plants and Home
Gardening A51

25 Ecology 483 Growing House Plants A51

Common House Plants A52
Overview 484 Growing Vegetables A61
Learning Outcomes 484 Common Vegetables and their Nutritional
Plants and the Environment 484 Values A62
Life Histories 489 Pruning A67
Natural Cycles 490 Major Types of Grafting A68
Succession 493 Additional Reading A74
KEY THEME: ecology Plant Population Appendix 5 Metric Equivalents and Conversion
Ecology 493 Tables A75
Impact of Humans on Plant Appendix 6 Periodic Table of the Elements A77
Communities 498
Loss of Biodiversity 501 Glossary G1
Restoration of the Land 503 Index I1
 About the Authors

Introductory Plant Biology was originally written by Kingsley

R. Stern (1927–2006), who spent more than 40 years as a devoted
botanist and teacher. It is estimated that Dr. Stern educated 15,000
students through classroom/laboratory teaching and inspired thou-
sands of botanists as the author of this book, which has sold more
than 275,000 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 was always Kingsley’s aspiration that
those who read Stern’s Introductory Plant Biology will share his life-
long love of botany.

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

office residence in Paradise, California. © Jan Monelo
In late 1999 and early 2000, Drs. Jim Bidlack and Shelley Jansky joined Kingsley Stern in editing and updating this textbook.
They worked with him on several editions and have since carried Dr. Stern’s legacy forward to educate and inspire young bota-
nists. This fourteenth edition reflects the same accuracy, content, and enthusiasm of the Stern writing style, along with revi-
sions 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 bachelor of sci- Shelley Jansky received a bachelor’s
ence degree in agronomy, with a soil and degree in biology from the University of
crop option, from Purdue University in Wisconsin–Stevens Point in 1982, and
1984 and continued his education with a a master of science degree and Ph.D. in
master’s degree in crop physiology at the plant breeding and plant genetics from
University of Arkansas in 1986. Upon the University of Wisconsin–Madison
© James E. Bidlack completing a Ph.D. in plant physiology at © Shelley H. Jansky in 1984 and 1986, respectively. Her
Iowa State University in 1990, Jim joined graduate work focused on developing
the teaching faculty at the University of Central Oklahoma methods to incorporate genes from wild relatives of potato
(UCO) where he is a professor of biology. His first paper was into the cultivated potato. Then, she spent four years as an
published from undergraduate research at Purdue University assistant professor at North Dakota State University, teach-
on the use of synthetic growth regulators to stimulate seed ger- ing courses in plant breeding and plant propagation, and
mination. Subsequent work in Arkansas, Iowa, and Oklahoma performing research in the potato breeding program. She
focused on soybean physiology, cell wall chemistry, alterna- taught courses in botany, genetics, and horticulture, and con-
tive crops, and photovoltaic cells, as well as teaching respon- tinued to perform potato genetics research at the University
sibilities in botany. Equipment and student salaries for Jim’s of Wisconsin–Stevens Point from 1990 until 2004. She
research projects have been funded by grants from the National was the chair of the Department of Biology and was pro-
Science Foundation and the United States Department moted to associate professor in 1992 and professor in 1995.
of Agriculture. About 20 refereed publications, as well as In 2004, she moved to Madison, where she is a research
hundreds of abstracts and popular articles, have resulted from geneticist with the U.S. Department of Agriculture and an
this work. Jim has been recognized with UCO’s Presidential associate professor in the Department of Horticulture at the
Partner’s Excellence in Teaching Award; University Merit University of Wisconsin–Madison. Her research program
Awards in Service, Research, and Teaching; the Biology Club focuses on using disease resistance and nutritional quality
Teaching Award; and the Pre-Med Teaching Award. Some of genes from wild relatives for the improvement of cultivated
Jim’s additional responsibilities have included participation on potato varieties. She received the University of Wisconsin–
NSF and USDA review panels, editor of the Biology Discipline Stevens Point Excellence in Teaching Award in 1992 and the
for the Multimedia Educational Resource for Learning and University Scholar Award in 2000. She has published 87 ref-
Online Teaching (MERLOT), vice president of The Genome ereed research articles and seven book chapters.
Registry, and president of Metabolism Foundation.
ix
Preface

Plants and algae are essential for life on earth as it exists today. aspects and other information of general interest pertaining to
They provide our world with oxygen and food, contribute an 16 major plant families or groups of families. Chapters 25 and
essential part of water and nutrient cycling in ecosystems, pro- 26 present an overview of the vast topic of ecology, although
vide clothing and shelter, and add beauty to our environment. ecological topics and applied botany are included in the pre-
Some scientists believe that if photosynthetic organisms exist ceding chapters as well. Some of these topics are broached in
on planets beyond our solar system, it would be possible to anecdotes that introduce the chapters, while others are men-
sustain other forms of life that depend upon them to survive. tioned in text boxes as well as the appendices.
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,
Learning Aids
acid rain, genetic engineering, organic gardening, Native A chapter outline, review questions, discussion questions,
American and pioneer uses of plants, pollution and recy- and additional reading lists are provided for each chapter.
cling, house plants, backyard vegetable gardening, natural New terms are defined as they are introduced, and those that
dye plants, poisonous and hallucinogenic plants, nutritional are boldfaced are included, with their pronunciation, in a
values of edible plants, and many other topics are discussed. glossary. A list of the scientific names of all organisms men-
To intelligently pursue such topics, one needs to understand tioned throughout the text is given in Appendix 1. Appendix
how plants are constructed, and how they function. To this 2 deals with biological controls and companion planting.
end, the text assumes little prior knowledge of the sciences Appendix 3 includes wild edible plants, poisonous plants,
on the part of the student, but covers basic botany, without medicinal plants, hallucinogenic plants, spices, tropical
excessively resorting to technical terms. The coverage, how- fruits, and natural dye plants. Appendix 4 gives horticultural
ever, includes sufficient depth to prepare students to go fur- information on house plants, along with brief discussions on
ther in the field, should they choose to do so. how to cultivate vegetables. Nutritional values of the veg-
The text is arranged so that certain sections can be etables are included. Appendix 5 covers metric equivalents
omitted in shorter courses. Such sections may include top- and conversion tables.
ics 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
New to this Edition
majors and nonmajors, these topics are presented at three The fourteenth edition is an exciting one, with dozens of new/
different levels. Some instructors will find one or two levels replacement photographs and updates in the text to make it
sufficient, whereas others will want to include all three. a modern textbook with great new topics. Most of the major
Both majors in botany and nonmajors who may initially changes in the book have been made as a result of new dis-
be disinterested in the subject matter of a required course fre- coveries and technologies in plant biology. This edition still
quently become engrossed if the material is related repeatedly retains the hallmark style and pedagogy that make it one of
to their popular interests. This is reflected, as intimated above, the most enduring and popular introductory plant biology
in the considerable amount of ecology and ethnobotany books on the market, and it now has a more contemporary
included with traditional botany throughout the book. look and presentation style. All chapters incorporate mea-
surable learning outcomes and updated additional readings.

Organization of the Text For instructors using Internet resources such as Connect and
the textbook website offered by McGraw-Hill, all learning
A relatively conventional sequence of botanical subjects is outcomes are directly tied to assessment within the ques-
followed. Chapters 1 and 2 cover introductory and background tion and test banks. With encouragement from reviewers,
information; Chapters 3 through 11 deal with structure and the new material has been eloquently incorporated directly
function; Chapters 12 and 13 introduce meiosis, genetics, and into the textbook narrative and as text boxes, to provide a
molecular biology. Chapter 14 discusses plant propagation blend of historically important principles as well as modern
and biotechnology; Chapter 15 introduces evolution; Chapter developments in plant biology. Some of the more interesting
16 deals with classification; Chapters 17 through 23 stress, in components that make this fourteenth edition more accurate
phylogenetic sequence, the diversity of organisms traditionally and up-to-date with our current understanding of plant biol-
regarded as plants; and Chapter 24 deals with ethnobotanical ogy include:
x
 Preface xi

Chapter 1 (What Is Plant Biology?): The scientific Chapter 17 (Domain (Kingdom) Bacteria, Domain
method section has been extensively revised. The Plant (Kingdom) Archaea, and Viruses): Some of the information
Biology and the Internet text box has been updated with on human diseases (such as AIDS and tuberculosis), as well
information about how the Internet has affected the scien- as human uses of archaebacteria (such as methane produc-
tific community. tion) has been updated. Additionally, a text box on cyano-
Chapter 2 (The Nature of Life): New examples have bacteria and algae blooms has been added to the chapter to
been added and text has been updated and revised for clarity. show the huge impact algae can have on ecosystems.
Chapter 3 (Cells): Several photographs have been Chapter 18 (Kingdom Protista): Two important and
replaced with higher-quality images. innovative text boxes have been added to this chapter—one
Chapter 4 (Tissues): The introduction has been rewrit- on green plant phylogeny and another on biofuels made
ten in language relevant to today’s college students. A flow- from algae. The text box on green plant phylogeny, in partic-
chart has been added to show how meristems give rise to ular, is an exciting new topic presented to introduce readers
tissues. Trichomes have been defined and added to the index. to the new approaches being used in taxonomy using DNA
Chapter 5 (Roots and Soils): The soil profile caption technology. Other modifications have also been made to this
has been enhanced. chapter to focus on the basic differences among life cycles
Chapter 6 (Stems): The caption for the cover image has and how these can be used to describe and distinguish
been expanded. among the algae phyla.
Chapter 7 (Leaves): Some discussion on incorporation Chapter 19 (Kingdom Fungi): More direct language
of plant pigments in photovoltaic cells has been added to the was incorporated into this chapter to make concepts easier
introduction. to understand. A great new picture of a puffball was also
Chapter 8 (Flowers, Fruits, and Seeds): The role of res- added to the chapter.
piration in seed germination is described in more detail. Chapter 20 (Introduction to the Plant Kingdom:
Chapter 9 (Water in Plants): The description of active Bryophytes): Antheridiophores and archegoniophores have
transport has been revised. been more accurately described and pointed out in one of
Chapter 10 (Plant Metabolism): A more straightforward the figures. Additionally, the important feature of hornworts
introduction is provided at the beginning of the chapter. having stomata has been added to the section on Phylum
Chapter 11 (Growth and Development): The distinction Anthocerophyta.
between determinate and indeterminate growth has been Chapter 21 (The Seedless Vascular Plants: Ferns and
clarified. The caption for the photograph on photoperiodism Their Relatives): The concept that seedless vascular plants
has been expanded. are sporophyte dominant, and the spores produced germi-
Chapter 12 (Meiosis and Alternation of Generations): nate into a free-living gametophye generation, is empha-
The introduction has been re-written to describe the conse- sized at the beginning of the chapter. More direct language
quences of meiosis and how it results in variability when farm- was also incorporated into this chapter to make it more
ers save seeds from hybrid crops. The distinction between x straightforward.
(ploidy) and n (gametophyte versus sporophyte generation) Chapter 22 (Introduction to Seed Plants: Gymnosperms):
has been clarified. The idea that the sporophyte becomes The phrase “common ancestor” is introduced in this chapter
more dominant in advanced plant lineages is introduced. to convey the more modern approach to plant classification
Chapter 13 (Genetics and Molecular Biology): The using DNA technology. New information is also presented
description of a DNA molecule has been enhanced. The on the production of paclitaxel (also known by its trade
description of 2n gametes has been expanded. The role of name Taxol) from new lines (derived from tissue culture) of
the monohybrid cross in illustrating the law of segregation yew trees.
is discussed. A section has been added to describe genetic Chapter 23 (Seed Plants: Angiosperms): More direct
engineering technologies to develop herbicide-resistant language was incorporated into this chapter to make it more
plants. straightforward and easier to understand.
Chapter 14 (Plant Breeding, Propagation, and Chapter 26 (Biomes): Some of the climatological data
Biotechnology): A new text box on “genome editing,” as a and specific information about biomes have been updated.
clever method for recombinant DNA technology with end- Appendix 2 (Biological Controls): The introduction has
less possibilities, has been added to the chapter. Additional been rewritten to discuss the consequences of improper use
updates on transgenic plants and rewording of the text has of pesticides. The section on the use of resistant plant variet-
also been completed to make the content more current. ies has been updated. The section on the control of weeds
Chapter 15 (Evolution): Some strategic rewording of has also been updated.
the text was done to provide a more straightforward presen- Appendix 4 (House Plants and Home Gardening): This
tation of evolutionary concepts. appendix has been extensively updated. New information is
Chapter 16 (Plant Names and Classification): More pre- presented on LED lights, transplanting of seedlings, direct
cise information is presented to show the global estimate of seeding, temperature and seed germination, the effect of
existing species to be about 8.7 million, and some other tact- overwatering on root growth, fertilizer application, pest con-
ful changes were made in the text to bring it up to date. trol, and inoculation of soil with Rhizobia.
 Required=Results
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xiv Preface

Acknowledgements and suggestions. Although too numerous to include here,

their contributions have been much appreciated. The follow-
Over 275 reviewers for the past few editions, along with ing is a list of reviewers for recent editions:
reviewers for this fourteenth edition, have helped to revise
and update Stern’s Introductory Plant Biology. In particu- Ligia Arango, Stone Child College
lar, David Spooner, USDA-ARS Botanist and Professor at Joseph Arditti, University of California–Irvine
the University of Wisconsin, provided excellent input for Mark H. Armitage, Azusa Pacific University
Chapter 16 on Plant Names and Classification. Additional Janice Asel, Mitchell Community College
contributions and encouragement were provided by Jan Tasneem K. Ashraf, Cochise College–Sierra Vista
Monelo; colleagues at the University of Central Oklahoma Ralph A. Backhaus, Arizona State University
and University of Wisconsin; and the design, editorial, and Nina L. Baghai-Riding, Delta State University
production staffs of McGraw-Hill Education. The authors Randy G. Balice, New Mexico Highlands University
extend thanks to the following reviewers who provided Susan C. Barber, Oklahoma City University
recent feedback on the text and the illustrations. Their help Paul W. Barnes, Southwest Texas State University
has been very useful in shaping the fourteenth edition of Sharon Bartholomew-Began, West Chester University
Stern’s Introductory Plant Biology. These reviewers include Robert W. Bauman, Jr., Amarillo College
the following: Dorothea Bedigian, Washington University
Patricia Bedinger, Colorado State University
Donovan Bailey, New Mexico State University
Maria Begonia, Jackson State University
Terese Barta, University of Wisconsin-Stevens Point
Robert A. Bell, University of Wisconsin–Stevens Point
Lisa Boucher, University of Nebraska-Omaha
Cynthia A. Bottrell, Scott Community College
Charles Cannon, Texas Tech University
Richard R. Bounds, Mount Olive College
William Cook, Midwestern State University
Richard G. Bowmer, Idaho State University
Roger del Moral, University of Washington
Rebecca D. Bray, Old Dominion University
Melinda Findlater, Texas Tech University
James A. Brenneman, University of Evansville
Dr. Gabrielle Forbes, Southwest Texas Junior College
George M. Briggs, State University of New York
Stephen W. Fuller, University of Mary Washington
Michelle Briggs, Lycoming College
Ed Gasque, University of Wisconsin-Stevens Point
George M. Brooks, Ohio University
Rick Hammer, Hardin-Simmons University
Suzanne Butler, Miami-Dade College
Paul Hankamp, College of San Mateo
William J. Campbell, Louisiana Technical University
Allan Scott Holaday, Texas Tech University
Ajoy G. Chakrabarti, South Carolina State University
Harry Janes, Rutgers University
Brad S. Chandler, Palo Alto College
Terry Jordan, College of Southern Maryland
Gregory Chandler, University of North Carolina–
Ari Jumpponen, Kansas State University
Wilmington
John Kiss, Miami University
James A. Christian, Louisiana Technical University
Jeffrey K. Lake, Adrian College
Richard Churchill, Southern Maine Technical College
Fengshan Ma, Wilfrid Laurier University
Jerry A. Clonts, Anderson College
Silvia Maciá, Barry University
John Cruzan, Geneva College
Rizana M. Mahroof, South Carolina State University
Kenneth J. Curry, University of Southern Mississippi
Karen McGinnis, Florida State University
David B. Czarnecki, Loras College
James Mickle, North Carolina State University
Stephen S. Daggett, Avila College
Carrie Monlux, California State University-Chico
Raviprakash G. Dani, Texas Tech University
Clark L. Ovrebo, University of Central Oklahoma
Roy Darville, East Texas Baptist University
Bob Peregoy, Spokane Community College
Cynthia Dassler, Ohio State University
Dennis T. Ray, University of Arizona
Bill D. Davis, Rutgers University
Flona Redway, Barry University
Jerry D. Davis, University of Wisconsin–LaCrosse
Traesha R. Robertson, Texas Tech University
John W. Davis, Benedictine College
Connie E. Rye, East Mississippi Community College
Semma Dhir, Fort Valley State University
Devinder Sandhu, University of Wisconsin-Stevens
Rebecca M. DiLiddo, Mount Ida College
Point
Susan C. Dixon, Walla Walla College
Mikel Stevens, Brigham Young University
Ben L. Dolbeare, Lincoln Land Community College
Qiang Sun, University of Wisconsin-Stevens Point
Patricia M. Dooris, Saint Leo College
Leslie Towill, Arizona State University
Donald Drake, University of Hawaii
Carol Wake, South Dakota State University
Tom Dudley, Angelina College
Upon reaching this milestone fourteenth edition, we would Jan Federic Dudt, Bartlesville Wesleyan College
also like to once again extend gratitude to the reviewers of Diane Dudzinski, Washington State Community College
earlier editions, who have provided considerable comments Kerry B. Dunbar, Dalton State College
 Preface xv

Robert Ediger, California State University–Chico Vic Landrum, Washburn University

H. Herbert Edwards, Western Illinois University James M. Lang, Greenville College
William Eisinger, Santa Clara University Brenda Price Latham, Merced College
Inge Eley, Hudson Valley Community College Cheryl R. Laursen, Eastern Illinois University
Thomas E. Elthon, University of Nebraska–Lincoln Peter J. Lemay, College of the Holy Cross
Frederick B. Essig, University of South Florida Donald C. Leynaud, Wabash Valley College
G. F. Estabrook, The University of Michigan Barbara E. Liedl, Central College
James Ethridge, Joliet Junior College John F. Logue, University of South Carolina–Sumter
Paul G. Fader, Freed-Hardeman University Elizabeth L. Lucyszyn, Medaille College
Bruce Felgenhauer, University of Louisiana–Lafayette Karen Lustig, Harper College
Jorge F. S. Ferreira, Southern Illinois University– Erin D. MacKenzie, Weatherford College
Carbondale Paul Mangum, Midland College
David G. Fisher, Maharishi University of Management Steve Manning, Arkansas State University–Beebe
Rosemary H. Ford, Washington College Michael H. Marcovitz, Midland Lutheran College
James Garner, Horry-Georgetown Technical College Bernard A. Marcus, Genesee Community College
Sibdas Ghosh, University of Wisconsin–Whitewater David Martin, Centralia College
Mike Gipson, Oklahoma Christian University Margaret Massey, Mississippi University for Women
Katherine Glew, University of Washington William J. Mathena, Kaskaskia College
Richard Glick, Winston-Salem State University Alicia Mazari-Andersen, Kwantlen University College
Charles Good, Ohio State University Andrew McCubbin, Washington State University
David L. Gorchov, Miami University of Ohio Joseph H. McCulloch, Normandale Community College
Scott A. Gordon, University of Southern Illinois Julie A. Medlin, Northwestern Michigan College
Govindjee, University of Illinois Richard G. Merritt, Houston Community College
Steve Greenwald, Gordon College Andrew S. Methven, Eastern Illinois University
Timothy C. Hall, Texas A & M University Timothy Metz, Campbell University
Mark Hammer, Wayne State College David H. Miller, Oberlin College
Susan Han, University of Massachusetts David W. Miller, Clark State Community College
Laszlo Hanzely, Northern Illinois University Subhash C. Minocha, University of New Hampshire
Joyce Phillips Hardy, Chadron State College Beth Morgan, University of Illinois, Urbana-Champaign
Nancy E. Harris, Elon College Dale M. J. Mueller, Texas A & M University
David Hartsell, Phillips Community College Lytton John Musselman, Old Dominion University
Jill F. Haukos, South Plains College Nusrat H. Naqvi, Southern University
David L. Herrin, University of Texas–Austin Joanna H. Norris, University of Rhode Island
Peter Heywood, Brown University Chuks A. Ogbonnaya, Mountain Empire College
Jeffrey P. Hill, Idaho State University Jeanette C. Oliver, Flathead Valley Community College
L. Michael Hill, Bridgewater College Sebastine O. Onwuka, Lesley College
H. H. Ho, State University of New York–New Paltz A. D. Owings, Southeastern Louisiana University
Elisabeth A. Hooper, Truman State University Julie M. Palmer, University of Texas–Austin
Susan Houseman, Southeastern Community College Richard A. Palmer, Fresno City College
Lauren D. Howard, Norwich University Carolyn Peters, Spoon River College
Vernon R. Huebschwerlen, Reedley Community College Martha M. Phillips, The College of St. Catherine
Patricia L. Ireland, San Jacinto College, South Jerry L. Pickering, Indiana University of Pennsylvania
William A. Jensen, Ohio State University Wayne S. Pierce, California State University–Stanislaus
Cindy Johnson-Groh, Gustavus Adolphus College Indiren Pillay, Southwestern Tennessee Community
Chad Jordan, North Carolina State University College
Toney Keeney, Southwest Texas State Mary Ann Polasek, Cardinal Stritch University
Sekender A. Khan, Elizabeth City State University Kumkum Prabhakar, Nassau Community College
Joanne M. Kilpatrick, Auburn University–Montgomery Tyre J. Proffer, Kent State University
Helen G. Kiss, Miami University Francis Putz, University of Florida
John Z. Kiss, Miami University of Ohio V. Raghaven, The Ohio State University
Kaoru Kitajima, University of Florida Mohammad A. Rana, St. Joseph College
Sharon Klavins, University of Wisconsin–Platteville Margene M. Ranieri, Bob Jones University
Roger C. Klockziem, Martin Luther College W. T. Rankin, University of Montevallo
Robert L. Koenig, Southwest Texas Junior College Linda Mary Reeves, San Juan College
David W. Kramer, Ohio State University–Mansfield Maralyn A. Renner, College of the Redwoods
Robert N. Kruger, Mayville State University Tom Reynolds, University of North Carolina–Charlotte
Martin LaBar, Southern Wesleyan University Stanley A. Rice, Southeastern Oklahoma State University
xvi Preface

Dennis F. Ringling, Pennsylvania College of Technology Rani Vajravelu, University of Central Florida
Daryl Ritter, Okaloosa-Walton Community College John Vanderploeg, Ferris State University
Suzanne M. D. Rogers, Salem International University Staria Vanderpool, Arkansas State University
Wayne C. Rosing, Middle Tennessee State University Delmar Vander Zee, Dordt College
Robert G. Ross, University of Puerto Rico C. Gerald Van Dyke, North Carolina State University
Jimmy Rozell, Tyler Junior College Leon Walker, University of Findlay
Manfred Ruddat, University of Chicago Ami Lea Wangeline, Colorado State University
Patricia Rugaber, Coastal Georgia Community Betty J. Washington, Albany State University
College Edgar E. Webber, Keuka College
Frances Rundlett, Georgia State University Christopher R. Wenzel, Eastern Wyoming College
Thomas H. Russ, Charles County Community College Cherie Wetzel, City College of San Francisco
Dennis J. Russell, University of Alaska Southeast Ingelia White, Windward Community College
C. L. Sagers, University of Arkansas Donald L. Williams, Sterling College
A. Edwards Salgado, Christian Brothers University Justin K. Williams, Sam Houston State University
Thomas Sasek, Northeast Louisiana University Marvin Williams, University of Nebraska–Kearney
Michael A. Savka, University of West Florida Dwina W. Willis, Freed-Hardeman University
Neil W. Sawyer, University of Wisconsin–Whitewater James A. Winsor, The Pennsylvania State University
Neil Schanker, College of the Siskiyous Chris Wolverton, Ohio Wesleyan University
Renee M. Schloupt, Delaware Valley College Kathleen Wood, University of Mary Hardin-Baylor
Bruce S. Serlin, DePauw University Richard J. Wright, Valencia Community College
Barbara Greene Shipes, Hampton University Todd Christian Yetter, Cumberland College
Richard H. Shippee, Vincennes University Brenda Young, Daemen College
Brian R. Shmaefsky, Kingwood College Rebecca Zamora, South Plains College
Shaukat M. Siddiqi, Virginia State University
Dilbagh Singh, Blackburn College
Del William Smith, Modesto Junior College Teaching and Learning
Supplements
James Smith, Boise State University
Joanna M. K. Smith
Steven Smith, University of Arizona
Nancy Smith-Huerta, Miami University Stern’s Introductory Plant Biology
F. Lee St. John, Ohio State University–Newark Lab Manual, 14th Edition by James
Spencer S. Stober, Alvernia College
Marshall D. Sundberg, Emporia State University Bidlack
Eric Sundell, University of Arkansas–Monticello
(ISBN: 9781260030143
Stan R. Szarek, Arizona State University
MHID: 1260030148)
Mesfin Tadesse, Ohio State University
Max R. Terman, Tabor College The laboratory manual that accompanies Stern’s Introductory
R. Dale Thomas, Northeast Louisiana University Plant Biology has been revised and updated. It is written for
Stephen L. Timme, Pittsburgh State University the student who is entering the study of botany. The exer-
Richard E. Trout, Oklahoma City Community College cises utilize plants to introduce biological principles and the
Jun Tsuji, Sienna Heights College scientific method. They are written to allow for maximum
Gordon E. Uno, University of Oklahoma flexibility in sequencing.
 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 on 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.
© Cliff Pelchat
OVERVIEW

This chapter introduces you to botany: what it is, how it developed, how it relates to our everyday lives, and its potential 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, the famous
botanist, Kingsley Stern, was invited to a friend’s
farm during spring break. One particular day,
Kingsley was returning to the farmhouse from a walk around
the farm, and heard groaning coming from inside. He
learned that his 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, the man had splashed some of the water in his
face, and traces of the poison had gotten into his eyes, caus-
ing 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 man’s eyes were bathed
in the milk, which contains an antidote for that poison, and
the pain subsided. Kingsley 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 poppies 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
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-
nivorous diet? Figure 1.1   Immature opium poppy capsules that were gashed with
Occasionally we hear or read of experiments—often a razor blade. Note the opium-containing latex oozing from the gashes.
© mafoto/Getty Images RF
associated with school science fairs—that suggest plants
respond in some positive way to music or soothing talk; con-
versely, some plants are said to grow poorly when they are more feet). When these giant trees are cut down, there is no
harshly yelled at. Do plants really respond to their surround- evidence of pumps of any kind within them. How, then, does
ings, and, if so, how and to what extent? water get from the roots below ground to the tops of these and
California’s huge coastal redwoods and Tasmania’s giant other trees? Do they have to expend large amounts of energy
gum trees can grow to heights of 90 or more meters (300 or to carry water such long distances (Fig. 1.3)?
2
 What Is Plant Biology? 3

Figure 1.2   Pitcher plants. Pitcher plants are carnivorous plants Figure 1.3   California coastal redwoods (Sequoia sempervirens).
that grow in nutrient-poor environments. Small insects trapped in Coastal redwoods may grow for thousands of years and some reach
their pitchers are digested and used as a source of minerals. © 2009 heights of nearly 100 meters (330 feet). © Kingsley Stern
Pia Liikala/Getty Images

Our tropical rain forests, which once covered 14% of green organisms have the exclusive capacity to produce oxy-
the earth’s land surface, now occupy only 6% of land area. gen while converting the sun’s energy into forms vital to the
They are disappearing at the rate of several acres a minute existence of both plant and animal life. At the same time,
as the plant life is cleared for agriculture, wood supplies plants remove the large amounts of carbon dioxide given
(primarily for fuel), cattle ranching, and other human off by all living organisms as they respire. In other words,
activities such as mining for gold. Experts estimate that virtually all living organisms are totally dependent on green
all rain forests may be destroyed within 40 years. Rain organisms for their existence. If some major disease were to
forests are home to 50% of all the species of living kill off all or most of the green organisms on land and in the
­organisms; it is estimated that 137 species are destroyed oceans and lakes, all the animals on land, in the sea, and in
every day due to rain-forest deforestation. What will be the air would soon starve. Even if some alternative source
the long-term impacts of these activities, and can they be of energy were available, animal life would suffocate within
reversed? 11 years—the time estimated for all the earth’s oxygen to
There is no doubt that our climate is changing. There is be completely used up if it were not replaced. Just how do
much debate, though, about the effects that global warming green plants capture the sun’s energy, use carbon dioxide,
will have on life as we know it. Are those who proclaim that and give off oxygen?
global warming will eventually have disastrous effects on This book tries to answer these and other questions about
modern civilization and living organisms simply exaggerat- living organisms—particularly those pertaining to plants, algae,
ing, or is there a scientific basis for the claims? Plant life fungi, and bacteria. Moreover, additional information about
constitutes more than 98% of the total biomass (collective plant biology related to future societies, conservation, and
dry weight of living organisms) of the earth. Plants and other human benefits is discussed.
4 Chapter 1

the future will have to include the return of organic material

KEY THEME : ecology to the soil after each harvest, instead of adding 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 abol-
Who Needs Plants? ished. Industrial pollutants will have to be rendered 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.
The Relationship of Humans Scientists and, increasingly, the general public have
become alarmed about the effects of human carelessness
to Their Environment on our environment. Damage to forests and lakes caused
by acid rain, contamination of ground water by nitrates and
It has been estimated that the total human population of the
pesticides, reduction of the ozone shield, major global cli-
world was less than 20 million in 6000 B.C. During the next
matic changes, depletion of aquifers, and loss of biodiversity
7,750 years, it rose to 500 million; by 1850, it had doubled
have gained widespread publicity.
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- Human and Animal Dependence
ulation size reached a milestone, exceeding 7 billion people.
The earth remains constant in size, but the human popula-
on Plants
tion continues to grow. Our dependence on green organisms to produce the oxy-
In feeding, clothing, and housing ourselves, we have gen in the air we breathe and to remove the carbon dioxide
had a major impact on our environment. We have drained we give off doesn’t stop there. Plants are also the sources
wetlands and cleared natural vegetation from vast areas of of products that are so much a part of human society that
land. California, for example, now has less than 5% of the we largely take them for granted. We know, of course, that
wetland it had 100 years ago. We have dumped wastes and our major crops, wheat, rice, corn, and potatoes, are plants
other pollutants into our waters and added pollutants to the (Fig. 1.4); but all foods, including meat, fish, poultry, eggs,
atmosphere. We have killed pests and plant disease organ- cheese, and milk, to mention just a few, owe their existence
isms with poisons. These poisons have also killed natural to plants. Condiments, such as spices (Fig. 1.5), and luxu-
predators and other useful organisms and, in general, have ries, such as perfumes, are produced by plants, as are some
disrupted the delicate balance of nature that existed before dyes, adhesives, digestible surgical stitching fiber, food sta-
humans began degrading their natural surroundings. bilizers, beverages (Fig. 1.6), and emulsifiers.
If we are to survive on this planet beyond the 21st century, Our houses are constructed with lumber from trees, which
there is little question that humans have to stop increasing in also furnish the cellulose for paper, cardboard, and synthetic
numbers, and the many unwise agricultural and industrial fibers. Some of our clothing, camping equipment, bedding,
practices that have accompanied the mushrooming of human curtains, and other textile goods are made from fibers of many
populations must be replaced with practices more in tune with different plant families (Fig. 1.7). Coal is fossilized plant mate-
restoring some ecological balance. Agricultural practices of rial, and oil came from microscopic green organisms or animals
 What Is Plant Biology? 5

Figure 1.5   Some of the spices derived from plants. © Kingsley

Stern

that either directly or indirectly were plant consumers. All med-

icines and drugs at one time came from plants, fungi, or bacte-
ria, and many important ones, including most of the antibiotics,
still do (Fig. 1.8). Microscopic organisms play a vital role in
recycling both plant and animal 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 even-
tually will disappear. Accordingly, the development of
alternative energy sources is receiving increased atten-
tion. In 2014, 13% of global energy consumption used
(a) renewable fuels.
The United States is the largest producer of ethanol in
the world. Corn, switchgrass, and other sources of carbo-
hydrates are currently used in the manufacture of ethanol,
which is blended with gasoline. Most cars in the United
States can run on fuel containing up to 10% ethanol. The
Energy Independence and Security Act of 2007 calls for
39 billion gallons of renewable 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
(b) persons, every one of whom needs food, clothing, and shel-
ter in order to survive. To ensure survival, we may need to
Figure 1.4   (a) Rice cakes being manufactured. Unprocessed rice learn not only how to cultivate food plants but also how to
kernels are heated, causing them to expand. The expanded kernels use plants to remove pollutants from the water, air, and soil
are then compressed into cakes, which are carried by a conveyor belt
(Fig. 1.9), to make land productive again, and to renew
to the packaging area. (b) A refrigerated produce area in a grocery
urban areas. In addition, we need to minimize the destruc-
store. (a-b) © Kingsley Stern
tion of plant habitats caused primarily by the huge increase
in the number of the earth’s inhabitants. This subject and
related matters are further discussed in Chapter 25.
6 Chapter 1

(a) 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.
Courtesy of Derrick Oosterhuis

Figure 1.8   A Penicillium colony. The tiny beads of fluid on the

surface contain penicillin, widely used as an antibiotic. © Kingsley Stern

(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 Figure 1.9   A polluted waterway. Source: U.S. Coast Guard photo
roasted. (a-b) © Kingsley Stern by Petty Officer 3rd Class Zac Crawford
 What Is Plant Biology? 7

At present the idea that humanity may not be able to or incorrect. To be accepted by scientists, the results of any
save itself may seem radical, but there are a few who have experiments designed to test the hypothesis must be repeat-
suggested that it might become necessary in the future to able and capable of being duplicated by others.
emigrate to other planets. Regardless of humanity’s future, A scientific experiment is typically carried out with
it is essential that our understanding of plants be used to sus- a test group of plants and a control group. The test group
tain life on this and maybe even other planets. Experiments receives the experimental treatment, while the control group
with portable oxygen generators have been in progress for is treated the same in all ways except that it does not receive
many years. Tanks of water teeming with tiny green algae the treatment. For example, if you would like to test the
are taken aboard a spacecraft and installed so that they are effect of light on seed germination, you would divide your
exposed to light for at least part of the time. The algae not seed lot and place some in the light and some in the dark.
only produce oxygen, which the spacecraft inhabitants can All other environmental conditions, such as temperature and
breathe, but also utilize the waste carbon dioxide produced humidity, would be the same for both sets of seeds. If you
by respiration. As the algae multiply, they can be fed to a observe a germination difference between the two sets of
special kind of shrimp, which in turn multiply and become seeds, then it may be due to light.
food for the space travelers. Other wastes are recycled by When a hypothesis is tested, data (bits of information)
different microscopic organisms. When this self-supporting are accumulated and may lead to the formulation of a useful
arrangement, called a closed system, is perfected, the range generalization called a principle. Several related principles
of spacecraft should greatly increase because heavy oxygen may lend themselves to grouping into a theory, which is
tanks will not be necessary, and the amount of food reserves not simply a guess. A theory is a group of generalizations
will be reduced. (principles) that help us understand something. We reject
Today, teams of botanists, anthropologists, and medi- or modify theories only when new principles increase our
cal doctors are interviewing medical practitioners and understanding of a phenomenon. An example of a theory
herbal healers in remote tropical regions and taking notes is the concept of natural selection and evolution. This the-
on various uses of plants by the local inhabitants. These ory describes how the diversity of life on earth came about
scientists are doing so in the hope of preserving at least and provides the foundation for many principles you will
some plants with potential for contributions to modern civi- encounter in this textbook.
lization before disruption of their habitats results in their While the testing of hypotheses is a rigorous, well-
extinction. 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
Botany as a Science scientists are those who can think “outside the box,” inte-
grating observations across fields or organisms to create a
The study of plants, called botany—from three Greek words, hypothesis that explains a complex phenomenon. While sci-
botanikos (botanical), botane (plant or herb), and boskein entific advances are often attributed to luck, there is a grow-
(to feed), and the French word botanique (botanical)— ing body of evidence that the methods used by scientists
appears to have had its origins with Stone Age peoples who actually harness the unexpected to the benefit of science.
tried to modify their surroundings and feed themselves. At When the results of an experiment do not follow predictions,
first, their interest in plants was mostly practical and cen- that experiment will be repeated with extra care and from a
tered around how plants might provide food, fibers, fuel, and new perspective. If the unexpected observation is observed
medicine. Eventually, however, an intellectual interest arose. again, then the scientist must develop a new hypothesis.
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.”
Diversification of Plant Study
Botanists are scientists who study plants. Plant anatomy, which is concerned chiefly with the inter-
A science is distinguished from other fields of study by nal structure of plants, was established through the efforts
its basis in measurable observations. The scientific method is of several scientific pioneers. Early plant anatomists of note
a series of steps that involves first asking a question, then for- included Marcello Malpighi (1628–1694) of Italy, who dis-
mulating a hypothesis, followed by conducting experiments, covered various tissues in stems and roots, and Nehemiah
and finally developing a theory. Grew (1628–1711) of England, who described the structure of
wood more precisely than any of his predecessors (Fig. 1.10).
Hypotheses Today, knowledge of plant anatomy is used to help us
find clues to the past, as well as for many practical purposes.
A hypothesis is simply a tentative, unproven explanation for For example, the related discipline of dendrochronology
something that has been observed. It may not be the correct deals with determining past climates by examining the width
explanation—testing will determine whether it is correct and other features of tree rings. We can also learn much from
8 Chapter 1

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
Figure 1.10   A thin section of Magnolia wood as seen through his system of classification is given in Chapter 16.
a light microscope. ×40. © Kingsley Stern
There are still thousands of plants, fungi, and other
organisms that have not yet been described or even discov-
archaeological sites by matching tree rings found in the wood ered. Although it obviously is already too late to identify
of ancient buildings to the rings of wood of known age. Plant species that were not described before they became extinct,
anatomy is also used to solve crimes. Forensic laboratories plant taxonomists around the world have united to try to
may use fragments of plant tissues found on clothing or identify and describe as many new organisms as possible—
under fingernails to determine where a crime took place or if many with food, medicinal, and other useful potential—
certain persons could have been present where the crime was before much more of their natural habitat disappears. Other
committed. The anatomy of leaves, stems, and other plant plant taxonomists, through the use of cladistics (analysis
parts is used to unravel and sort out relationships among of shared features) and molecular techniques, are refining
plants. A form of plant anatomy, known as paleobotany, our knowledge of plant relationships. By the year 2000 we
involves the study of plant fossils. had acquired so much new information about natural rela-
Plant physiology, which is concerned with plant func- tionships that some major reclassification took place (see
tion, was established by J. B. van Helmont (1577–1644), Chapter 16). The molecular knowledge and techniques are
a Flemish physician and chemist, who was the first to also contributing to the improvement of many of our food
­demonstrate that plants do not have the same nutritional crops, although some of the changes are controversial.
needs as animals. In a classic experiment, van Helmont Many plant taxonomists specialize in certain groups of
planted a willow branch weighing 5 pounds in an earthen- plants. For example, pteridologists specialize in the study of
ware tub filled with 90.7 kilograms (200 pounds) of dry ferns; bryologists study mosses and plants with similar life
soil. He covered the soil to prevent dust from settling on it cycles.
from the air. The willow produced roots and grew, and after The discipline of plant geography, the study of how
5 years, he reweighed the willow and the soil. He found that and why plants are distributed where they are, did not
the soil weighed only 56.7 grams (2 ounces), less than it develop until the 19th century (Fig. 1.11). The allied field of
had at the beginning of the experiment, but that the willow plant ecology, which is the study of the interaction of plants
had gained 76.7 kilograms (169 pounds). He concluded that with one another and with their environment, also developed
the tree had added to its bulk and size from the water it had in the 19th century.
absorbed. We know now that most of the weight came as a After the publication in 1962 of a best-seller entitled
result of photosynthetic activity (discussed in Chapter 10), Silent Spring (authored by Rachel Carson), public awareness
but van Helmont deserves credit for landmark experimenta- of the field of ecology as a whole increased considerably. In
tion in plant physiology. this book, based on more than 4 years of literature research,
Modern plant physiologists use cloned genes (units of Carson noted that more than 500 new toxic chemicals are
heredity that are found mostly within the nuclei of cells) put to use annually as pesticides in the United States alone,
to learn in precise detail much more about plant functions, and she detailed how these chemicals and other pollutants
including how plants conduct materials internally; how tem- are having a negative impact on all facets of human life and
perature, light, and water are involved in growth; why plants the environment.
flower; and how plant growth regulatory substances are pro- The study of the form and structure of plants, plant
duced, to mention just a few. morphology, was developed during the 19th century, and
 What Is Plant Biology? 9

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. © Nadezda Zavitaeva/
Shutterstock.com

during the 20th century much of our basic knowledge about components perform and integrate a variety of functions,
the form and life cycles of plants was incorporated into the including that of sexual reproduction. The mid-20th-century
plant sciences as we know them today. During this time, the development of electron microscopes (see Chapter 3) further
number of scientists engaged in investigating plants also spurred cell research and led to vast new insights into cells
greatly increased. and new forms of cell research that continues to the present.
Genetics, the science of heredity, was founded by the Economic botany and ethnobotany, which focus on
Austrian monk Gregor Mendel (1822–1884), who per- practical uses of plants and plant products, had their origin
formed classic experiments with pea plants. Today, vari- in antiquity as humans discovered, used, and eventually cul-
ous bran­ches of genetics include plant breeding, which tivated plants for food, fiber, medicines, and other purposes.
has greatly improved the yields and quality of crop plants, Today there is increased interest in herbal medicines (see
and genetic engineering. Genetic engineering involves the Appendix 3) and many other uses of plants by the general
transfer of genes from one organism to another and has public. Research is being conducted in collaboration with
already improved the pest, frost, and disease resistance indigenous peoples with an eye to discovering new medi-
of some crop plants. Genetic engineering is being used cines and other useful plant products previously unknown in
to develop better agricultural, medicinal, and other use- developed countries.
ful plants. It is also being used to control human, animal, There is still a vast amount of botanical information to
and plant diseases. In recent years, the field of genomics, be discovered. Dozens of research journals publish thou-
which focuses on genes and their function, has burst onto sands of botanical scientific papers every year. In recent
the genetics scene and now impacts nearly all genetics years, open source journals have allowed free online access
research. to research findings for scientists all over the world. While
Cell biology (previously called cytology), the sci- scientists have identified about 2 million species of organ-
ence of cell structure and function, received a boost from isms, the earth likely contains between 5 and 100 million
the discovery of how cells multiply and how their various species.
10 Chapter 1

Plant Biology on the Internet

P eople use the Internet for direct communication, edu-

cation, and recreation, among other things, and do
so through the use of various devices. The recent
inception of wireless connection has increased worldwide
use of the Internet because it allows access to information
virtually anywhere and by any device that can pick up a sig-
nal. Computers have revolutionized the study of every field
of plant biology. Portable computers with Internet access
(Box Figure 1.1) can be taken to the field to help navigate
through the wilderness, locate and identify specimens, and
interact directly with scientists at other locations to exchange
information . Even more popular are the numerous handheld
devices such as the Android, BlackBerry, iPhone, and other
smartphones, which are smaller and less expensive than the
bulkier systems. These handheld devices include mobile
phone access, which makes them among the most popular
Internet access tools on the planet. Box Figure 1.1   A botany student uses the Internet to identify
and learn about a botanical specimen in the field. © James E. Bidlack
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.
5. Australian National Botanic Gardens provides a wealth of
Sites that include materials on botany education, botanical
botanical and biological information about Australia. http://
gardens, plant databases, and videos about plants are a few
www.anbg.gov.au/gardens/
of the many resources available on the Web. The following
6. The United States Department of Agriculture contains
are some interesting websites that can be explored. Try them
news and information about the nation’s agricultural
out sometime! Maybe you’ll find a good idea for a research
economy. There is an excellent section on the history of
paper.
American agriculture from 1776 to 2000. http://www.usda.
1. The Multimedia Educational Resource for Learning gov/
and Online Teaching (MERLOT) includes many learning 7. GardenWeb is an information center for gardening enthu-
objects for plant biology and other disciplines. This web- siasts. http://www.gardenweb.com/
site includes a search engine as well as peer-reviewed 8. California Flora Database contains geographic and eco-
learning materials for many topics in higher education. logical distribution information for 7,975 California vascu-
http://www.merlot.org/ lar plant taxa, as well as additional habitat information for
2. The Missouri Botanical Garden is one of the oldest rare taxa and species of the Sierra Nevada. http://www.
botanical institutions in the United States. It is a center for calflora.org/
botanical research and science education. http://www. 9. Common Conifers of the Pacific Northwest provides infor-
mobot.org/ mation about the conifers of Oregon, including a dichoto-
3. The New York Botanical Garden is situated on 250 acres mous key for their identification. http://www.oregonstate.
in the Bronx and includes 27 outdoor gardens and plant edu/trees/
collections, the nation’s most beautiful Victorian 10. Carnivorous Plants Database includes over 3,000 entries,
conservatory, and a 40-acre presettlement forest. http:// giving an exhaustive nomenclatural synopsis of all
www.nybg.org/ carnivorous plants. http://www.omnisterra.com/bot/
4. The Arnold Arboretum of Harvard University is the cp_home.cgi
nation’s oldest arboretum. The site includes a catalog 11. Plants in Motion provides short video clips of different
of over 5,000 kinds of woody plants cultivated in the types of plant movement, including germination, flower-
arboretum as well as educational and visitor information. ing, and tropic responses. http://plantsinmotion.bio.
http://www.arboretum.harvard.edu/ indiana.edu/
 What Is Plant Biology? 11

SUMMARY plants. In recent years, molecular and cladistical

investigations have resulted in modifications of
Linnaeus’s system.
1. Why do some plants produce poisons, while others are
edible and useful? Why are some plants carnivorous? 12. During the 19th century, plant ecology, plant
How and why do plants respond to their environment? geography, and plant morphology developed, and by
What is the future of tropical rain forests? What can be the beginning of the 20th century, genetics and cell
done about global warming and other environmental biology became established. Much remains yet to be
problems? This book addresses these questions and discovered and investigated.
more about plants.
2. Human populations have increased dramatically in the
REVIEW QUESTIONS
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 1. How and to what extent have humans affected their
people threatens the survival of not only humans but natural environment?
other organisms as well. 2. What is meant by the scientific method?
3. We are totally dependent on green organisms because 3. To what extent is animal life dependent on green
they alone can convert the sun’s energy into forms organisms for its existence?
that are usable by, and vital to the very existence of, 4. In terms of biological experiments, what are hypotheses
animal life. and controls?
4. We largely take plants and plant products for granted. 5. What is the oldest branch of botany, and why did it
Animals, animal products, many luxuries and precede other branches?
condiments, and other useful substances such as fibers, 6. What are the basic features of each of the other
lumber, coal, medicines, and drugs either depend on branches of botany?
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 DISCUSSION QUESTIONS
them. Plants will undoubtedly play a vital role in space
exploration as portable oxygen generators. 1. Since humans survived on wild plants for thousands of
6. Teams of scientists are interviewing medical years, might it be desirable to return to that practice?
practitioners and herbal healers in the tropics to locate 2. What factors are involved in determining if and when
little-known plants used by local inhabitants before the humans might not be able to sustain themselves on this
plants become extinct. planet?
7. Botany, the study of plants, apparently began with 3. How do you suppose that Stone Age peoples discovered
Stone Age peoples’ practical uses of plants. Eventually, medicinal uses for plants?
botany became a science as intellectual curiosity about 4. Why do you suppose that many of the early botanists
plants arose. were also medical doctors?
8. A science involves the observation, recording, 5. Consider the following hypothesis: “The majority
organization, and classification of facts. The verifying of mushrooms that grow in grassy areas are not
or discarding of facts is done chiefly from known poisonous.” How could you go about testing this
samples. The scientific method involves following hypothesis scientifically?
a routine series of steps and generally assuming and
testing hypotheses.
9. The microscope has had a profound effect on studies ADDITIONAL READING
in the biological sciences and led to the discovery
of cells. Capon, B. 2010. Botany for gardeners. 3d ed. Portland, OR:
10. Plant anatomy and plant physiology developed during Timber Press.
the 17th century. J. B. van Helmont was the first to Carson, R. L. 2002. Silent spring. Boston: Houghton Mifflin
demonstrate that plants have nutritional needs different Harcourt. Chalker-Scott, L. 2015. How plants work:
The science behind the amazing things plants do. OR:
from those of animals. During the 17th century,
Timber Press.
Europeans engaged in botanical exploration on other
Coyle, H. M. 2004. Forensic botany. New York: Taylor and Francis.
continents and took plants back to Europe.
Jacobs, P. F., and J. Brett. 2004. Guide to information sources
11. During the 18th century, Linnaeus produced the in the botanical sciences. Englewood, CO: Libraries
elements of a system of naming and classifying Unlimited.
 Rev.confirming Pages

12 Chapter 1

Johnson, T. 1998. CRC ethnobotany desk reference. Boca Raton, van Wyk, B. 2005. Food plants of the world: An illustrated guide.
FL: CRC Press. Portland, OR: Timber Press.
Minnus, P. E. 2000. Ethnobotany. Norman: University of
Oklahoma Press. LEARNING ONLINE
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 Visit our website at http://connect.mheducation.com for additional
plants, 1620–1900. Portland, OR: Timber Press. information and learning tools.

bid82749_ch01_001-012.indd 12 03/15/17 05:03 PM

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T HOU seemest to have been long from me; let nothing any longer
detain thee, but my sister’s necessity, or father’s authority. I
am in a comfortable state of health, through divine goodness, to
which be glory for ever. See that thou love and admire that fountain
of our life, and peace: and be ever mindful that ’tis all thy business to
love, and serve, and praise thy Creator and Redeemer. I have no
other business but this to write to thee about: but this is all our
business. What we use to call business is but vanity and pastime in
comparison of this. Remember that ’tis thy one end, to glorify God,
and enjoy him for ever. Learn well that lesson, and know, that it is the
only thing necessary. Every morning remember that thy serving and
pleasing God is the whole business of that day, and therefore set out
accordingly with an express design and intention to please God in
thy eating, drinking, visiting, conversing, and duties throughout the
day. My most dear heart, I have nothing in the world that doth
concern thee, or me so much to write of to thee, as this. Oh that thou
mayst still be laying up in heaven! Still furthering thy account; still
adding to the heap, and increasing thy glorious reward? Nothing is
done for God, but thou shalt hear of it again. What is not done for
God, is but so much lost. Those things which others do, being led by
their natural affections and desires, those things do thou with holy
aims, for spiritual ends, and then God will put it on the account, as so
much done for him. So it is, my dearest, God keeps a true account.
See that thou believe it, and so plough in hope, and sow in hope,
pray and hear with an eye to the sure reward. Let thy hopes be
strong and lively, and then thy hands will be strong, and thy
resolutions and affections will be strong. My time is very precious,
and I would not lose an inch of it. See thou to it, that my time in
writing this letter be not lost time. Love God the more, and set thy
heart the straiter towards him, and practise this one thing, in every
action look to thy end, and then I have got well, and thou better by
these counsels. My dearest, I love thee in truth and tenderness, but
my love signifies little, unless it serve thine eternal good.

I rest thine own,

JOS. ALLEINE.
 [To his wife.]

My dear heart,

M Y heart
* is now a little at rest to write to thee. I have been these
three days much disturbed. Strong sollicitations I have had
from several hands, to accept very honourable preferment;
but I have not found the invitations to suit with the inclinations of my
own heart, as I was confident they would not with thine. I have sent
away my friends satisfied with the reasons of my refusal, and now
can say, Soul, return unto thy rest. But alas, that such things should
disturb me! I would live above this lower region, that no passages
whatsoever might put me out of frame, or unsettle me from my
desired rest. I would have my heart fixed upon God, so as no
occurrences might disturb my tranquility, but I might be still in the
same quiet and even frame. Well, though I am apt to be unsettled,
yet I am like a bird out of the nest, I am never at quiet till I am in my
old way of communion with God; like the needle in the compass, that
is restless, till it be turned towards the pole.

I can say through grace, with my soul have I desired thee in the
night, and with my spirit within me have I sought thee early; my heart
is early and late with God, ’tis the business and delight of my life to
seek him. But alas, how long shall I spend my days in wishing, when
my glorified brethren spend theirs in enjoying? As the poor
imprisoned captive sighs under his irons, and can only look through
the grate, and long for the liberty which others enjoy: such is my
condition. I can only look through the grate of this prison, my flesh; I
see Abraham, and Isaac, and Jacob, sitting down in the kingdom of
God. But alas, I myself must stand without, longing, praying, waiting,
for what they are enjoying. Happy souls! When shall these fetters of
mine be knocked off? When shall I be set at liberty from this prison
of my body? You are cloathed with glory, when I am cloathed with
dust. I dwell in flesh, in a house of clay, when you dwell with God in a
house not made with hands, eternal in the heavens.
 I must be continually clogged with this cumbersome body, when
you have put on incorruption and immortality. What continual
molestation am I subject to by reason of this flesh? What pains doth
it cost me to keep this earthen vessel from breaking! It must be
exercised; and which is worst of all, cherished with time-devouring
sleep; so that I live but little of the short time I have allotted me here.
But oh blessed souls, you are swallowed up of immortality and life,
your race is run, and you have received your crown. How cautious
must I be to keep me from dangers! How apt am I to be troubled with
the cares and fears of this life, when your souls are taken up with
God and Christ, and ’tis your work to be still contemplating, and
admiring that love that redeemed you from all this. What pains must I
be at to repair the ruinous building of this earthly tabernacle, which
when I have done, I am sure will shortly fall about my ears; when you
are got far above mortality, and are made equal with the angels. Oh!
I groan earnestly to be cloathed upon with my house which is from
heaven, being willing rather to be absent from the body and present
with the Lord! Oh, when shall I come and appear before him? When
shall I receive the purchase of my Saviour, the fruit of my prayers,
the harvest of my labours, the end of my faith, the salvation of my
soul? Alas, what do I here? This is not my resting place, my treasure
is in heaven. Oh when shall I be where my heart is? Wo is me that
dwell in the tents of Kedar! Oh that I had wings like a dove, that I
might fly away and be at rest! Then would I hasten my escape from
the storm and tempest, and be out of the reach of fears,
disturbances, and distractions. How long shall I live at such a
distance from my God, at such a distance from my country? Alas,
how can I sing the Lord’s song in a strange land? No, I will hang my
harp upon the willows, and sit down and weep when I remember
Sion. But yet my flesh shall rest in hope, and I will daily bathe my
soul in the sweet thoughts of my blessed home. I will rejoice in
hopes of what I do not yet enjoy, and content myself with the taste of
what I shall shortly have my fill of.
 The Lord grant the request I daily pour out before him, and make
us furtherances to each other’s soul, that we may quicken and
promote and forward one another in his ways! Help me by thy
prayers, as thou dost always. The God of all peace and comfort be
with thee my sweet love! Farewell!

Thine beyond expression,

JOS. ALLEINE.

L E T T E R XXVI.
[God is a satisfying Portion.]

My most dear friend,

H AD not my right hand long since forgot her cunning, and the
Almighty shook the pen out of my hand, I should long e’er this
have written to thee; but it is a wonder of divine power and goodness
that my soul had not before this time dwelt in silence, and that death
had not put the long period to all my writing and converse.
 Long is the song of love that I have to tell thee. I rejoice in the
constancy of thy love, that the waters of so long a silence, and so
great a distance have not yet quenched it. But thy desires are
towards me, and thy heart is with me, though providence hath
hindered me from thy much-desired company. I will assure thee it
hath been a pleasure to my heart a good part of this summer, to
hope that I should come one half of the way to give thee a meeting.
But such is my weakness hitherto, that I am forced to put off those
hopes till the spring, when, if God gives me strength to ride, I intend
to see thee before mine own home. I thank thee for all the dear
expressions of thy fervent love: my expences have been vast; but
surely goodness and mercy hath followed me, and do follow me in
every place, and in every change of my condition; so that as to
temporals, I have lack of nothing, and as for spirituals I abound and
superabound, and the streams of my comforts have been full and
running over. The joy of the Lord hath been my strength at the
weakest, and in the multitude of my thoughts within me, his comforts
have refreshed my soul. I have found God a satisfying portion to me,
and have sat under his shadow with full delights, and his fruit is most
sweet to my taste: he is my strength ♦and my song, for I will talk of
him, and write of him with perpetual pleasure. Through grace I can
say, methinks I am now in my element, since I have begun to make
mention of him, I am rich in him and happy in him, and my soul saith
unto him with David, Thou hast made me most blessed for ever
more. Happy is the hour that ever I was born, to be made partaker of
so blissful a treasure, so endless a felicity, such angelical a
prerogative, as I have in him: O how sweet are his converses, how
delightful it is to triumph in his love.

♦ duplicate word “and” removed

 Methinks the story of the lepers comes not unaptly to my mind,
who said one to another when they had eat and drunk and carried
away silver and gold and raiment, and went and hid it, We do not
well; this day is a day of good tidings, and we hold our peace. It is fit
that I should be cloathed with shame; I acknowledge before God,
who trieth the hearts I am unworthy, everlastingly unworthy. But it is
not fit that he should lose his praise; nay rather let him be the more
adored, and magnified and admired for ever and ever. Bless the
Lord, O my soul, bless the Lord, O my friend; let us exalt his name
together. He is my solace in my solitude; he is my standing
comforter, my tried friend, my sure refuge, my safe retreat; he is my
paradise, he is my heaven; and my heart is at rest in him: and I will
sit and sing under his shadow, as a bird among the branches. And
whither should I go but unto him? Shall I leave the fatness of the
olive, and the sweetness of the fig-tree, and of the vine, and go and
put my trust under the shadow of the bramble? No, I have made my
everlasting choice: this is my rest for ever, he is my well-beloved, in
whom I am well-pleased. Suffer me to boast a little: here I may glory
without vanity, and I can praise him without end or measure; but I
have nothing to say of myself: I find thou dost over-value me; set the
crown upon the head of Christ; let nothing be great with thee but
him, give him the glory. God that knoweth all things, knoweth my
poverty, how little, how low, and how mean I am, and how short I
come of the attainments of the saints, who yet themselves come so
exceedingly short of the rule that God hath set before us. I often
think of the complaint of the devout Monsieur De Renty [I feel myself
very poor this week; and very defective in the love of God; if you
would know wherein you can pleasure me, love God more: that what
is wanting in me may be made up in the abundance of your love:] in
this thou mayest highly pleasure me: love God a little the better,
praise him a little the more for my sake; let me have this to please
myself in, that God is a little the better loved for me, and that I have
blowed up, if it be but one spark of divine love in the bosom of my
dearest friend, towards him.
 Thy cautions are acceptable to me, I desire to provide for
manifold changes and storms. I know I am not yet in the harbour; O
pray with me that I may not enter into temptation; for I am very weak
in spirit, as well as in body, God knoweth. Somewhere or other I
must break off, and thou wilt say, it is time to shut up. For once only
know, that I am thy daily orator, and will be whilst I am. And yet once
more, I must have room to add my thankful acknowledgment of
thine. With our most dear affections to you both, I commend you to
the God of love, still abiding,

Thy fast and sure

F R I E N D.

Bath, October 12, 1668.

L E T T E R XXVII.
Dear Cousin,

T HE welcome tidings of your safe arrival at Barbadoes is come to

my ears; as also the news of your escape from a perilous
sickness, for which I bless the Lord. I have considered, that God had
bereft you of a careful father, and that your mother takes but little
care for you; so that you have none nearer than myself to watch for
your soul, and to charge and admonish you in the Lord.
 But yet, be not discouraged by these things, but look to heaven,
fly unto Jesus, put away every known sin, set upon the conscientious
performance of every known duty; make Christ your choice, embrace
him upon his own terms; deliver up yourself, body and soul to him:
see that you have no reserves nor limitations in your choice of him;
give him your very heart; cast away your worldly hopes and
expectations, make religion your business.

These things do, and you shall be sure of a friend in heaven; and,
if I may be any comfort to you, you shall not fail, while I live, to have
one friend on earth to care for you. You are gone far from me, even
to the uttermost parts of the earth: but I have sent these letters to call
after you; yea, not only to call, but to cry in your ears. O what is like
to become of your soul! Where is that immortal soul of yours like to
be lodged for ever? Amongst devils or angels? Upon a bed of
flames, or in the joys of paradise?

Go aside; retire from the noise of the world, and say to yourself,
Oh my soul! Whither art thou going? Do not I know, that I must be
converted or condemned? That I must be sanctified, or I can never
be saved? Oh my soul! What seekest thou? What is my chief care?
Is it for this world, or the world to come? Do I first seek the kingdom
of heaven, and the righteousness thereof? Do I think heaven will
drop into my mouth? That glory and immortality will be got with a wet
finger, with cold prayers, and heartless wishes, while the world has
my heart? Do I think to be crowned, and yet never fight? To gain the
race and never run? To enter at the strait gate, and never strive? To
overcome principalities and powers, and never wrestle? No, no; Oh
my soul, either lay by the hopes of heaven for ever, or rouse up
thyself, put forth thy strength after God and glory. Either lay by thy
worldly hopes, or thy hopes of immortality; away with thy sins, or let
Christ go for ever. Think not to have Christ and the world too, to
serve God and mammon: if thou follow the world, thou must die: the
Lord hath spoken it, and all the world can never reverse it. Thus
reason the case with your own soul, and give not rest to thyself night
nor day, till you are gotten off from the world, broken off from every
known sin, and got safe into Christ.
 Dear cousin, I charge you by the Lord, to observe these things.
Pray over them, weep over them, read them again and again; do not
pass them over as slight and ordinary things. Your soul is at stake; it
is your salvation which is concerned in them; think not that I am in
jest with you. I travel in birth with you, till Christ be formed in you.
Why should you die? Oh repent and live, lay hold on eternal life, win
Christ, and you win all. Oh be thankful to the Lord, that now you are
fatherless and friendless, yet you have one remembrancer to warn
you to flee from the wrath to come. God forbid that I should find you
at last in the place of torments, for your not embracing these
counsels. To conclude, I charge you as a minister, as a father, take
heed of these three things:

1. Lest the gain of the world prove the loss of your soul:

2. Lest company draw you from God:

3. Lest a lofty or a worldly heart should thrust you out of the

kingdom of heaven.

Oh labour whatever you do for an humble heart. Be little, be vile

in your own eyes; seek not after great things; be poor in spirit:
without this, heaven will be no place for you. Your lot is fallen in a
place of great wickedness, where your soul is in much danger, where
your temptations are many, and your helps for heaven but few:
where good examples are rare, and many will entice ♦you to sin and
vanity. O! look about you, consider your danger, fear lest you should
miscarry for ever. I can but warn you and pray for you: but though
you have none to oversee you, remember the eye of God is upon
you, to observe all your actions, and that he will surely bring all your
practices into judgment. I commend you to the Lord, and remain,

Your loving and careful uncle,

JOS. ALLEINE.

August 19, 1668.

 ♦ “yo” replaced with “you”

L E T T E R XXVIII.
Dear friend,

* OUR letter was exceeding welcome to me, not only as reviving

Y the remembrance of our old friendship, but also, as bringing
me news of some spiritual good that you received by me,
which is the best tidings that I can receive: For what do I live for, but
to be useful to souls in my generation? *I desire no other business
than to please and honour my God, and serve my generation in that
short allowance of time I have here. Shall I commend to you the
lesson that I am about to learn? It is, to be entirely devoted to the
Lord, that I may be able to say after the apostle, To me to live is
Christ. I would not be serving God only for a day in the week, or an
hour or two in the day: but every day, and all the day. I am ambitious
to come up to that of our Lord and Master, To do always those things
that please God. I plainly see that self-seeking is self-undoing; and
that then we promote ourselves best, when we please God most. I
find, that when I have done all, if God be not pleased, I have done
nothing; and if I can but approve myself to God, my work is done: I
reckon I do not live that time I do not live to God.
 *I am fain to cut off so many hours from my days, and so many
years from my life, as I have lived to myself. I find no enemy so
dangerous as myself, and O that others may take warning by my
hurt: O that I had lived wholly to God! Then had every day and every
hour that I have spent, been found upon my account at that day:
then had I been rich indeed, in treasure laid up there, whither I am
apace removing; then I had been every day and hour adding to the
heap, and increasing the reward which God of his mere grace hath
promised, even to the meanest work that is done to him. I perceive I
am an eternal loser by acting no more for God; for what is done to
myself is lost; but what is done for God, is done for ever, and shall
receive an everlasting reward. Verily, if there be a world to come,
and an eternal state after this short life, it is our only wisdom to be
removing, and, as it were transplanting and transporting what we
can, from hence into that country to which we are shortly to be
removed, that what we are now doing we may reap the fruit of for
ever.
 Well, let us be wholly swallowed up in religion, and know no other
interest but Jesus Christ. I cannot say, I have already attained; but
this is what my heart is set to learn. That in all that I do, whether
sacred or civil actions, still I may be doing but one work, and driving
on one design, That God may be pleased by me, and glorified in me;
That not only my praying, preaching, alms, may be found upon my
account; but even my eating, drinking, sleeping, visits, discourses,
because they are done to God. Too often do I miss my mark; but I
will tell you what are the rules I set myself: Never to lie down but in
the name of God; nor barely for natural refreshment, but that a
wearied servant of Christ may be recruited and fitted to serve him
better the next day. Never to rise up but with this resolution, I will go
forth this day in the name of God, and will make religion my
business, and spend the day for eternity. Never to enter upon my
calling, but first thinking, I will do these things as unto God, because
he requireth these things at my hands, in the place and station he
hath put me into. Never to sit down to the table, but resolving, I will
not eat meerly to please my appetite, but to strengthen myself for my
Master’s work. Never to make a visit, but to leave something of God
where I go; and in every company to leave some good savor behind.
This is that which I am pressing hard after: and if I strive not to walk
by these rules, let this paper be a witness against me.

I perceive you are otherwise persuaded in some things than I am:

but however, I trust we meet in our end. May it be your whole study
to gain souls, and to build them up in holiness, which is with too
many the least of their cares. One duty (miserably neglected) I shall
be bold to recommend from my own experience, and that is, the
visiting your whole flock from house to house, and enquiring into
their spiritual estates particularly, and dealing plainly and truly with
them about their conversion to God.

You see how free I am with you: but I know your candour. I
rejoice in your happy yoke-fellow: salute her from your old friend,
and accept of the unfeigned respects of him who is, Sir,

Your real and faithful friend,

JOS. ALLEINE.
 L E T T E R XXIX.
[To a minister in prison.]

Worthy Sir,

I T was but a little after my release from my own confinement, but I

heard of yours: and now write to you, as one that hath taken a
higher degree than ever, being commenced prisoner of Christ. I was
once affected with the picture of a devout man, to whom a voice
came down from heaven, saying, Quid vis fieri pro te? To which he
answered, Nihil domine, nifi pati ac contemini pro te. Undoubtedly,
Sir, it is our real glory to be throughout conformed to Jesus Christ,
not only in his sanctity, but in his sufferings. I doubt not your
consolations in Christ superabound in all your tribulations for him.
Yet let me add this, that you have a whole shoal of promises come in
to you, which you had not before; I mean all the promises to
suffering saints, in which they have not so immediate a part, unless
in a suffering state. And doubtless he hath got well, that hath gotten
such a number of exceeding great and precious promises.

I can tell you little good of myself: but this I can tell you, that the
promises of God were never so sweet to me, as since my
imprisoned state. It shames me that I have let such a treasure lie by
so long, and have made so little use of it. Never did my soul know
the heaven of a believer’s life, till I learnt to live a life of praise, and
to set home the unspeakable riches of the divine promises, to which,
through grace, I am made an heir. I verily perceive that all our work
were done, if we could but prevail with ourselves and others to live
like believers; to tell all the world by our carriage, that there is such
pleasantness in Christ’s ways, such beauty in holiness, such reward
to obedience, as we profess to believe!
 It is but a little while that prisons shall hold us, or that we shall
dwell in dirty flesh. Porphyry tells us of Plotinus, that he was
ashamed to see himself in the body; to see a divine and immortal
soul in a prison of flesh (for so they held the body to be;) but the
worst shackles are those of sin. Well, they must shortly fall off; our
Lord doth not long intend us for this lower region: surely he is gone
to prepare a place for us: yea, and he will come again, and receive
us to himself, that where he is, we may be also. And what have we
to do, but to believe, and wait, and love, and long, and look out for
his coming, in which is all our hope? ’Twill be time enough for us to
be preferred then. We know before hand who shall then be
uppermost. Our Lord hath shewed us where our places shall be,
even at his own right hand; and what he will say to us, Come ye
blessed. Surely we shall stand in his judgment: he hath promised to
stand our friend: let us look for the joyful day: and sure as there is a
God, this day will come, and then it shall go well with us. What if
bonds and banishment abide us for a season? This is nothing but
what our Lord hath told us, The world shall rejoice, but ye shall
lament: ye shall be sorrowful, but your sorrow shall be turned into
joy. Oh how reviving are his words! I will see you again, and your
heart shall rejoice, and your joy no man taketh from you.

If that miserable wretch leapt chearfully off the ladder, saying, I

shall be a queen in hell: with what joy should we do and suffer for
God, knowing that we shall be crowned in heaven? They are
wonderful preparations that are making for us: the Lord make us
meet to be partakers. *It was the highest commendation that ever
that worthy R. Baxter received, which fell from his scoffing adversary
Tilenus, Totum, puritanismum totus spirat. Oh that this may be true
of us and ours!

Let your true yoke-fellow, and my Christian friends with you, have
my hearty commendation: and these counsels I pray you give them
from me.
 1. To habituate themselves, both as to their thoughts and
discourses, more throughly than ever to holiness. Brethren, I would
teach you the lesson that I resolve to learn, that your minds and
tongues may as naturally run upon the things of heaven, as others
on the things of this world. Why should it not be thus? I am sure God
and heaven as well deserve to be thought on, and talked of as froth
and vanity. There are many that have in a great measure learnt this
lesson, and why should not we? What if it be hard at first? Every
thing is so to a beginner. And is not ours a religion of self-denial? If
we do but force ourselves awhile to holy thoughts, and heavenly
discourse, it will grow habitual to us, and then it will be most natural,
familiar, and sweet. O what gainers will you be, if you learn this
lesson?
 ’Tis the shame of religion, that Christians are so unlike
themselves, unless upon their knees. Our lives and language should
tell the world what we are, and whither we are going. Christians, let
little things content you in the world, but aspire after great things in
the grace of God. Many little think what high degrees of holiness
they may grow up to even in this life with pains and diligence. Sirs,
be you men of great designs: think it not enough if you have
wherewith to bear your charges to heaven; but aspire to be great in
the court of heaven, favourites of the Most High, of tall growth,
singular communion, that you may burn and shine in your place, that
you may savour of heaven wherever you come, and that there may
be an even-spun thread of holiness running through your whole
course. ’Tis our disgrace, that there is so little difference to be seen
in the ordinary conversation of believers and other men. Is it not a
shame, that when we are in company with others, this should be all
the difference that is to be seen, that we will not curse and swear? If
you will honour the gospel, bring forth your religion out of your
closets into your shops, trades, visits, and exemplify the rules of
religion in the management of all your relations, and in your ordinary
converse. Let there be no place or company that you come into, in
which you do not drop something of God; this will be the glory of
religion, and we shall never convince the world ’till we come to this.
May you come, my brethren, out of your prisons with your faces
shining, having your minds seasoned, and your tongues tipt with
holiness! May your mouths be as a well of life, from whence may
flow the holy streams of edifying discourse! May you ever remember,
as you are sitting in your houses, going by the way, lying down,
rising up, what the Lord doth then require of you.
 2. To improve their present retirements from the world, for the
settling their spiritual estates. ’Tis a common complaint amongst
Christians, that they want assurance. Oh, if any of you that wanted
assurance when you came to prison, may carry that blessing out,
what happy gainers would you be? Now you are called more than
ever to self-searching. Now bring your graces to the touchstone. Be
much in self observation. Rest not in probable hopes. Think not that
it is enough that you can say, you hope ’tis well. Be restless till you
can say, that you know ’tis well; that you know you are passed from
death to life.

*Think not that this is a privilege that only a few may expect.
Observe but these three things:

1. To take heed of laying the marks of salvation either too high or

too low;

2. To be much in observing the frame, and bent, and workings of

your own heart:

3. To be universally conscientious, and to be constant in even

and close walking, and then I doubt not but you will have a settled
assurance, and know and feel that peace of God that passeth all
understanding.

I wish your prison may be a paradise of peace, and a Patmos of

divine discoveries, Lord Jesus set to thy Amen. I am, Sir,

Your unworthy brother and companion in the kingdom and

patience of Jesus,

JOS. ALLEINE.

January 10, 1664.

 L E T T E R XXX.
To the most beloved people, the servants of God in
Taunton, salvation.

Most dearly beloved and longed for,

my joy and crown.

M Y heart’s desire and prayer for you is, that you may be saved.
This is that which I have been praying and studying, and
preaching for these many years: and this is the end of my suffering,
and writing at this present time. I seek not other gifts, give me your
hearts, let me but part between your sins and you: suffer me but to
save you; give me leave to carry you over to Jesus Christ, and I will
not ask you any more. I will serve you gladly, I will suffer for you
thankfully, so I may but save you. Do not wonder why I follow you so
pressingly, why I call upon you so frequently; let not my importunity
be grievous to you, all this is but to save you. Christ did not think his
blood, and shall I think my breath too dear in order to your salvation;
what pity is it, that any of you should miscarry at last, under the
power of ignorance, or by a profane negligence, or a formal and
lifeless profession of strict godliness?
 Beloved, I am afraid of you, lest (as to many of you) I have run in
vain. I cannot but thankfully acknowledge, that there are not a few of
you who are the joy of your ministers, and the glory of Christ. But it
cannot be dissembled, that far the greater number give little ground
to hope, they are in the state of salvation. And must not this be a
pinching thought to a compassionate teacher, that he cannot
persuade men, but that the most of them will wilfully throw away
themselves? Is it not a woeful sight, to behold the devils driving a
great part of our miserable flocks, (as they did once the herd of
swine) violently down the hill, till they be choaked in the water,
drowned in the gulph of endless perdition? Ah miserable spectacle!
What through the wilful blindness of some, the looseness and
sensuality of others, the halving, and cold, and customary religion of
others, how great a number of our poor flocks, is Satan like to carry
utterly away from us, after all that hath been done to save him?