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Zoology Tenth Edition
Stephen A. Miller
College of the Ozarks—Professor Emeritus
zoology.miller@gmail.com
John P. Harley
Eastern Kentucky University—Professor Emeritus
zoology.harley@gmail.com.
ZOOLOGY, TENTH EDITION
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Library of Congress Cataloging-in-Publication Data
Miller, Stephen A.
Zoology / Stephen A. Miller, College of the Ozarks, John P. Harley, Eastern
Kentucky University.—Tenth edition.
pages cm
ISBN 978-0-07-783727-3 (alk. paper)
1. Zoology. I. Harley, John P. II. Title.
QL47.2.M55 2016
590—dc23 2015008555
The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website
does not indicate an endorsement by the authors or McGraw-Hill Education, and McGraw-Hill Education does
not guarantee the accuracy of the information presented at these sites.
mheducation.com/highered
B R I E F C O N T E N T S
Preface ix 16 Ambulacraria: Echinoderms and Hemichordates 301

17 Chordata: Urochordata and Cephalochordata 320
18 The Fishes: Vertebrate Success in Water 330
1 Zoology: An Evolutionary and Ecological Perspective 1
19 Amphibians: The First Terrestrial Vertebrates 352
2 Cells, Tissues, Organs, and Organ Systems of
Animals 11 20 Reptiles: Diapsid Amniotes 371
3 Cell Division and Inheritance 36 21 Birds: Reptiles by Another Name 389
4 Evolution: History and Evidence 59 22 Mammals: Synapsid Amniotes 409
5 Evolution and Gene Frequencies 78 23 Protection, Support, and Movement 435
6 Ecology: Preserving the Animal Kingdom 93 24 Communication I: Nervous and Sensory Systems 455
7 Animal Taxonomy, Phylogeny, and Organization 112 25 Communication II: The Endocrine System
and Chemical Messengers 485
8 Animal-Like Protists: The Protozoa 129
26 Circulation and Gas Exchange 506
9 Multicellular and Tissue Levels of Organization 148
27 Nutrition and Digestion 529
10 The Smaller Lophotrochozoan Phyla 172
28 Temperature and Body Fluid Regulation 553
11 Molluscan Success 197
29 Reproduction and Development 576
12 Annelida: The Metameric Body Form 220
13 The Smaller Ecdysozoan Phyla 241 Glossary Online
14 The Arthropods: Blueprint for Success 255 Credits 598
15 The Pancrustacea: Crustacea and Hexapoda 273 Index 601
iii
C O N T E N T S
Preface ix CHAPTER 4
EVOLUTION: HISTORY AND EVIDENCE 59
CHAPTER 1 Chapter Outline 59
Pre-Darwinian Theories of Change 59
ZOOLOGY: AN EVOLUTIONARY Darwin’s Early Years and His Journey 60
AND ECOLOGICAL PERSPECTIVE 1 Early Development of Darwin’s Ideas
Chapter Outline 1 of Evolution 61
Zoology: An Evolutionary Perspective 2 The Theory of Evolution by Natural Selection 63
Zoology: An Ecological Perspective 5 Microevolution, Macroevolution, and Evidence
WILDLIFE ALERT 8 of Macroevolutionary Change 66
Summary 9 Summary 76
Concept Review Questions 9 Concept Review Questions 77
Analysis and Application Questions 10 Analysis and Application Questions 77
CHAPTER 2 CHAPTER 5
CELLS, TISSUES, ORGANS, AND ORGAN
SYSTEMS OF ANIMALS 11 EVOLUTION AND GENE FREQUENCIES 78
Chapter Outline 78
Chapter Outline 11
Populations and Gene Pools 78
What Are Cells? 11
Must Evolution Happen? 79
Why Are Most Cells Small? 13
Evolutionary Mechanisms 80
Cell Membranes 13
Species and Speciation 86
Movement Across Membranes 15
Rates of Evolution 89
Cytoplasm, Organelles, and Cellular Components 19
Molecular Evolution 90
The Nucleus: Information Center 25
Mosaic Evolution 91
Levels of Organization in Various Animals 26
Summary 91
Tissues 26
Concept Review Questions 92
Organs 31
Analysis and Application Questions 92
Organ Systems 32
Summary 34
Concept Review Questions 34 CHAPTER 6
ECOLOGY: PRESERVING
THE ANIMAL KINGDOM 93
CHAPTER 3
Chapter Outline 93
CELL DIVISION AND INHERITANCE 36 Animals and Their Abiotic Environment 93
Chapter Outline 36 Biotic Factors: Populations 95
Eukaryotic Chromosomes 37 Biotic Factors: Interspecific Interactions 97
The Cell Cycle and Mitotic Cell Division 39 Communities 99
Meiosis: The Basis of Sexual Reproduction 41 Trophic Structure of Ecosystems 101
DNA: The Genetic Material 43 Cycling within Ecosystems 103
Inheritance Patterns in Animals 50 Ecological Problems 105
WILDLIFE ALERT 55 WILDLIFE ALERT 109
iv
Contents v
CHAPTER 7 CHAPTER 11
ANIMAL TAXONOMY, PHYLOGENY, MOLLUSCAN SUCCESS 197
AND ORGANIZATION 112 Chapter Outline 197
Chapter Outline 112 Evolutionary Perspective 197
Taxonomy and Phylogeny 112 Molluscan Characteristics 198
Patterns of Organization 120 Class Gastropoda 200
Higher Animal Taxonomy 124 Class Bivalvia 204
Summary 128 Class Cephalopoda 208
Concept Review Questions 128 Class Polyplacophora 213
Analysis and Application Questions 128 Class Scaphopoda 214
Class Monoplacophora 215
Class Solenogastres 215
CHAPTER 8
Class Caudofoveata 215
ANIMAL-LIKE PROTISTS: THE PROTOZOA 129 Further Phylogenetic Considerations 216
Chapter Outline 129 WILDLIFE ALERT 217
Evolutionary Perspective of the Protists 129 Summary 218
Life within a Single Plasma Membrane 131 Concept Review Questions 219
Symbiotic Lifestyles 133 Analysis and Application Questions 219
Protists and Protozoan Taxonomy 133
Further Phylogenetic Considerations 143 CHAPTER 12
Summary 146
Concept Review Questions 147 ANNELIDA: THE METAMERIC BODY FORM 220
Analysis and Application Questions 147 Chapter Outline 220
Evolutionary Perspective 220
Annelid Structure and Function 223
CHAPTER 9 Clade (Class) Errantia 228
MULTICELLULAR AND TISSUE LEVELS Clade (Class) Sedentaria 230
OF ORGANIZATION 148 Basal Annelid Groups 236
Chapter Outline 148 Further Phylogenetic Considerations 237
Evolutionary Perspective 148 Summary 239
Phylum Porifera 151 Concept Review Questions 240
Phylum Cnidaria 156 Analysis and Application Questions 240
Phylum Ctenophora 165
WILDLIFE ALERT 167 CHAPTER 13
Further Phylogenetic Considerations 168
Summary 170 THE SMALLER ECDYSOZOAN PHYLA 241
Concept Review Questions 171 Chapter Outline 241
Analysis and Application Questions 171 Evolutionary Perspective 241
Phylum Nematoda (Roundworms) 242
Other Ecdysozoan Phyla 249
CHAPTER 10
THE SMALLER LOPHOTROCHOZOAN PHYLA 172 Summary 253
Chapter Outline 172 Concept Review Questions 253
Evolutionary Perspective 172 Analysis and Application Questions 254
Platyzoa: Phylum Platyhelminthes 174
Platyzoa: Smaller Phyla 185 CHAPTER 14
Other Lophotrochozoans 190
Further Phylogenetic Considerations 194 THE ARTHROPODS: BLUEPRINT FOR SUCCESS 255
Summary 195 Chapter Outline 255
Concept Review Questions 196 Evolutionary Perspective 255
Analysis and Application Questions 196 Metamerism and Tagmatization 256
vi Contents
The Exoskeleton 257 CHAPTER 18

The Hemocoel 259
Metamorphosis 260 THE FISHES: VERTEBRATE SUCCESS IN WATER 330
Subphylum Trilobitomorpha 260 Chapter Outline 330
Subphylum Chelicerata 261 Evolutionary Perspective 330
Subphylum Myriapoda 269 Survey of Fishes 333
Further Phylogenetic Considerations 271 Evolutionary Pressures 340
Summary 271 WILDLIFE ALERT 347
Concept Review Questions 272 Further Phylogenetic Considerations 348
Analysis and Application Questions 272 Summary 351
CHAPTER 15
THE PANCRUSTACEA: CHAPTER 19
CRUSTACEA AND HEXAPODA 273
AMPHIBIANS: THE FIRST
Chapter Outline 273
TERRESTRIAL VERTEBRATES 352
Subphylum Crustacea 274 Chapter Outline 352
WILDLIFE ALERT 282 Evolutionary Perspective 352
Subphylum Hexapoda 284 Survey of Amphibians 354
Further Phylogenetic Considerations 296 Evolutionary Pressures 357
WILDLIFE ALERT 367
Summary 299
Concept Review Questions 300 Amphibians in Peril 367
Analysis and Application Questions 300 Further Phylogenetic Considerations 368
Summary 369
CHAPTER 16 Analysis and Application Questions 370
AMBULACRARIA: ECHINODERMS AND
HEMICHORDATES 301 CHAPTER 20
Chapter Outline 301 REPTILES: DIAPSID AMNIOTES 371
Chapter Outline 371
Phylum Echinodermata 302
Phylum Hemichordata 312
Survey of the Reptiles 374
WILDLIFE ALERT 313
WILDLIFE ALERT 378
Evolutionary Pressures 379
Summary 318
Summary 387
CHAPTER 17
CHORDATA: UROCHORDATA AND CHAPTER 21
CEPHALOCHORDATA 320 BIRDS: REPTILES BY ANOTHER NAME 389
Chapter Outline 320 Chapter Outline 389
Evolutionary Perspective 320 Evolutionary Perspective 389
Phylum Chordata 321 Evolutionary Pressures 392
Further Phylogenetic Considerations 326 WILDLIFE ALERT 406
Contents vii
CHAPTER 22 Endocrine Systems of Vertebrates Other Than

Birds or Mammals 493
MAMMALS: SYNAPSID AMNIOTES 409 Endocrine Systems of Birds and Mammals 495
Chapter Outline 409 Some Hormones Are Not Produced by Endocrine
Evolutionary Perspective 409 Glands 503
Diversity of Mammals 411 Evolution of Endocrine Systems 503
Evolutionary Pressures 414 Summary 504
WILDLIFE ALERT 425 Concept Review Questions 504
Human Evolution 427 Analysis and Application Questions 505
Summary 433
Concept Review Questions 434 CHAPTER 26
CIRCULATION AND GAS EXCHANGE 506
CHAPTER 23 Chapter Outline 506
Internal Transport and Circulatory Systems 506
PROTECTION, SUPPORT, AND MOVEMENT 435 Transport Systems in Invertebrates 506
Chapter Outline 435 Transport Systems in Vertebrates 509
Protection: Integumentary Systems 435 The Hearts and Circulatory Systems of Bony Fishes,
Movement and Support: Skeletal Systems 440 Amphibians, and Reptiles 512
Movement: Nonmuscular Movement The Hearts and Circulatory Systems of Birds,
and Muscular Systems 445 Crocodilians, and Mammals 514
Summary 453 The Lymphatic System Is an Open,
Concept Review Questions 454 One-Way System 516
Analysis and Application Questions 454 Gas Exchange 516
Vertebrate Respiratory Systems 518
CHAPTER 24 Human Respiratory System 523
Evolution of Respiratory Pigments 526
COMMUNICATION I: NERVOUS Summary 527
AND SENSORY SYSTEMS 455 Concept Review Questions 527
Chapter Outline 455 Analysis and Application Questions 528
Neurons: The Basic Functional Units
of the Nervous System 455 CHAPTER 27
Neuron Communication 457
Invertebrate Nervous Systems 461 NUTRITION AND DIGESTION 529
Vertebrate Nervous Systems 462 Chapter Outline 529
Sensory Reception 467 Evolution of Nutrition 529
Invertebrate Sensory Receptors 469 The Metabolic Fates of Nutrients in Heterotrophs 530
Vertebrate Sensory Receptors 473 Digestion 533
Summary 482 Animal Strategies for Getting and Using Food 534
Concept Review Questions 483 Diversity in Digestive Structures: Invertebrates 537
Analysis and Application Questions 483 Diversity in Digestive Structures: Vertebrates 539
The Mammalian Digestive System 544
CHAPTER 25 Summary 551
COMMUNICATION II: THE ENDOCRINE Analysis and Application Questions 552
SYSTEM AND CHEMICAL MESSENGERS 485
Chapter Outline 485 CHAPTER 28
Chemical Messengers 485
Hormones and Their Feedback Systems 487 TEMPERATURE AND BODY FLUID
Mechanisms of Hormone Action 488 REGULATION 553
Some Hormones of Invertebrates 489 Chapter Outline 553
An Overview of the Vertebrate Endocrine System 492 Homeostasis and Temperature Regulation 553
viii Contents
Control of Water and Solutes (Osmoregulation Sexual Reproduction in Vertebrates 580

and Excretion) 562 Examples of Reproduction among Various
Invertebrate Excretory Systems 562 Vertebrate Classes 581
Vertebrate Excretory Systems 565 The Human Male Reproductive System 584
Summary 574 The Human Female Reproductive System 587
Concept Review Questions 574 Prenatal Development and Birth in a Human 591
Analysis and Application Questions 575 Summary 596
CHAPTER 29 Analysis and Application Questions 597
REPRODUCTION AND DEVELOPMENT 576 Glossary Online

Chapter Outline 576 Credits 598
Asexual Reproduction in Invertebrates 576 Index 601
Sexual Reproduction in Invertebrates 579
P R E F A C E
Beginning with the first outlines in 1986, we have envisioned been updated with new population statistics, examples, illus-
Zoology as a general zoology textbook for use in one-semester trations, and photographs.
courses. Our plan was that Zoology should be adaptable to a Major content changes in the 10th edition of Zoology
variety of course organizations; that it should be filled with reflect the changes in our understanding of animal phylogeny
relevant, up-to-date zoological information; and that it should that have come to light in the past few years. These changes
not overwhelm introductory-level students with unnecessary should not surprise anyone given the dynamic and vital state
terminology. As teachers with over 80 years of combined of affairs in modern animal phylogenetics. A comparison
experience in college and university classrooms and laborato- of the expanded table of contents from the 9th edition to
ries, we know that a book is good only if it is read. Feedback that of the 10th edition will highlight some of the changes
from reviewers, professors, teachers, and students tells us that in chapters 7 through 22. Most noticeably, chapters 10 and
Zoology’s informative and friendly writing does encourage its 13 have been completely reorganized. Chapter 10 is now
use by students in ways that other textbooks do not. devoted to the smaller lophotrochozoan phyla, and chapter
We are honored that this book has had a part in the 13 is devoted to the ecdysozoans other than the Panarthrop-
development of students we will never know personally. We oda. Chapter 12 is reorganized to reflect new interpretations
recognize that our part in the training of future zoologists of phylogeny within the annelids. Chapters 14 and 15 are
and biologists is modest. A general zoology course is as good reorganized in recognition of the validity of the Pancrustacea.
as the professors and teachers who inspire their students Chapters 16 and 17 are reorganized to group echinoderms
to delve into a book’s pages. Over the life of Zoology we and hemichordates into a single ambulacrarian chapter.
have been a team of two authors, numerous editors, zoology In addition to organizational changes, chapters 7 through
teachers and professors, and students who have contributed 22 contain many new photographs, newly drawn cladograms,
to keeping this textbook alive and lively into its 10th edition. revised illustrations, and content additions and revisions. For
In preparing for the 10th edition of Zoology, we have taken example, we have added more information on vertebrate
seriously the feedback we have received. Every chapter has teeth in chapters 18 through 22, new material on the reptili-
been carefully scrutinized, and many of the changes incor- omorphs and the basal tetrapod/reptile transition in chapter
porated into the revisions summarized later in this preface 19, and new information on human evolution in chapter 22.
are the result of reviewer, instructor, and student comments. In previous editions, some small phyla were omitted
Preparation for the 11th edition begins now, and we wel- from the survey chapters to keep the size of the book man-
come your comments. We can be reached at the following ageable. We have found a way to introduce descriptions and
email addresses: Stephen Miller at zoology.miller@gmail.com phylogenetic relationships of these “lesser-known phyla”
and John Harley at zoology.harley@gmail.com. without adversely affecting the book’s length. These phyla
Every edition of Zoology brings something new and are presented in tables at the end of chapters 9, 12, and 16,
exciting from McGraw-Hill. As authors and teachers, we are and we hope that these tables will help students understand
excited about the LearnSmart and SmartBook adaptive more of the amazing diversity within the animal kingdom.
learning features that are available with this edition. Learn- Chapters 23 through 29 retain their clear, concise, system-
Smart and SmartBook allow students to progress through by-system coverage of animal organ systems. These chap-
chapters with greater confidence knowing that they under- ters contain new photographs and text revisions that illustrate
stand concepts being studied. We have found these tools comparative aspects of animal structure and function. For
user friendly, and we encourage you to take time to investi- example, chapter 25 describes insulin production within
gate how they can enhance student learning in your course. bivalve intestines, and chapter 26 has expanded coverage of
More information is presented on LearnSmart and SmartBook the evolution of the sinus venosus and the SA node.
in “Teaching and Learning Resources.”
CONTENT AND ORGANIZATION PEDAGOGY

We have maintained from the inception of this text that Integrated Learning Outcomes
evolutionary and ecological perspectives captivate students. and Critical Thinking
These perspectives are fundamental to students understand-
ing the unifying principles of zoology. Chapters 1 through 6 We have retained pedagogical elements useful to science fac-
present cellular, evolutionary, and ecological concepts that ulty in identifying measurable learning outcomes. Learning
unite zoology to biology as a whole. These chapters have Outcomes and Section Reviews, including section review
ix
x Preface
questions, have been retained in the 10th edition for each species statistics have been updated. Ecological problems are
major section of each chapter. Answers for section review discussed including an assessment of eight critical environ-
questions are available to instructors on the Zoology website. mental processes: biodiversity loss, nitrogen cycling, phos-
These elements allow students to self-test and instructors to phorus cycling, climate change, ocean acidification, land and
document student learning. In addition, instructors and stu- freshwater use, and ozone depletion. The ecological per-
dents using Connect Zoology can access auto-gradable and spective is reinforced by boxed readings entitled Wildlife
interactive assessment material tied to learning outcomes from Alerts. Wildlife Alerts first appeared in the 4th edition and
the text. These Connect features include the new LearnSmart have been very well received by students and professors.
and SmartBook adaptive learning tools and are described Each boxed reading depicts the plight of selected animal
under “Teaching and Learning Resources.” species or broader ecosystem issues relating to preserv-
Each chapter ends with a set of Concept Review ing animal species. These readings have been revised, and
Questions and Analysis and Application Questions. some new readings have been added. Chapter 6 has a new
These questions have been carefully reviewed and revised reading on species translocation as a tool in conservation
as needed. They allow students to test their understanding of biology. Chapter 18 has a new reading on the problem of
chapter concepts and to apply concepts they have learned in invasive species. Chapter 20 has a new reading on the plight
each chapter. Suggested answers to these questions are avail- of the Eastern diamondback rattlesnake (Crotalus adaman-
able to instructors through Connect. The glossary has been teus). Students who read and study this book should have
moved to the Connect site as well. In the mobile information an enhanced understanding of ecological principles and how
age, it seems students are quicker to check a definition elec- human ignorance and misplaced values have had detrimental
tronically than to flip to the back of the book. The glossary effects on our environment in general and on specific animal
will also be available in SmartBook. groups in particular.
An Evolutionary The Process of Science

and Ecological Focus To help students understand that science is a process, not
just a body of facts, How Do We Know boxed readings are
Zoology emphasizes ecological and evolutionary concepts and
retained in this edition and they highlight research results
helps students understand the process of science through ele-
that provide insight into biological processes. Chapter 9 has
ments of chapter organization and boxed readings. Each chapter
a boxed reading entitled “How Do We Know about Sponge
in chapters 8 through 22 begins with a section entitled Evolu-
Defenses?” This reading describes how zoologists investi-
tionary Perspective. This section discusses the relationship
gated sponge defense mechanisms. Chapter 19 has a boxed
of the phylum or phyla covered in the current chapter to the
reading entitled “How Do We Know about Amphibian Skin
animal kingdom as a whole and to animals discussed in previ-
Toxins?” This reading describes how scientists are studying
ous chapters. Students are frequently reminded to consult the
antibacterial and anticancer effects of amphibian skin toxins.
animal kingdom cladogram on the inside front cover and the
Students learn that these studies have implications for study-
geological time chart on the inside back cover. Similarly, each
ing naturally occurring compounds that may aid in the devel-
survey chapter ends with a section entitled Further Phylo-
opment of novel pharmaceutical drugs.
genetic Considerations. This section discusses phylogenetic
relationships of groups (subphyla or classes) within the phylum
or phyla being studied and is a point of transition between Digital Assets and Media Integration
chapters. The discussion in this section is usually supported by
a cladogram illustrating important phylogenetic relationships. Beginning with the 9th edition of Zoology, digital resources
To further explain and support evolutionary con- were integrated into the book through the Connect Zoology
cepts, a set of themed boxed readings entitled Evolution- site. Many of the sections within most chapters are linked
ary Insights is present throughout the book. These boxes to animations of biological processes and to MP3 files. This
provide detailed examples of principles covered in a chapter media integration is indicated within the printed text by the
and provide insight into how evolutionary biology works. For icons shown below. These media assets are available through
example, chapter 4 includes a reading on big-cat biogeogra- Connect.
phy that illustrates how a variety of sources of evidence are MP3 files. These short three-to-five minute audio
used to paint a picture of the history of one group of animals. files serve as a review of material in certain sections of
Chapter 5 has a reading on speciation of Darwin’s finches the book and help students with the pronunciation of
that illustrates how speciation can occur. Chapter 18 has a scientific terms and processes.
reading on the evolution of the vertebrate limb, and chapter Animations. The authors have selected animations
25 has a reading on the evolution of hormone receptors. from McGraw-Hill’s library of animations that will
The ecological perspective of Zoology is stressed through- enhance students’ understanding of the material
out chapters 1 to 22. Human population and endangered within the chapter.
Preface xi
NEW TO THE TENTH EDITION larger genetic drift concept. Cichlid fish are used as an
additional example of rapid evolutionary change in “Rates
As with earlier revisions of Zoology, the focus for this revision of Evolution.”
has been on presenting evolutionary and ecological concepts
• Chapter 6 (Ecology: Preserving the Animal
clearly and accurately using examples from current literature
Kingdom)
as convincingly as possible. The revisions highlighted below
should impress students with the excitement experienced in The discussion of density-dependent factors influencing
zoology as new information clarifies zoological concepts and populations has been expanded. The discussion of cryp-
informs our understanding of phylogenetic relationships. sis has been expanded. New examples illustrate chemi-
• Chapter 1 (Zoology: An Evolutionary and Ecological cal, auditory, and visual crypsis. The section “Ecological
Perspective) Problems” has been revised. It has been updated with
Table 1.1 has been updated with the addition of com- population statistics from 2014 and new statistics on rates
parative genomics and bioinformatics as a specialization of population growth. Problems associated with the aging
in zoology. The use of cichlid fish as an example of the of the human population are now included. The human
importance of evolutionary and ecological perspectives age pyramids in figure 6.13 have been revised to sup-
within zoology has been expanded. The concept of port this updated discussion. A new “Wildlife Alert” that
evolutionary plasticity is introduced. Population, world discusses species translocation as a tool in ecosystem
resource, rainforest depletion, and threatened and endan- restoration has been added. It points out the usefulness
gered species statistics have been updated with figures of species introductions and reintroductions as well as the
from 2014. Table 1.5 is new and compares human popu- risks associated with introducing nonnative species into
lation projections in major world regions. ecosystems (see the new “Wildlife Alert” in chapter 20).
• Chapter 2 (Cells, Tissues, and Organ Systems of • Chapter 7 (Animal Taxonomy, Phylogeny, and
Animals) Organization)
This chapter, including table 2.3, has been updated to The discussion in the section “Animal Systematics” is
include discussion of a newly discovered organelle, the expanded. It now includes a comparison of the concepts
exosome. New information is presented on hydrogen per- of homology and homoplasy. The discussion of phyloge-
oxide as a metabolite that induces oxidative damage and netic systematics (cladistics) has been revised. The terms
mediates aging. “plesiomorphies” and “apomorphies” are discussed. The
hypothetical cladogram (formerly figure 7.5) used to illus-
• Chapter 3 (Cell Division and Inheritance) trate cladistic principles has been replaced by a simplified
Coverage of the cell cycle has been expanded, includ- vertebrate cladogram (now figure 7.4). The new figure
ing the discussion of the G0 phase. Figure 3.3 is replaced depicts familiar character states that are used to support
to accompany this expanded discussion. The discus- the discussion of cladistics. After studying figure 7.4,
sion of mitotic cell division now includes a discussion of students can “graduate to” figure 7.5—a more detailed
prometaphase, and figure 3.5 has been revised to more version of vertebrate phylogeny. The discussion of evolu-
clearly illustrate the concepts of mitotic cell division. tionary systematics is also expanded, including the “adap-
Figure 3.6 has been revised to include an illustration of tive zone” concept. The phylogenetic species concept is
crossing-over in meiosis. Figure 3.15 has been redrawn to discussed in more detail. In “Higher Animal Taxonomy,”
clearly illustrate primary and secondary nondisjunction. figure 7.12 has been redrawn and is an abbreviated
A new “How Do We Know” box on Thoroughbred horse version of the larger, highly revised cladogram on the
inbreeding illustrates the dangers of reducing genetic inside front cover of the textbook. Figure 7.12 (and the
diversity through inbreeding. The “Wildlife Alert” on pre- expanded cladogram) reflects the taxonomic revisions
serving genetic diversity provides new information on the that will be described in chapters 8 through 22.
endangered status of the cheetah (Panthera uncia). • Chapter 8 (Animal-Like Protists: The Protozoa)
• Chapter 4 (Evolution: History and Evidence)
Figure 8.1 has been replaced with a new cladogram
New information is presented on the evolution of the showing the phylogeny of six protist supergroups.
horse, and figure 4.10 has been revised to support this
• Chapter 9 (Multicellular and Tissue Levels of
coverage.
Organization)
• Chapter 5 (Evolution and Gene Frequencies)
Chapter 9 opens with a revised discussion of the origin of
The discussion of genetic drift has been revised and now multicellularity, including selective advantages of multicel-
includes the concept of fixation of an allele. “Founder lularity and requirements for the evolution of multicellu-
Effect” and “Bottleneck Effect” are organized into sub- larity. Colonial and coenocytial hypotheses are discussed.
headings to more clearly define their relationship to the Figure 9.1 has been revised to reflect updated animal
xii Preface
phylogeny. “Animal Origins” has additional detail on the clade Cycloneuralia. The relationships of these phyla
animal/choanocyte relationships. “Further Phylogenetic to the Panarthropoda are described in a revised “Further
Relationships” presents new evidence that suggests that Phylogenetic Considerations” and presented in a clado-
the Ctenophora, not the Porifera, is a sister taxon to all gram in figure 13.16.
other animals. Table 9.4 is new and features two lesser-
• Chapter 14 (The Arthropods: Blueprint for Success)
known basal animal phyla: Placozoa and Acoelomorpha.
Chapter 14 has received major organizational revisions
• Chapter 10 (The Smaller Lophotrochozoan Phyla)
that reflect arthropod phylogeny. Coverage of the Crusta-
Chapter 10 has received major revisions and now describes cea has been moved to chapter 15, and coverage of the
lophotrochozoan phyla other than Mollusca (chapter 11) Myriapoda has been moved to chapter 14.
and Annelida (chapter 12). The “Evolutionary Perspective”
• Chapter 15 (Pancrustacea: Crustacea and Hexapoda)
has been rewritten to explain why the new chapter orga-
nization makes phylogenetic sense, and it also describes Chapter 15 is devoted to the clade Pancrustacea. Discus-
the lophophore and the trochophore larval stage—the two sion of the clade Panarthropoda is described in “Further
features that unite the lophotrochozoans. Members of the Phylogenetic Considerations” and includes brief descrip-
clade Platyzoa (Platyhelminthes, Gastrotricha, Microgna- tions of Tardigrada and Onychophora. The discussion of
thozoa, Gnathostomulida, Rotifera, and Acanthocephala) arthropod phylogeny includes new information support-
are described first. They are followed by Cycliophora, ing the validity of the mandulate and chelicerate lineages.
Nemertea, Ectoprocta, and Brachiopoda. Three of these It also presents new information that suggests that the
phyla have not been featured in previous editions of this traditional subphylum Crustacea is paraphyletic. Hexa-
textbook. “Further Phylogenetic Considerations” has been poda is presented as a monophyletic lineage within the
rewritten to focus on lophotrochozoan relationships. The crustacean phylogeny.
questionable validity of the clade Platyzoa and the para-
• Chapter 16 (Amulacraria: Echinoderms
phyly of Turbellaria are discussed. Figure 10.29 is a new
and Hemichordates)
cladogram depicting lophotrochozoan relationships.
Chapter 16 has received organizational revisions that
• Chapter 11 (Molluscan Success)
reflect our current understanding of deuterostome
New information on bivalve burrowing and cephalopod phylogeny. The discussion of the hemichordates has been
sensory perception is provided. moved from chapter 17 to reflect their closer ties to the
Echinodermata. The “Evolutionary Perspective” has been
• Chapter 12 (Annelida: The Metameric Body Form)
revised to include more information on the clade Ambu-
Chapter 12 has received extensive revision that reflects lacraria and deuterostome evolution in general. “Further
recent changes in our understanding of the phylogenetic Phylogenetic Considerations” has been revised to include
relationships within the Annelida. The “Evolutionary Per- discussion of the growing body of evidence of the ances-
spective” describes the traditional class “Polychaeta” as tral status of pharyngeal slits in the deuterostome lineage.
paraphyletic, and it explains the reinstatement of “Erran- The cladogram in figure 16.19 has been revised to support
tia” and “Sedentaria” as two major clades within Annelida. the discussion of deuterostome phylogeny. Table 16.2
An updated discussion of annelid structure and function is a new table that provides information on two lesser-
is then followed by descriptions of the clades Errantia and known phyla. The Chaetognatha and Xenoturbellida are
Sedentaria. Nereis and Glycera are used as representative described as “Phyla of Uncertain Affinities.”
errantians. Various tubeworms, siboglinids, echiurians,
• Chapter 17 (Chordata: Urochordata
and clitellates are described as representative sedentar-
and Cephalochordata)
ians. Chaetopteridae and Sipuncula are described as basal
annelid groups. The reinterpretation of annelid phylogeny Chapter 17 has received minor revisions apart from mov-
is described in a revised “Further Phylogenetic Consider- ing the Hemichordata into chapter 16. The recognition
ations” and shown in a revised cladogram in figure 12.24. that pharyngeal slits arose early in deuterostome evolu-
A new table 12.2 presents descriptions and phylogenetic tion means that these structures are not unique to the
relationships of three lesser-known lophotrochozoan chordates, but they are adapted for important functions
phyla: Entoprocta, Phoronida, and Mesozoa. in most chordates. “Further Phylogenetic Considerations”
presents a revised discussion of the relationships between
• Chapter 13 (The Smaller Ecdysozoan Phyla)
the chordate subphyla. The cladogram in figure 17.10 has
Chapter 13 has received major organizational revisions. been revised to support this discussion.
It covers the ecdysozoan phyla other than Arthropoda,
• Chapter 18 (The Fishes: Vertebrate Success in Water)
Onycophora, and Tardigrada. The five phyla discussed
in chapter 13 (Nematoda, Nematomorpha, Kinorhyncha, Chapter 18 has received minor revisions. It includes a
Priapulida, and Loricifera) are described as members of new boxed reading “Wildlife Alert: Invasive Species—A
Preface xiii
Growing Problem in a Shrinking World.” This reading points out that different hominin species were contem-
uses the red lionfish (Pterois volitans) as an example to poraries of one another and may have interacted. Table
alert students to the risks associated with accidental or 22.3 (Significant Events in Hominin Evolution) and Figure
intentional release of species into nonnative ecosystems. 22.20 (Human Evolution) have been updated to support
the revised discussion of human evolution.
• Chapter 19 (Amphibians: The First Terrestrial
Vertebrates) • Chapter 25 (Communication II: The Endocrine
System and Chemical Messengers)
New information is presented on amphibian phylogeny in
the “Evolutionary Perspective.” “Evolutionary Pressures” A short discussion has been added on the possible role of
contains expanded coverage of amphibian teeth, heart insulin in carbohydrate regulation in bivalves. Table 25.1
structure, and heart function. “Further Phylogenetic Con- (Some Major Endocrine Tissues and Hormones) now
siderations” has been expanded to include discussion of lists additional hormones and their principal functions:
the reptiliomorph lineage and evolution of the synapsid peptide YY3-36, adiponectin, irisin, and ghrelin. The “Evo-
lineage from ancient tetrapods. This discussion is sup- lutionary Insights” box has been expanded to include
ported by the revised cladogram in figure 19.3 and a pho- discussion of the evolutionary conservation of hormonal
tograph of a diadectomorph fossil in figure 19.19. control of parental behavior and the effects of the resul-
tant parental behavior on infant development.
• Chapter 20 (Reptiles: Diapsid Amniotes )
The organization of chapter 20 better reflects diapsid
phylogeny. The evolutionary perspective and the revised ACKNOWLEDGMENTS
cladogram in figure 20.3 complement the reptiliomorph
discussion in chapter 19. The survey of reptiles is orga- We wish to thank reviewers who provided feedback and
nized into three headings: Testudines, Archosauria, and analysis of the revision plan for the 10th edition. In the midst
Lepidosauria. While the traditional reptilian order names of their busy teaching and research schedules, they took time
are retained, the new organization reflects reptilian phy- to consider the revisions we were making to the table of
logeny and makes very clear the position of Aves within contents and offer constructive advice that greatly improved
the reptilian lineage. The birds are still covered in a sepa- the 10th edition. One person in particular has become
rate chapter 21 out of respect for zoological tradition and a friend and valued advisor for us. As the 9th edition was
in recognition of the importance of distinctive avian char- being released, we began an ongoing email dialog with Todd
acteristics. “Evolutionary Pressures” contains expanded Tupper of Northern Virginia Community College. His feed-
coverage of reptilian teeth and temperature regulation. back, and feedback and questions from his students, have
A new “Wildlife Alert: The Eastern Diamondback Rattle- been especially valuable in the development of the 10th edi-
snake (Crotalus adamanteus)” has been added to chapter tion of Zoology. His comments and photographs were par-
20. It was written by guest contributors actively working ticularly valuable in the revisions for chapters 19 and 20, and
to preserve this magnificent reptile. he should receive most of the credit for the new “Wildlife
Alert” on the Eastern diamondback rattlesnake in chapter 20.
• Chapter 21 (Birds: Reptiles by Another Name) Thank you, Todd!
New information has been added to chapter 21 on
ancient theropods and the evolution of flight. The blurred
distinction between bird and nonbird within the thero-
REVIEWERS
pod lineage is emphasized. The presentation of avian Chris Brown, Tennessee Tech University
taxonomy reflects recent genome-scale findings. In “Evo- David M. Hayes, Eastern Kentucky University
lutionary Pressures” new information has been added Jennifer Skillen, Sierra College
on the unidirectional air flow through crocodylian lungs, Todd Tupper, Northern Virginia Community College
reinforcing the archosaurian affinities of birds and croco-
dylians. The coverage of thermoregulation has been reor-
ganized for clarity of presentation. SPECIAL THANKS
• Chapter 22 (Mammals: Synapsid Amniotes) AND DEDICATIONS
“Evolutionary Pressures” has new information on mam- The publication of a textbook requires the efforts of many
malian teeth. The description of mammalian placentas people. We are grateful for the work of our colleagues at
has been clarified. The presentation of human evolution McGraw-Hill Education who have shown extraordinary
has been updated to reflect our current understanding patience, skill, and commitment to this textbook. Rebecca
of the very bush-like hominin phylogeny. The coverage Olson, our Brand Manager, has helped shape Zoology through
emphasizes that adaptations for bipedal locomotion prob- its recent editions and has skillfully managed Zoology’s tran-
ably occurred more than once within our lineage. It also sition into the interactive electronic world. Her wisdom and
xiv Preface
skill are evident in the 10th edition. Elizabeth Sievers, Lead Most importantly, we wish to extend appreciation to
Product Developer, coordinated all of the tasks involved with our families for their patience and encouragement. Janice A.
publishing this edition. We learned to expect her emails at Miller lived through many months of planning and writing
all hours of the day, and we are still amazed at her ability of the 1st edition of Zoology. She died suddenly two months
to guide reviews, manuscript, figure and table revisions, and before it was released. Our wives, Carol A. Miller and Donna
new photographs into their proper places in the final version L. Harley, have been supportive throughout the revision pro-
you have in front of you. Thank you for your patience with cess. Carol, an accomplished musician, spent many hours
us on the many occasions that we submitted revised material proofreading Zoology for grammatical errors. Over the past
and then resubmitted the same with additional changes. We 20 years, she has become a much better zoologist than her
know that we must have caused you moments of frustra- husband has become a musician—something about practic-
tion beyond words. Lisa A. Bruflodt served as Content Project ing got in his way. We appreciate the sacrifices that our fami-
Manager for this edition. We appreciate her efficiency and lies have made during the writing and revision of this text.
organization. We dedicate this book to our families.
Preface xv
TEACHING AND LEARNING RESOURCES

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xvi Preface
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Preface xvii
Instructor’s Resources
Within Connect you will find presentation materials and other resources for your course including:
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or custom-made classroom materials.
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xviii Preface
GENERAL ZOOLOGY updates in animal taxonomy and evolution. The pedagogy

LABORATORY MANUAL includes learning outcomes and numbered section headings.
Learning outcomes are correlated with Learning Outcome
ISBN: 0-07-747929-7 Reviews and Analytical Thinking questions in worksheets.
Seventh Edition, by Stephen A. Miller, is an excellent corol- The pedagogy makes this laboratory manual more interac-
lary to the text. This laboratory manual includes photographs tive and student learning more easily assessed. A Laboratory
and illustrations, activities on the scientific method, cladistics, Resource Guide with information on materials and proce-
ecological and evolutionary principles, and animal structure dures as well as answers to worksheet questions accompany-
and function. The Seventh Edition includes major content ing the lab exercises can be found in the Zoology website.
1
Zoology:
An Evolutionary
and Ecological Perspective
Chapter Outline
1.1 Zoology: An Evolutionary Perspective
Evolutionary Processes
Animal Classification and Evolutionary
Generations of Luo fishermen on Lake Victoria, Africa have caught cichlid fish, Relationships
including tilapia, as a mainstay of their economy. Recent introductions of the 1.2 Zoology: An Ecological Perspective
Nile perch (Lates niloticus) has changed the Lake Victoria ecosystem and the fish- World Resources and Endangered
ing economy of the lake. Animals
Zoology (Gr. zoon, animal 1 logos, to study) is the study of animals. It is one of
the broadest fields in all of science because of the immense variety of animals and
the complexity of the processes occurring within animals. There are, for exam-
ple, more than 28,000 described species of bony fishes and more than 400,000
described (and many more undescribed) species of beetles! It is no wonder that
zoologists usually specialize in one or more of the subdisciplines of zoology. They
may study particular functional, structural, or ecological aspects of one or more
animal groups (table 1.1), or they may choose to specialize in a particular group
of animals (table 1.2).
Ichthyology, for example, is the study of fishes, and ichthyologists work to
understand the structure, function, ecology, and evolution of fishes. These stud-
ies have uncovered an amazing diversity of fishes. One large family of bony fish,
Cichlidae, contains 2,000 to 3,000 species. Members of this family include the
familiar Tilapia species that grace our dinner plates and a host fish that hobbyists
maintain in freshwater aquaria. Cichlid species range in length from 2.5 cm to
1 m and have an enormous variety of color patterns (figure 1.1), habitats, and body
forms. Ichthyologists have described a wide variety of feeding habits in cichlids.
These fish include algae scrapers like Eretmodus that nip algae with chisel-like
teeth; insect pickers like Tanganicodus; and scale eaters like Perissodus. All cich-
lids have two pairs of jaws. The mouth jaws are used for scraping or nipping
food, and the throat jaws are used for crushing or macerating food before it is
swallowed.
Many cichlids mouth brood their young. A female takes eggs into her mouth
after the eggs are spawned. She then inhales sperm released by the male, and
fertilization and development take place within the female’s mouth! Even after the
eggs hatch, young are taken back into the mouth of the female if danger threatens
(figure 1.2). Hundreds of variations in color pattern, body form, and behavior in
this family of fishes illustrate the remarkable diversity present in one relatively
small branch of the animal kingdom. Zoologists are working around the world to
understand and preserve this enormous diversity.
2 CHAPTER ONE
T A B L E 1 . 1
E XAMPLES OF S PECIALIZATIONS IN Z OOLOGY
SUBDISCIPLINE DESCRIPTION
Anatomy Study of the structure of entire organisms

and their parts
Cytology Study of the structure and function of cells
Comparative Study of the structure, function, and evo-
Genomics and lution of the genetic composition of
Bioinformatics groups of animals using computer-based
computational methods
Ecology Study of the interaction of organisms with
their environment
Embryology Study of the development of an animal from (a)
the fertilized egg to birth or hatching
Genetics Study of the mechanisms of transmission of
traits from parents to offspring
Histology Study of tissues
Molecular biology Study of subcellular details of structure
and function
Parasitology Study of animals that live in or on other
organisms at the expense of the host
Physiology Study of the function of organisms and
their parts
Systematics Study of the classification of, and the evolution-
ary interrelationships among, animal groups
(b)
T A B L E 1 . 2 FIGURE 1.1
E XAMPLES OF S PECIALIZATIONS IN Z OOLOGY
Cichlids. Cichlids of Africa exist in an amazing variety of color
BY T AXONOMIC C ATEGORIES
patterns, habitats, and body forms. (a) This dogtooth cichlid
(Cynotilapia afra) is native to Lake Malawi in Africa. The female of
SUBDISCIPLINE DESCRIPTION the species broods developing eggs in her mouth to protect them
from predators. (b) The fontosa (Cyphontilapia fontosa) is native to
Lake Tanganyika in Africa.
Entomology Study of insects
Herpetology Study of amphibians and reptiles
Animals share a common evolutionary past and evolutionary
Ichthyology Study of fishes forces that influenced their history. Evolutionary processes are
Mammalogy Study of mammals remarkable for their relative simplicity, yet they have had awe-
Ornithology Study of birds some effects on life-forms. These processes have resulted in
an estimated 4 to 10 million species of animals living today.
Protozoology Study of protozoa
(Over 1 million animal species have been described.) Many
more, about 90%, existed in the past and have become extinct.
Zoologists must understand evolutionary processes if they are
1.1 Z OOLOGY: A N to understand what an animal is and how it originated.
E VOLUTIONARY P ERSPECTIVE
Evolutionary Processes
LEARNING OUTCOMES Organic evolution (L. evolutus, unroll) is change in the genetic
1. Formulate a hypothesis regarding the evolutionary origin makeup of populations of organisms over time. It is the
of contrasting color patterns in two closely related spe- source of animal diversity, and it explains family relation-
cies of fish. ships within animal groups. Charles Darwin published
2. Explain how our taxonomic system is hierarchical. convincing evidence of evolution in 1859 and proposed a
Zoology: An Evolutionary and Ecological Perspective 3
Uganda
Lake Kivu Kenya

Lake Victoria
Democratic Lake Tanganyika
Republic of the
Congo Tanzania
Mozambique
Zambia
FIGURE 1.2
A Scale-Eating Cichlid. Scale-eaters (Perissodus microlepis)
attack from behind as they feed on scales of prey fish. Two body Lake Malawi
forms are maintained in the population. In one form, the mouth is Malawi
asymmetrically curved to the right and attacks the prey’s left side.
The second form has the mouth curved to the left and attacks the
prey’s right side. Both right- and left-jawed forms are maintained FIGURE 1.3
in the population and prey do not become wary of being attacked Lakes Victoria, Kivu Tanganyika, and Malawi. These lakes
from one side. Perissodus microlepis is endemic (found only in) to have cichlid populations that have been traced by zoologists
Lake Tanganyika. A male with its brood of young is shown here. to an ancestry that is approximately 200,000 years old. Cichlid
populations originated in Lake Kivu and Lake Tanganyika and then
spread to the other lakes.
mechanism that could explain evolutionary change. Since
that time, biologists have become convinced that evolution Animal Classification
occurs. The mechanism proposed by Darwin has been con-
firmed and now serves as the nucleus of our broader under-
and Evolutionary Relationships
standing of evolutionary change (see chapters 4 and 5). Evolution not only explains why animals appear and function
Understanding how the diversity of animal structure as they do, but also explains family relationships within the
and function arose is one of the many challenges faced by animal kingdom. Zoologists have worked for many years to
zoologists. For example, the cichlid scale eaters of Africa feed understand the evolutionary relationships among the 2,000 to
on the scales of other cichlids. They approach a prey cichlid 3,000 cichlid species. Groups of individuals are more closely
from behind and bite a mouthful of scales from the body. related if they share more of their genetic material (DNA)
The scales are then stacked and crushed by the second set of with each other than with individuals in other groups. (You
jaws and sent to the stomach and intestine for protein diges- are more closely related to your brother or sister than to your
tion. Michio Hori of Kyoto University found that there were cousin for the same reason. Because DNA determines most
two body forms within the species Perissodus microlepis. of your physical traits, you will more closely resemble your
One form had a mouth that was asymmetrically curved to the brother or sister.) Genetic studies suggest that the oldest pop-
right, and the other form had a mouth that was asymmetri- ulations of African cichlids are found in Lakes Tanganyika
cally curved to the left. The asymmetry results in right-jawed and Kivu, and from these the fish invaded African rivers and
fish approaching and biting scales from the left side of their Lakes Malawi, Victoria, and other smaller lakes (figure 1.3).
prey and the left-jawed fish approaching and biting scales The history of these events is beginning to be u nderstood
from the right side of their prey. Both right- and left-jawed and represents the most rapid known origin of species of any
fish have been maintained in the population; otherwise, the animal group. For example, the origin of Lake Victoria’s cich-
prey would eventually become wary of being attacked from lid species has been traced to an invasion of ancestral cich-
one side. The variety of color patterns within the species lids, probably from Lake Kivu approximately 100,000 years
Topheus duboisi has also been explained in an evolution- ago. Today, Lake Kivu has only 15 species of cichlids. This
ary context. Different color patterns arose as a result of the invasion continued up to about 40,000 years ago when vol-
isolation of populations among sheltering rock piles sepa- canic eruptions isolated the fauna of Lakes Kivu and Victoria.
rated by expanses of sandy b ottom. Breeding is more likely That time period is long from the perspective of a human
to occur within their isolated populations because fish that lifetime, but it is a blink of the eye from the perspective of
venture over the sand are exposed to predators. evolutionary time. There is firm geological evidence that Lake
4 CHAPTER ONE
How Do We Know about Genetic

Relationships among Animals?
A
s shown by the example readily measured. With the advent are related. Because proteins, like
of Lake Victorian cichlids, of molecular biological techniques, enzymes, are encoded by DNA,
zoologists often ask ques- zoologists have added to their rep- variations in the structure of a pro-
tions about genetic relationships ertoire of tools the analysis of varia- tein also reflect genetic relationships.
among groups of animals. These tion in a series of enzymes, called The genetic relationships of cich-
family relationships are depicted in allozymes, and DNA structure. lids described in this chapter were
tree diagrams throughout this book. These techniques allow zoologists investigated using a combination of
Early studies of genetic relationships to directly observe genetic relation- morphological characteristics and
involved the analysis of inherited ships because the more DNA that molecular techniques. These topics
morphological characteristics like two individuals, or groups of individu- are discussed in more detail in
jaw and fin structure that can be als, share, the more closely they chapters 3, 4, and 5.
Victoria nearly dried out and then refilled 14,700 years ago. microlepis—is recognized throughout the world by its two-
This event probably did not result in the extinction of all part species name. Verbal or written reference to a species refers
cichlids in the lake because the lake basin may have retained to an organism identified by this two-part name. The species
smaller bodies of water, and thus refuges for some cichlid epithet is generally not used without the accompanying genus
species. After Lake Victoria refilled, these refuge popula- name or its abbreviation (see chapter 7). Above the genus level,
tions provided the stock for recolonizing the lake. More than organisms are grouped into families, orders, classes, phyla,
500 species of cichlids inhabited Lake Victoria by the begin- kingdoms, and domains, based on a hierarchy of relatedness
ning of the twentieth century. Many of these species evolved (figure 1.4). Organisms in the same species are more closely
in fewer than 15,000 years. This very rapid evolution is a phe- related than organisms in the same genus, and organisms in
nomenon referred to as evolutionary plasticity (see chapter 5). the same genus are more closely related than organisms in
Like all organisms, animals are named and classified into the same family, and so on. When zoologists classify animals
a hierarchy of relatedness. Although Carl von Linne (1707–1778) into taxonomic groupings they are making hypotheses about
is primarily remembered for collecting and classifying plants, the extent to which groups of animals share DNA, even when
his system of naming—binomial nomenclature—has also they study variations in traits like jaw structure, color pat-
been adopted for animals. A two-part name describes each terns, and behavior, because these kinds of traits ultimately
kind of organism. The first part is the genus name, and the are based on the genetic material.
second part is the species epithet. Each kind of organism Evolutionary theory has affected zoology like no
(a species)—for example, the cichlid scale-eater Perissodus other single theory. It has impressed scientists with the
Species
Domain Kingdom Phylum Class Order Family Genus
Name
Muscidae Musca Musca

domestica
Arthropoda Hexapoda Diptera
Tabanidae Tabanus Tabanus

opacus
Eukarya Animalia
Osteichthyes Perciformes Cichlidae Perissodus Perissodus

microlepis
Chordata
Mammalia Primates Hominidae Homo Homo

sapiens
FIGURE 1.4
Hierarchy of Relatedness. The classification of a housefly, horsefly, cichlid fish, and human illustrates how the classification system
depicts degrees of relatedness.
fundamental unity of all of life. As the cichlids of Africa in further habitat loss. To make matters worse, when Nile
illustrate, evolutionary concepts hold the key to under- perch are caught, their excessively oily flesh must be dried.
standing why animals look and act in their unique ways, Fishermen cut local forests for the wood needed to smoke
live in their particular geographical regions and habitats, the fish. This practice has resulted in severe deforestation
and share characteristics with other related animals. around Lake Victoria. The resulting runoff of soil into the
lake has caused further degradation. Decreased water quality
not only presented problems for the survival of individual
SECTION REVIEW 1.1 cichlids, but also increased turbidity that interfered with criti-
cal behavioral functions. Many of these species rely on their
The knowledge of evolutionary processes helps zoologists
bright colors as visual cues during mating. Mouth-brooding
understand the great diversity of structure and function pres-
species rely on vision to pick up developing eggs. The loss
ent in animals. Evolution also helps zoologists understand rela-
of Lake Victorian cichlids may be the largest extinction
tionships among animals. These evolutionary relationships are
event of vertebrate species in modern human history.
ultimately based on shared DNA, they are reflected in inher-
There are some hopeful signs in this story. Although
ited morphological characteristics, and they are represented
many Lake Victorian species have been lost forever, some
by groupings in the classification system. The hierarchical
cichlids are recovering. Heavy fishing pressure on the Nile
nature of the naming system is reflected in groups becoming
perch has reduced its population density. (It still comprises
more inclusive as one moves from species to domain.
more than 50% of catch weight—down from about 90% in the
Why can taxonomists use similarities in DNA, simi- 1980s.) This decline has promoted the recovery of some cichlids
larities in morphological characteristics, or both when that feed on small animals in the upper portions of open-water
investigating taxonomic (evolutionary) relationships areas. (The Nile perch is predominately a bottom-dwelling
among animals? predator.) One cichlid (Haplochromis pyrrhocephalus) is far-
ing better than most other cichlid species. Over a 20-year
period, scientists have observed rapid evolution of increased
1.2 ZOOLOGY: AN ECOLOGICAL gill surface area and associated changes in head morphol-
ogy, which have allowed this species to survive the lowered
PERSPECTIVE oxygen concentrations now present in Lake Victoria.
The Lake Victoria example also illustrates how ecologi-
LEARNING OUTCOMES cal decisions made for economic reasons can have far-ranging
1. Explain how the failure to understand ecological rela- economic and ecological consequences. Nile perch are mar-
tionships among animals and their environment has keted to Nairobi, the Middle East, and Europe to restaurants
resulted in detrimental environmental consequences. and fish markets. The hide is used in belts and purses, and
2. Analyze the relationships between human population the urinary bladder is used in oriental soup stock and as filter
growth and threats to world resources. material by European alcohol producers. Catching, process-
ing, and marketing such large fish to diverse foreign markets
Just as important to zoology as an evolutionary perspective have resulted in the fishing and processing industries being
is an ecological perspective. Ecology (Gr. okios, house + taken from the hands of local fishermen and processors. These
logos, to study) is the study of the relationships between functions are primarily the work of large-boat fishing fleets
organisms and their environment (see chapter 6). Through- and large fish processing corporations. Changes in the local
out our history, humans have depended on animals, and economy to agriculture have resulted in deforestation of the
that dependence too often has led to exploitation. We surrounding landscapes, and untreated sewage and agricul-
depend on animals for food, medicines, and clothing. We tural and industrial runoff have further polluted Lake Victoria.
also depend on animals in other, more subtle ways. This
dependence may not be noticed until human activities upset
the delicate ecological balances that have evolved over World Resources
hundreds of thousands of years. and Endangered Animals
In the 1950s, the giant Nile perch (Lates niloticus) was
There is grave concern for the ecology of the entire world, not
introduced into Lake Victoria in an attempt to increase the
just Africa’s greatest lakes. The problems, however, are most
lake’s fishery (figure 1.5). This voracious predator reduced
acute in developing countries, which are striving to attain the
the cichlid population from 80% to less than 1% of the total
same wealth as industrialized nations. Two problems, global
fish biomass (total mass of all fish in the lake). Predation
overpopulation and the exploitation of world resources, are
by the Nile perch has also resulted in the extinction of 65%
the focus of our ecological concerns.
of the cichlid species. Because many of the cichlids fed on
algae, the algae in the lake grew uncontrolled. When algae
died and decayed, much of the lake became depleted of Population
its oxygen. The introduction of nonnative water hyacinth, Global overpopulation is at the root of virtually all other
which has overgrown portions of the lake, has resulted environmental problems. Human population growth is
6 CHAPTER ONE
FIGURE 1.5
Introduction of the Nile perch (Lates niloticus) in an attempt to improve Lake Victoria’s fishery has resulted in the extinction of many cichlid
species and has indirectly contributed to decreased water quality and deforestation.
expected to continue in the twenty-first century. Virtu- T a b l e 1 . 3

ally all of this growth is in less developed countries, where W ORLD P OPULATION P ROJECTIONS FOR M AJOR
5.4 billion out of a total of 7.3 billion humans now live. Since W ORLD R EGIONS: 2010 AND 2050 (P ROJECTED )
a high proportion of the population is of childbearing age,
WORLD REGION 2012 2050 (PROJECTED)
the growth rate will increase in the twenty-first century. By
the year 2050, the total population of India (1.65 billion) is Africa 1.03 2.39
expected to surpass that of China (1.31 billion) and the Asia 4.16 5.16
total world population will reach 9.6 billion. The 2010 U.S.
Europe 0.74 0.71
population was 160 million. In 2050, it is projected to increase
to 401 million. Even though Africa does not have the high- Latin America and Caribbean 0.60 0.78
est human population, its population is increasing more rapidly North America 0.35 0.45
than other major regions of the world (table 1.3). As the human
Population sizes are based on figures from the United Nations Department of
population grows, the disparity between the wealthiest and Economics and Social Affairs (2014) and expressed in billions of people.
poorest nations is likely to increase.
World Resources and results in the extinction of many plant and animal spe-
cies, especially in tropical forests. Forest preservation would
Human overpopulation is stressing world resources.
result in the identification of new species of plants and ani-
Although new technologies continue to increase food pro-
mals that could be important human resources: new foods,
duction, most food is produced in industrialized countries
drugs, building materials, and predators of pests (figure 1.6).
that already have a high per-capita food consumption. Max-
Nature also has intrinsic value that is just as important as
imum oil production is expected to continue in this mil-
its provision of resources for humans. Recognition of this
lennium. Continued use of fossil fuels adds more carbon
intrinsic worth provides important aesthetic and moral
dioxide to the atmosphere, contributing to the greenhouse
impetus for preservation.
effect and climate change. Deforestation of large areas of
the world results from continued demand for forest prod-
ucts, fuel, and agricultural land. This trend contributes to Solutions
climate change by increasing atmospheric carbon dioxide An understanding of basic ecological principles can help
from burning forests and impairing the ability of the earth prevent ecological disasters like those we have described.
to return carbon to organic matter through photosynthesis. Understanding how matter is cycled and recycled in nature,
Deforestation also causes severe regional water shortages how populations grow, and how organisms in our lakes and
(a) (b)
FIGURE 1.6
Tropical Rain Forests: A Threatened World Resource. (a) A Brazilian tropical rain forest. (b) A bulldozer clear-cutting a rain forest in
the Solomon Islands. Clear-cutting for agriculture causes rain forest soils to quickly become depleted, and then the land is often abandoned
for richer soils. Cutting for roads breaks continuous forest coverage and allows for easy access to remote areas for exploitation. Loss of
tropical forests results in the extinction of many valuable forest species.
8 CHAPTER ONE
WILDLIFE ALERT
An Overview of the Problems
Extinction has been the fate of most plant and animal species. It (Recall that it is estimated that there are between 4 and 100 million
is a natural process that will continue. In recent years, however, species of animals living today.) An endangered species is in
the threat to the welfare of wild plants and animals has increased imminent danger of extinction throughout its range (where it
dramatically—mostly as a result of habitat destruction. Tropical lives). A threatened species is likely to become endangered
rain forests are one of the most threatened areas on the earth. in the near future. Box figure 1.1 shows the number of endan-
It is estimated that rain forests once occupied 14% of the earth’s gered and threatened species in different regions of the United
land surface. Today this has been reduced to approximately 6%. States. Clearly, much work is needed to improve these alarming
Each year we lose about 150,000 km2 of rain forest. This is an statistics.
area of the size of England and Wales combined. This decrease In the chapters that follow, you will learn that saving spe-
in habitat has resulted in tens of thousands of extinctions. Accu- cies requires more than preserving a few remnant individuals. It
rately estimating the number of extinctions is impossible in areas requires a large diversity of genes within species groups to pro-
like rain forests, where taxonomists have not even described most mote species survival in changing environments. This genetic
species. We are losing species that we do not know exist, and we diversity requires large populations of plants and animals.
are losing resources that could lead to new medicines, foods, and Preservation of endangered species depends on a multifaceted
textiles. Other causes of extinction include climate change, pollu- conservation plan that includes the following components:
tion, and invasions from foreign species. Habitats other than rain
forests—grasslands, marshes, deserts, and coral reefs—are also 1. A global system of national parks to protect large tracts of
being seriously threatened. land and wildlife corridors that allow movement between
No one knows how many species living today are close to natural areas
extinction. As of 2014, the U.S. Fish and Wildlife Service lists 2. Protected landscapes and multiple-use areas that allow con-
1,531 species in the United States as endangered or threatened. trolled private activity and also retain value as a wildlife habitat
The IUCN has assessed 71,000 species worldwide and of these 3. Zoos and botanical gardens to save species whose extinc-
more than 20,000 species are listed as endangered or threatened. tion is imminent
25
56 15
15 12 16
10
61 20
21 26
12 22 33
18 17
28 20
19 23 27 21
43 25
44 44 37 35
36 28 DC 23
22 7
319 36 55 70 30
64
67 104
52 25
35 43
431
46 137 76
107
31
128
BOX FIGURE 1.1 Map Showing Approximate Numbers of Endangered and Threatened Species in the United States.
Because the ranges of some organisms overlap two or more states, the sum of all numbers is greater than the sum of all endangered and threatened spe-
cies. The total number of endangered and threatened species in the United States is 1,531. The total number of listed animals is 655, with fish having the
greatest number of listed species..
forests use energy is fundamental to preserving the environ- SECTION REVIEW 1.2
ment. There are no easy solutions to our ecological problems.
Unless we deal with the problem of human overpopulation, As with the introduction of the Nile perch into Lake Victoria,
however, solving the other problems will be impossible. We our failure to understand complex ecological relationships
must work as a world community to prevent the spread of among animals often results in detrimental consequences
disease, famine, and other forms of suffering that accompany that require many decades, or even evolutionary time
overpopulation. Bold and imaginative steps toward improved frames, to heal. Many of these detrimental consequences
social and economic conditions and better resource manage- are direct or indirect results of the overpopulation of our
ment are needed. planet by our own species.
“Wildlife Alerts” that appear within selected chapters What is another example of how the careless disregard
of this text remind us of the peril that an unprecedented of ecological relationships has resulted in detrimental
number of species face around the world. Endangered or environmental consequences? (If you cannot think of
threatened species from a diverse group of animal phyla are an example on your own, see the “Wildlife Alert” boxes
highlighted. in subsequent chapters.)
S UMMARY 2. A change in the genetic makeup of populations of organisms

over time is a definition of
1.1 Zoology: An Evolutionary Perspective a. binomial nomenclature.
Zoology is the study of animals. It is a broad field b. organic evolution.
that requires zoologists to specialize in one or more c. evolution.
subdisciplines. d. ecology.
Animals share a common evolutionary past and evolutionary 3. Which of the following do zoologists use to study the genetic
forces that influenced their history. relationships among animals?
Evolution explains how the diversity of animals arose. a. Inherited morphological characteristics
Evolutionary relationships are the basis for the classification of b. Enzyme structure
animals into a hierarchical system. This classification system
c. DNA structure
uses a two-part name for every kind of animal. Higher levels of
classification denote more distant evolutionary relationships. d. All of the above are used by zoologists to study genetic
relationships.
1.2 Zoology: An Ecological Perspective
Animals share common environments, and ecological prin- 4. Which one of the following statements is true?
ciples help us understand how animals interact within those a. Members of the same class are always more closely related
environments. to each other than members of the same order.
Human overpopulation is at the root of virtually all other b. Members of different orders may be more closely related to
environmental problems. It stresses world resources and each other than members of the same family.
results in pollution, climate change, deforestation, and the c. Members of the same family are more closely related to
extinction of many plant and animal species. each other than members of different orders.
d. Members of the same order are always more closely related
C ONCEPT R EVIEW Q UESTIONS to each other than members of the same class.
5. All of the following may result from deforestation except one.
1. At least three of the following are examples of specialization Select the exception.
in zoology. Select the one choice that is not a specialization in a. Climate change is promoted.
zoology or select choice “e.” b. Extinction of many plant and animal species occurs.
a. Ichthyology c. Regional water shortages occur.
b. Mammalogy d. Long-term improvement in the standard of living in less
c. Ornithology developed countries occurs.
d. Histology e. Loss of important human resources such as new drugs and
e. All of the above are examples of specializations in zoology. food occurs.
10 CHAPTER ONE
6. By the year 2050, most human population growth will occur 3. Many of the ecological problems facing our world concern
in _____________ and result in a world population of events and practices that occur in less developed countries.
about _____________. Many of these practices are the result of centuries of cultural
a. less developed countries; 7 billion evolution. What approach should people and institutions of
b. less developed countries; 9.6 billion developed countries take in helping encourage ecologically
minded resource use?
c. less developed countries; 20.5 billion
4. Why should people in all parts of the world be concerned
d. developed countries; 5.5 billion
with the extinction of cichlids in Lake Victoria?
e. developed countries; 10.2 billion
A NALYSIS AND A PPLICATION

Q UESTIONS Enhance your study of this chapter with study tools and practice
tests. Also ask your instructor about the resources available through
1. How is zoology related to biology? What major biological con- Connect, including a media-rich eBook, interactive learning tools,
cepts, in addition to evolution and ecology, are unifying prin- and animations.
ciples shared between the two disciplines?
2. What are some current issues that involve both zoology and
questions of ethics or public policy? What should be the role
of zoologists in helping resolve these issues?
2
Cells, Tissues,
Organs, and Organ
Systems of Animals
This photomicrograph is a longitudinal section through skeletal muscle tissue,

one of the four major tissue types discussed in this chapter.
Chapter Outline
2.1 What Are Cells?
Because all organisms are made of cells, the cell is as fundamental to an under- 2.2 Why Are Most Cells Small?
standing of zoology as the atom is to an understanding of chemistry. In the hier- 2.3 Cell Membranes
archy of biological organization, the cell is the simplest organization of matter that Structure of Cell Membranes
Functions of Cell Membranes
exhibits all of the properties of life (figure 2.1). Some organisms are single celled; 2.4 Movement across Membranes
others are multicellular. An animal has a body composed of many kinds of special- Simple Diffusion
ized cells. A division of labor among cells allows specialization into higher levels Facilitated Diffusion
Osmosis
of organization (tissues, organs, and organ systems). Yet, everything that an animal Filtration
does is ultimately happening at the cellular level. Active Transport: Energy Required
Bulk Transport
2.5 Cytoplasm, Organelles, and Cellular
Components
2.1 W HAT A RE C ELLS ? Cytoplasm
Ribosomes: Protein Workbenches
Endoplasmic Reticulum: Production
LEARNING OUTCOMES and Transport
Golgi Apparatus: Packaging, Sorting,
1. Differentiate between a prokaryotic and eukaryotic cell. and Export
Lysosomes: Digestion
2. Describe the three parts of a eukaryotic cell. and Degradation
Microbodies: A Diverse Category
Cells are the functional units of life, in which all of the chemical reactions neces- of Organelles
sary for the maintenance and reproduction of life take place. They are the small- Mitochondria: Power Generators
est independent units of life. There are two basic types of cells: prokaryotic and Cytoskeleton: Microtubules,
Intermediate Filaments,
eukaryotic. The prokaryotes lack nuclei and other membrane-bound organelles. and Microfilaments
These simpler (prokaryotic or prokaryotes; “before nucleus”) cells are classified Cilia and Flagella: Movement
into two domains: Archaea and Eubacteria. The Archaea have unique characteristics Centrioles and Microtubule-
Organizing Centers
and also share features with Eubacteria and the third domain, Eukarya. Eukaryotic Vacuoles: Cell Maintenance
cells are larger and more complex than prokaryotic cells. Since animals and protista Vaults: Mysterious Symmetrical Shells
are composed of eukaryotic cells, this cell type will be emphasized in this chapter. Exosomes
2.6 The Nucleus: Information Center
Table 2.1 compares prokaryotic and eukaryotic cells. Nuclear Envelope: Gateway
All eukaryotes (“true nucleus”) have cells with a membrane-bound nucleus to the Nucleus
containing DNA. In addition, eukaryotic cells contain many other structures called Chromosomes: Genetic
Containers
organelles (“little organs”) that perform specific functions. Eukaryotic cells also Nucleolus: Preassembly Point for
have a network of specialized structures called microfilaments and microtubules Ribosomes
organized into the cytoskeleton, which gives shape to the cell and allows intracel- 2.7 Levels of Organization in Various
Animals
lular movement. 2.8 Tissues
All eukaryotic cells have three basic parts: Epithelial Tissue: Many Forms
and Functions
1. The plasma membrane is the outer boundary of the cell. It separates the Connective Tissue: Connection
and Support
internal metabolic events from the environment and allows them to proceed in Nervous Tissue: Communication
organized, controlled ways. The plasma membrane also has specific receptors Muscle Tissue: Movement
for external molecules that alter the cell’s function. 2.9 Organs
2.10 Organ Systems
12 CHAPTER TWO
Animal TABLE 2.1

C OMPARISON OF P ROKARYOTIC
AND E UKARYOTIC C ELLS
Organ systems
Increasing complexity
COMPONENT PROKARYOTE EUKARYOTE
Organization of
Organs genetic material
True membrane- Absent Present
bound nucleus
Tissues
DNA complexed No Yes
with histones
Number of One More than one
chromosomes
Cells Nucleolus Absent Present
Living
Mitosis occurs No Yes
Nonliving
Genetic Partial, unidirectional Meiosis and fusion
Organelles recombination transfer of DNA of gametes
Mitochondria Absent Present
Chloroplasts Absent Present
Membranes Plasma Usually no Yes
membrane
Decreasing complexity
with sterols
Flagella Submicroscopic in Microscopic in
Macromolecules size; composed size; membrane
of only one fiber bound; usually
20 microtubules
in 9 1 2 pattern
Simple molecules
Endoplasmic Absent Present
reticulum
Golgi apparatus Absent Present
Atoms Cell walls Usually chemically Chemically simpler
complex
Simpler organelles
FIGURE 2.1 Ribosomes 70S 80S (except in
Structural Hierarchy in a Multicellular Animal. At each level, mitochondria
function depends on the structural organization of that level and and chloroplasts)
those below it.
Lysosomes and Absent Present
peroxisomes
Microtubules Absent or rare Present
2. Cytoplasm (Gr. kytos, hollow vessel 1 plasm, fluid) Cytoskeleton May be absent Present
is the portion of the cell outside the nucleus. The Vacuoles Present Present
semifluid portion of the cytoplasm is called the Vesicles Present Present
cytosol. Suspended within the cytosol are Differentiation Rudimentary Tissues and organs
the organelles.
3. The nucleus (pl., nuclei) is the cell control center.
It contains the chromosomes and is separated SECTION REVIEW 2.1
from the cytoplasm by its own nuclear envelope. Prokaryotes are small cells that lack complex internal organiza-
The nucleoplasm is the semifluid material in the tion. The two prokaryotic domains are Archaea and Eubacteria.
nucleus. Eukaryotic cells exhibit compartmentalization and various organ-
elles that carry out specific functions. The three parts of a eukary-
Because cells vary so much in form and function, no
otic cell are the plasma membrane, cytoplasm, and nucleus.
“typical” cell exists. However, to help you learn as much as
possible about cells, figure 2.2 shows an idealized version of What are some similarities between eukaryotic cells
a eukaryotic cell and most of its component parts. and the prokaryotic cells of Eubacteria and Archaea?
Cells, Tissues, Organs, and Organ Systems of Animals 13
Microtubule-organizing
centers (with centriole pairs)
Lysosome Microtubules
Mitochondrion
Plasma
membrane Radius ( r ) 1 cm 2 cm 4 cm
2 2
Surface area (SA) 12.57 cm 50.26 cm 201.06 cm 2
Nuclear
envelope Volume (V) 4.19 cm 3 33.51 cm 3 268.08 cm 3
Nucleus
SA/V 3.0 1.50 0.75
Nucleolus
Chromatin
Surface area of a sphere = 4π r 2
Volume of sphere = 4 /3 πr 3
Golgi
apparatus
Polyribosome FIGURE 2.3

(polysome)
The Relationship between Surface Area and Volume. As the
Vesicle radius of a sphere increases, its volume increases more rapidly than
its surface area. (SA/V 5 surface-area-to-volume ratio.)
Free ribosomes
SECTION REVIEW 2.2
Rough Smooth
endoplasmic endoplasmic A cell needs a surface area large enough to allow efficient
reticulum reticulum movement of nutrients into the cell and waste material out of
FIGURE 2.2 the cell. Small cells have a lot more surface area per volume
A Generalized Animal Cell. Understanding of the structures in than large cells. For example, a 4-cm cube has a surface-area-
this cell is based mainly on electron microscopy. The sizes of some to-volume ratio of only 5.5:1, but a 1-cm cube has a ratio of 6:1.
organelles and structures are exaggerated to show detail.
If the cell radius of a cell increases 10 times, the sur-
face area will increase by 100 times. How much will
2.2 W HY A RE M OST the volume increase?
C ELLS S MALL ?
2.3 C ELL M EMBRANES
LEARNING OUTCOMES
1. Explain why most cells are small. LEARNING OUTCOME
2. Determine how surface area changes as a function 1. Relate the structure of the plasma membrane to the
of volume. function of the membrane.
Most cells are small and can be seen only with the aid of a The plasma membrane surrounds the cell. Other membranes
microscope. (Exceptions include the eggs of most vertebrates inside the cell enclose some organelles and have properties
[fishes, amphibians, reptiles, and birds] and some long nerve similar to those of the plasma membrane.
cells.) One reason for the small size of cells is that the ratio of
the volume of the cell’s nucleus to the volume of its cytoplasm
must not be so small that the nucleus, the cell’s major control
Structure of Cell Membranes
center, cannot control the cytoplasm. In 1972, S. Jonathan Singer and Garth Nicolson developed the
Another aspect of cell volume works to limit cell size. fluid-mosaic model of membrane structure. According to this
As the radius of a cell lengthens, cell volume increases more model, a membrane is a double layer (bilayer) of proteins and
rapidly than cell surface area (figure 2.3). The need for nutri- phospholipids and is fluid rather than solid. The phospholipid
ents and the rate of waste production are proportional to cell bilayer forms a fluid “sea” in which specific proteins float like
volume. The cell takes up nutrients and eliminates wastes icebergs (figure 2.4). Being fluid, the membrane is in a constant
through its surface plasma membrane. If cell volume becomes state of flux—shifting and changing, while retaining its uniform
too large, the surface-area-to-volume ratio is too small for an structure. The word mosaic refers to the many different kinds of
adequate exchange of nutrients and wastes. proteins dispersed in the phospholipid bilayer.
14 CHAPTER TWO
Carbohydrate
Outside cell
Fibrous
proteins
Glycolipid
Double layer of
phospholipid
molecules
Phospholipid
“head”
Cytoplasmic side
of membrane
Globular Phospholipid
Cholesterol protein “tail”
molecules
FIGURE 2.4
Fluid-Mosaic Model of Membrane Structure. Intrinsic globular proteins may protrude above or below the lipid bilayer and may move
about in the membrane. Peripheral proteins attach to either the inner surface or the outer surface.
The following are important points of the fluid-mosaic Hydrophilic (polar)

model: head
1. The phospholipids have one polar end and one nonpolar Water
end. The polar ends are oriented on one side toward the Region stiffened
by cholesterol
outside of the cell and into the fluid cytoplasm on the Hydrophobic
(nonpolar) tail
other side, and the nonpolar ends face each other in the
middle of the bilayer. The “tails” of both layers of phos-
pholipid molecules attract each other and are repelled
by water (they are hydrophobic, “water dreading”). As More fluid
region
a result, the polar spherical “heads” (the phosphate por-
tion) are located over the cell surfaces (outer and inner) Cholesterol
and are “water attracting” (they are hydrophilic).
2. Cholesterol is present in the plasma membrane and Phospholipid Phospholipid
organelle membranes of eukaryotic cells. The cholesterol
FIGURE 2.5
molecules are embedded in the interior of the mem-
The Arrangement of Cholesterol between Lipid Molecules
brane and help make the membrane less permeable to
of a Lipid Bilayer. Cholesterol stiffens the outer lipid bilayer and
water-soluble substances. In addition, the relatively rigid causes the inner region of the bilayer to become slightly more fluid.
structure of the cholesterol molecules helps stabilize the Only half the lipid bilayer is shown; the other half is a mirror image.
membrane (figure 2.5).
3. The membrane proteins are individual molecules
attached to the inner or outer membrane surface they form glycolipids on the surface of the plasma
(peripheral proteins) or embedded in it (intrinsic pro- membrane. Surface carbohydrates and portions of the
teins) (see figure 2.4 ). Some intrinsic proteins are links proteins and lipids make up the glycocalyx (“cell coat”)
to sugar-protein markers on the cell surface. Other (figure 2.6). This arrangement of distinctively shaped
intrinsic proteins help move ions or molecules across groups of sugar molecules of the glycocalyx acts as a
the membrane, and still others attach the membrane to molecular “fingerprint” for each cell type. The glyco-
the cell’s inner scaffolding (the cytoskeleton) or to vari- calyx is necessary for cell-to-cell recognition and the
ous molecules outside the cell. behavior of certain cells, and it
4. When carbohydrates unite with proteins, they form is a key component in coordi- MP3
Membrane Structure
glycoproteins, and when they unite with lipids, nating cell behavior in animals.
How Do Zoologists Investigate the Inner

Workings of the Tiny Structures within a Cell?
T he small size of cells is the

greatest obstacle to dis-
covering their nature and
the anatomy of the tiny structures
within cells. The evolution of sci-
Most commonly used are the light
microscope, the transmission
electron microscope (TEM), the
scanning electron microscope,
the fluorescence microscope, the
modern cell biology developed
from an integration of cytology with
biochemistry, the study of molecules
and the chemical processes of
metabolism. Throughout this book,
ence often parallels the invention scanning tunneling microscope, many photographs are presented
of instruments that extend human and the atomic force microscope. using various microscopes to show
senses to new limits. Cells were Microscopes are the most impor- different types of cells and the various
discovered after microscopes were tant tools of cytology, the study of cell tiny structures within. From these
invented, and high-magnification structure. But simply describing the photographs, it will become apparent
microscopes are needed to see the diverse structures within a cell reveals that similarities among cells reveal
smallest structures within a cell. little about their function. Today’s the evolutionary unity of life.
Extracellular
specific cell identification markers that differentiate one cell
fluid type from another.
(outside of The ability of the plasma membrane to let some sub-
cell)
Sugar molecules stances in and keep others out is called selective permeability
(L. permeare or per, through 1 meare, pass) and is essen-
Glycolipid
Glyco- tial for maintaining a “steady state” within the cell. How-
calyx ever, before you can fully understand how substances pass
Proteins into and out of cells and organelles, you must know how
Glyco-
protein
the molecules of those substances move from one place
to another.
SECTION REVIEW 2.3

The major components of the plasma membrane are as follows:
a phospholipid bilayer, cholesterol, membrane proteins, and
Lipid the glycocalyx. This structure creates the outer boundary of the
Protein bilayer
fibers cell, it separates the internal metabolic events from the environ-
ment, and it allows the events to proceed in an organized, con-
trolled way. The plasma membrane also has specific structures
Cytoplasm
for movement of materials into and out of the cell and recep-
tors for external molecules that alter the cell’s function.
FIGURE 2.6
If the plasma membrane of a cell were just a single layer
The Glycocalyx, Showing the Glycoproteins and Glycolipids. Note
that all of the attached carbohydrates are on the outside of the plasma of phospholipids, how would this affect its function?
membrane.
Functions of Cell Membranes 2.4 M OVEMENT ACROSS
Cell membranes (1) regulate material moving into and out of

M EMBRANES
the cell, and from one part of the cell to another; (2) separate
the inside of the cell from the outside; (3) separate various
LEARNING OUTCOMES
organelles within the cell; (4) provide a large surface area on 1. Differentiate the different processes by which material can
which specific chemical reactions can occur; (5) separate cells move into and out of the cell through the plasma membrane.
from one another; and (6) are a site for receptors containing 2. Explain the movement of water by osmosis.
16 CHAPTER TWO
Molecules can cross membranes in a number of ways, both of the short-distance transport of substances moving into
by using their own energy and by relying on an outside and out of cells. Figure 2.7 shows the diffusion of sugar
energy source. Table 2.2 summarizes the various kinds of particles away from a sugar
Animation MP3
transmembrane movement, and the sections that follow dis- cube placed in water. Diffusion Diffusion
cuss them in more detail.
Simple Diffusion Facilitated Diffusion

Molecules move randomly at all temperatures above absolute Polar molecules (not soluble in lipids) may diffuse through
zero (2273° C) (due to spontaneous molecular motion) from protein channels (pores) in the lipid bilayer (figure 2.8). The
areas where they are highly concentrated to areas where they protein channels offer a continuous pathway for specific mol-
are less concentrated, until they are evenly distributed in a state ecules to move across the plasma membrane so that they
of dynamic equilibrium. This process is simple diffusion never come into contact with the hydrophobic layer or the
(L. diffundere, to spread). Simple diffusion accounts for most membrane’s polar surface.
TABLE 2.2
D IFFERENT T YPES OF M OVEMENT ACROSS P LASMA M EMBRANES
TYPE OF MOVEMENT DESCRIPTION EXAMPLE IN THE BODY OF A FROG
Simple diffusion No cell energy is needed. Molecules move “down” a A frog inhales air containing oxygen, which
concentration gradient. Molecules spread out randomly moves into the lungs and then diffuses into
from areas of higher concentration to areas of lower the bloodstream.
concentration until they are distributed evenly in a state of
dynamic equilibrium.
Facilitated diffusion Carrier (transport) proteins in a plasma Glucose in the gut of a frog combines with
membrane temporarily bind with molecules carrier proteins to pass through the gut cells
and help them pass across the membrane. into the bloodstream.
Other proteins form channels through which
molecules move across the membrane.
Osmosis Water molecules diffuse across selectively permeable Water molecules move into a frog’s red
membranes from areas of higher concentration to blood cell when the concentration of water
areas of lower concentration. molecules outside the blood cell is greater
than it is inside.
Filtration Essentially protein-free plasma moves across capillary A frog’s blood pressure forces water and
walls due to a pressure gradient across the wall. dissolved wastes into the kidney tubules
during urine formation.
Active transport Specific carrier proteins in the plasma membrane Sodium ions move from inside the neurons
bind with molecules or ions to help them cross the of the sciatic nerve of a frog (the sodium-
membrane against a concentration gradient. Cellular potassium pump) to the outside of the
energy is required. neurons.
Endocytosis The bulk movement of material into a cell by the
formation of a vesicle.
Pinocytosis The plasma membrane encloses small amounts of fluid The kidney cells of a frog take in fluid to
droplets (in a vesicle) and takes them into the cell. maintain fluid balance.
Phagocytosis The plasma membrane forms a vesicle around a The white blood cells of a frog engulf and
solid particle or other cell and draws it into the digest harmful bacteria.
phagocytic cell.
Receptor-mediated Extracellular molecules bind with specific receptor The intestinal cells of a frog take up large
endocytosis proteins on a plasma membrane, causing the membrane molecules from the inside of the gut.
to invaginate and draw molecules into the cell.
Exocytosis The bulk movement of material out of a cell. A vesicle The sciatic nerve of a frog releases a chemical
(with particles) fuses with the plasma membrane (neurotransmitter).
and expels particles or fluids from the cell across the
plasma membrane. The reverse of endocytosis.
(a) (b) (c) (d)
FIGURE 2.7
Simple Diffusion. When a sugar cube is placed in water (a), it slowly dissolves (b) and disappears. As this happens, the sugar molecules
diffuse from a region where they are more concentrated to a region (c) where they are less concentrated. Even distribution of the sugar
molecules throughout the water is diffusion equilibrium (d).
FIGURE 2.8 FIGURE 2.9

Transport Proteins. Molecules can move into and out of Facilitated Diffusion and Carrier (Transport) Proteins. Some
cells through integrated protein channels (pores) in the plasma molecules move across the plasma membrane with the assistance
membrane without using energy. of carrier proteins that transport the molecules down their
concentration gradient, from a region of higher concentration to a
region of lower concentration. A carrier protein alternates between
Large molecules and some (e.g., glucose and amino acids)
two configurations, moving a molecule across a membrane as
of those not soluble in lipids require assistance in passing across the shape of the protein changes. The rate of facilitated diffusion
the plasma membrane. These molecules use facilitated depends on how many carrier proteins are available in the
diffusion, which, like simple diffusion, requires no energy input. membrane and how fast they can move their specific molecules.
To pass across the membrane, a molecule temporarily binds with
a carrier (transport) protein in the plasma membrane and is trans-
ported from an area of higher concentration to Animation Recent studies show that water, despite its polarity, can
an area of lower concentration (figure 2.9). Facilitated cross cell membranes, but this flow is limited. Water flow in
Diffusion
living cells is facilitated by specialized water channels called
aquaporins. Aquaporins fall into two general classes: those that
Osmosis are specific only for water, and others that allow small hydro-
The diffusion of water across a selectively permeable mem- philic molecules (e.g., urea, glycerol) to cross the membrane.
brane from an area of higher concentration to an area of lower The term tonicity (Gr. tonus, tension) refers to the relative
concentration is osmosis (Gr. osmos, pushing). Osmosis is just concentration of solutes in the water inside and outside the cell.
a special type of diffusion, not a different method (figure 2.10). For example, in an isotonic (Gr. isos, equal 1 tonus, tension)
18 CHAPTER TWO
Selectively permeable Water molecule Isotonic Hypertonic Hypotonic

membrane Sugar molecule Solution Solution Solution
1 2 1 2
Normal cells Shriveled cells Cells swell and

eventually burst
(a) (b) (c)
Time
FIGURE 2.11
(a) (b) Effect of Salt Concentration on Red Blood Cell Volumes.
(a) An isotonic solution with the same salt concentration inside
FIGURE 2.10 and outside the cell has no effect on the size of the red blood cell.
Osmosis. (a) A selectively permeable membrane separates the (b) A hypertonic (high-salt) solution causes water to leave the red
beaker into two compartments. Initially, compartment 1 contains blood cell, which shrinks. (c) A hypotonic (low-salt) solution results
sugar and water molecules, and compartment 2 contains only in an inflow of water, causing the red blood cell to swell. Arrows
water molecules. Due to molecular motion, water moves down the indicate direction of water movement.
concentration gradient (from compartment 2 to compartment 1) by
osmosis. The sugar molecules remain in compartment 1 because
they are too large to pass across the membrane. (b) At osmotic
equilibrium, the number of sugar molecules in compartment 1 does
not increase, but the number of water molecules does.
solution, the solute concentration is the same inside and outside

a red blood cell (figure 2.11a). The concentration of water mol-
ecules is also the same inside and outside the cell. Thus, water
molecules move across the plasma membrane at
the same rate in both directions, and there is no MP3
Osmosis
net movement of water in either direction.
In a hypertonic (Gr. hyper, above) solution, the solute
concentration is higher outside the red blood cell than it is inside.
Because the concentration of water molecules inside the cell is
FIGURE 2.12
higher than it is outside, water moves out of
the cell, which shrinks (figure 2.11b). This con- Animation Filtration. The high blood pressure in the capillary forces small
Osmosis molecules through the capillary membrane. Larger molecules cannot pass
dition is called crenation in red blood cells. through the small openings in the capillary membrane and remain in the
In a hypotonic (Gr. hypo, under) solution, the solute con- capillary. Arrows indicate the direction of small molecule movement.
centration is lower outside the red blood cell than it is inside.
Conversely, the concentration of water molecules is higher out-
Active Transport: Energy Required
side the cell than it is inside. As a result, water moves into the
cell, which swells and may burst (figure 2.11c). Active-transport processes move molecules across a selec-
tively permeable membrane against a concentration gradient—
that is, from an area of lower concentration to an area of
Filtration higher concentration. This movement against the concentra-
Filtration is a process that forces small molecules across tion gradient requires ATP energy.
selectively permeable membranes with the aid of hydrostatic The active-transport process is similar to facilitated dif-
(water) pressure (or some other externally applied force, fusion, except that the carrier protein in the plasma mem-
such as blood pressure). For example, in the body of an ani- brane must use energy to move the molecules against their
mal such as a frog, filtration is evident when blood pressure concentration gradient (figure 2.13). These carrier proteins
forces water and dissolved molecules through the permeable are called uniporters if they transport a single type of mol-
walls of small blood vessels called capillaries (figure 2.12). ecule or ion, symporters if they transport two molecules or
In filtration, large molecules, such as proteins, do not pass ions in the same direction, and antiporters if they transport
through the smaller membrane pores. Filtration also takes two molecules or ions in the opposite direction.
place in the kidneys when blood pressure forces water and One active-transport mechanism, the sodium-potassium
dissolved wastes out of the blood vessels and into the kidney pump, helps maintain the high concentrations of potassium
tubules in the first step in urine formation. ions and low concentrations of sodium ions inside nerve cells
Endocytosis
Invagination Formation Formation
of pouch of vesicle
Extracellular
fluid
Extracellular
substances
Cytoplasm now within
vesicle
Exocytosis
Joining of Secretion of
vesicle cellular
with plasma product
membrane
Secretion
now in
extracellular
fluid
FIGURE 2.14
Endocytosis and Exocytosis. Endocytosis and exocytosis are
FIGURE 2.13 responsible for the bulk transport of molecules into and out of a cell.
Active Transport. During active transport, a molecule combines
with a carrier protein whose shape is altered as a result of the
combination. This change in configuration, along with ATP energy, In the process of exocytosis (Gr. exo, outside), the
helps move the molecule across the plasma membrane against a secretory vesicles fuse with the plasma membrane and release
concentration gradient. their contents into the extracellular environment (figure 2.14).
This process adds new membrane material, which replaces
the plasma membrane lost during exocytosis.
that are necessary for the transmission of electrical impulses.
Another active-transport mechanism, the calcium pump, keeps
the calcium concentration hundreds of Animation SECTION REVIEW 2.4
Sodium-Potassium
times lower inside the cell than outside. Pump The different processes by which material moves into and
out of the cell through the plasma membrane include simple
diffusion, facilitated diffusion, osmosis, filtration, active trans-
Bulk Transport port, bulk transport, endocytosis, (pinocytosis, phagocytosis,
Large molecules cannot be transported through the plasma and receptor-mediated endocytosis) and exocytosis. Water
membrane by the processes described in the previous sections. passes through the plasma membrane and through aqua
Endocytosis and exocytosis together provide bulk transport porins in response to solute concentration differences inside
into and out of the cell, respectively. (The Animation and outside the cell. This transport process is called osmosis.
Endocytosis and
term “bulk” is used because many mole- Exocytosis If you require that drugs be given to you by an intrave-
cules are moved at the same time.) nous (IV) process, what should the concentration of sol-
In endocytosis (Gr. endon, within), the plasma mem- utes in the IV solution be relative to your red blood cells?
brane envelops large particles and molecules (figure 2.14)
and moves them in bulk across the membrane. The three
forms of endocytosis are pinocytosis, phagocytosis, and 2.5 C YTOPLASM, O RGANELLES,
receptor-mediated endocytosis.
Pinocytosis (Gr. pinein, to drink 1 cyto, cell) is the
AND C ELLULAR C OMPONENTS
nonspecific uptake of small droplets of extracellular fluid.
Phagocytosis (Gr. phagein, to eat 1 cyto, cell) is similar to
LEARNING OUTCOMES
pinocytosis except that the cell takes in solid material rather 1. Relate the structure of the major cellular organelles to
than liquid. Receptor-mediated endocytosis involves a spe- their function.
cific receptor protein on the plasma membrane that “recog- 2. Explain the function of the cytoskeleton.
nizes” an extracellular molecule and binds with it. The reaction
stimulates the membrane to indent and create a vesicle con- Many cell functions that are performed in the cytoplasmic
taining the selected molecule. compartment result from the activity of specific structures
20 CHAPTER TWO
called organelles. Organelles effectively compartmentalize

a cell’s activities, improving efficiency and protecting cell
contents from harsh chemicals. Organelles also enable cells
to secrete various substances, derive energy from nutrients,
degrade debris and waste materials, and reproduce. Table 2.3
summarizes the structure and function
of these organelles, and the sections that MP3
Cellular Organelles
follow discuss them in more detail.
Cytoplasm
The cytoplasm of a cell has two distinct parts: (1) The
cytomembrane (or endomembrane) system consists of
well-defined structures, such as the endoplasmic reticulum,
Golgi apparatus, vacuoles, and vesicles. (2) The fluid cytosol (a)
suspends the structures of the cytomembrane system and con-
tains various dissolved molecules.
Ribosomes: Protein Workbenches

Ribosomes are non-membrane-bound structures that are the
sites for protein synthesis. They contain almost equal amounts
of protein and a special kind of ribonucleic acid called ribo-
somal RNA (rRNA). Some ribosomes attach to the endoplas-
mic reticulum (see next section), and some float freely in the
cytoplasm. Whether ribosomes are free or attached, they usu-
ally cluster in groups connected by a strand of another kind of
ribonucleic acid called messenger RNA (mRNA). These clus-
ters are called polyribosomes or polysomes (see figure 2.2).
(b)
Endoplasmic Reticulum:
FIGURE 2.15
Production and Transport
Endoplasmic Reticulum (ER). (a) Ribosomes coat rough ER.
The endoplasmic reticulum (ER) is a complex, membrane- Notice the double membrane and the lumen (space) within it.
bound labyrinth of flattened sheets, sacs, and tubules that (b) Smooth ER lacks ribosomes.
branches and spreads throughout the cytoplasm. The ER is con-
tinuous from the nuclear envelope to the plasma membrane
Proteins that ribosomes synthesize are sealed off in little
(see figure 2.2) and is a series of channels that helps various
packets called transfer vesicles. Transfer vesicles pass from
materials to circulate throughout the cytoplasm. It also is a stor-
the ER to the Golgi apparatus and fuse with it (figure 2.16b).
age unit for enzymes and other proteins and a point of attach-
In the Golgi apparatus, the proteins are concentrated and
ment for ribosomes. ER with attached ribosomes is rough ER
chemically modified. One function of this chemical modifica-
(figure 2.15a), and ER without attached ribosomes is smooth ER
tion seems to be to mark and sort the proteins into different
(figure 2.15b). Smooth ER is the site for lipid production, detox-
batches for different destinations. Eventually, the proteins are
ification of a wide variety of organic molecules, and storage of
packaged into secretory vesicles, which are released into the
calcium ions in muscle cells. Most cells contain both types of
cytoplasm close to the plasma membrane. When the vesicles
ER, although the relative proportion varies among cells.
reach the plasma membrane, they fuse with it and release
their contents to the outside of the cell by exocytosis. Golgi
Golgi Apparatus: apparatuses are most abundant in cells that secrete chemical
Packaging, Sorting, and Export substances (e.g., pancreatic cells secreting digestive enzymes
and nerve cells secreting neurotransmitters). As noted in the
The Golgi apparatus or complex (named for Camillo Golgi, next section, the Golgi apparatus also produces lysosomes.
who discovered it in 1898) is a collection of membranes asso-
ciated physically and functionally with the ER in the cytoplasm
(figure 2.16a; see also figure 2.2). It is composed of flattened
Lysosomes: Digestion and Degradation
stacks of membrane-bound cisternae (sing., c isterna; L. closed Lysosomes (Gr. lyso, dissolving 1 soma, body) are membrane-
spaces serving as fluid reservoirs). The Golgi a pparatus sorts, bound spherical organelles that contain enzymes called acid
packages, and secretes proteins and lipids. hydrolases, which are capable of digesting organic molecules
TABLE 2.3
S TRUCTURE AND F UNCTION OF E UKARYOTIC C ELLULAR C OMPONENTS
COMPONENT STRUCTURE/DESCRIPTION FUNCTION
Centriole Located within microtuble-organizing center; Forms basal body of cilia and flagella; functions in
contains nine triple microtubules mitotic spindle formation
Chloroplast Organelle that contains chlorophyll and is Traps, transforms, and uses light energy to convert
involved in photosynthesis carbon dioxide and water into glucose and oxygen
Chromosome Made up of nucleic acid (DNA) and protein Controls heredity and cellular activities
Cilia, flagella Threadlike processes Cilia and flagella move small particles past fixed cells
and are a major form of locomotion in some cells
Cytomembrane system The endoplasmic reticulum, Golgi apparatus, Organelles, functioning as a system, modify, package,
vacuoles, and vesicles and distribute newly formed proteins and lipids
Cytoplasm Semifluid enclosed within plasma membrane; consists Dissolves substances; houses organelles and
of fluid cytosol and cytomembrane system vesicles
Cytoskeleton Interconnecting microfilaments and microtubules; Assists in cell movement; provides support; site for
flexible cellular framework binding of specific enzymes
Cytosol Fluid part of cytoplasm; enclosed within plasma Houses organelles; serves as fluid medium for
membrane; surrounds nucleus metabolic reactions
Endoplasmic reticulum (ER) Extensive membrane system extending Storage and internal transport; rough ER is a site
throughout the cytoplasm from the plasma for attachment of ribosomes; smooth ER makes
membrane to the nuclear envelope lipids
Exosome Cell-derived vessicles Carry signals to distant parts of an animal’s body
Golgi apparatus Stacks of disklike membranes Sorts, packages, and routes cell’s synthesized
products
Lysosome Membrane-bound sphere Digests materials
Microbodies Vesicles that are formed from the incorporation of Isolate particular chemical activities from the rest
lipids and proteins and that contain oxidative of the cell
and other enzymes; for example, peroxisomes
Microfilament (actin filament) Rodlike structure containing the protein actin Gives structural support and assists in cell
movement
Microtubule Hollow, cylindrical structure Assists in movement of cilia, flagella, and
chromosomes; transport system
Microtubule-organizing center Cloud of cytoplasmic material that contains Dense site in the cytoplasm that gives rise to large
centrioles numbers of microtubules with different functions
in the cytoskeleton
Mitochondrion Organelle with double, folded membranes Converts energy into a form the cell can use
Nucleolus Rounded mass within nucleus; contains RNA Preassembly point for ribosomes
and protein
Nucleus Spherical structure surrounded by a nuclear Contains DNA that controls cell’s genetic program
envelope; contains nucleolus and DNA and metabolic activities
Plasma membrane The outer bilayered boundary of the cell; Protection; regulation of material movement;
composed of protein, cholesterol, and cell-to-cell recognition
phospholipids
Ribosome Contains RNA and protein; some are free Site of protein synthesis
and some attach to ER
Vacuole Membrane-surrounded, often large, sac in the Storage site of food and other compounds; also
cytoplasm pumps water out of a cell (e.g., contractile vacuole)
Vaults Cytoplasmic ribonucleoproteins shaped like Dock at nuclear pores: believed to transport
octagonal barrels messenger RNA from the nucleus to the
ribosomes
Vesicle Small, membrane-surrounded sac; contains Site of intracellular digestion, storage, or transport
enzymes or secretory products
22 CHAPTER TWO
Lysosome engulfing Lysosomes digesting

damaged organelle phagocytized material
Plasma
membrane
Phagolysosome
Lysosome
Cisternae
Food vacuole
(a)
Transfer vesicle Phagocytosis of
from ER food particle
Budding
vesicle
Transport
vesicle with
enzymes
Rough ER Golgi apparatus Smooth ER Food particle
FIGURE 2.17
Lysosome Formation and Function. Lysosomes arise from the
Secretory Golgi apparatus and fuse with vesicles that have engulfed foreign
vesicles material to form digestive vesicles (phagolysosomes). These
Cytoplasm vesicles function in the normal recycling of cell constituents.
hydrogen peroxide. (If these oxidative enzymes were not

Plasma Exocytosis
isolated within microbodies, they would disrupt metabolic
membrane pathways.) Hydrogen peroxide is dangerous to cells because
(b) Outside of cell of its violent chemical reactivity. It is generated during mito-
chondrial respiration and is an inducer of oxidative damage
FIGURE 2.16 and a mediator of aging. The enzyme in the peroxisome is
Golgi Apparatus. (a) The Golgi apparatus consists of a stack catalase, which breaks down hydrogen peroxide to water
of cisternae. Notice the curved nature of the cisternae. (b) The
Golgi apparatus stores, sorts, packages, and secretes cell products.
and oxygen, which are both beneficial to cells.
Secretory vesicles move from the Golgi apparatus to the plasma
membrane and fuse with it, releasing their contents to the outside
of the cell via exocytosis. Mitochondria: Power Generators
Mitochondria (sing., mitochondrion) are double-membrane-
bound organelles that are spherical to elongated in shape.
(lipids, proteins, nucleic acids, and polysaccharides) under A small space separates the outer membrane from the inner
acidic conditions. The enzymes are synthesized in the ER, membrane. The inner membrane folds and doubles in on
transported to the Golgi apparatus for processing, and then itself to form incomplete partitions called cristae (sing., crista;
secreted by the Golgi apparatus in the form of lysosomes or as figure 2.18). The cristae increase the surface area available for
vesicles that fuse with lysosomes (figure 2.17). Lysosomes fuse the chemical reactions that trap usable energy for the cell. The
with phagocytic vesicles, thus exposing the Animation space between the cristae is the matrix. The matrix contains
vesicle’s contents to lysosomal enzymes. Lysosomes
ribosomes, circular DNA, and other material. Because they
convert energy to a usable form, mitochondria are frequently
called the “power generators” of the cell. Mitochondria usually
Microbodies: A Diverse multiply when a cell needs additional high-energy molecules.
Category of Organelles
Eukaryotic cells contain a variety of enzyme-bearing, membrane- Cytoskeleton: Microtubules,
enclosed vesicles called microbodies. The distribution of Intermediate Filaments,
enzymes into microbodies is one main way eukaryotic cells
organize their metabolism.
and Microfilaments
One specific type of microbody is the peroxisome. In most cells, the microtubules, intermediate filaments, and
Peroxisomes contain enzymes that catalyze the removal of microfilaments form the flexible cellular framework called
electrons and associated hydrogen atoms from, for example, the cytoskeleton (“cell skeleton”) (figure 2.19). This latticed
Outer mitochondrial 25 nm 10 nm 7 nm
membrane
Tubulin Actin
subunit subunit
Matrix
Crista
Microtubule Intermediate Microfilament
filament
(a) (b) (c)
FIGURE 2.20
Three Major Classes of Protein Fibers Making Up the
Cytoskeleton of Eukaryotic Cells. (a) Microtubules consist
of globular protein subunits (tubulins) linked in parallel rows.
(b) Intermediate filaments in different cell types are composed
of different protein subunits. (c) The protein actin is the key
subunit in microfilaments.
Inner mitochondrial
membrane framework extends throughout the cytoplasm, connecting
FIGURE 2.18 the various organelles and cellular components.
Mitochondrion. Mitochondrial membranes, cristae, and matrix.
Microtubules are hollow, slender, cylindrical structures
The matrix contains DNA, ribosomes, and enzymes. in animal cells. Each microtubule is made of spiraling sub-
units of globular proteins called tubulin subunits (figure 2.20a).
Microtubules function in the movement of organelles, such as
secretory vesicles, and in chromosome movement during divi-
sion of the cell nucleus. They are also part of a transport system
within the cell. For example, in nerve cells, they help move
materials through the long nerve processes. Microtubules are an
important part of the cytoskeleton in the cytoplasm, and they
are involved in the overall shape changes that cells undergo
during periods of specialization.
Intermediate filaments are a chemically heteroge-
neous group of protein fibers, the specific proteins of which
can vary with cell type (figure 2.20b). These filaments help
maintain cell shape and the spatial organization of organelles,
as well as promote mechanical activities within the cytoplasm.
Microfilaments (actin filaments) are solid strings of
protein (actin) molecules (figure 2.20c). Actin microfilaments
are most highly developed in muscle cells as myofibrils,
which help muscle cells to shorten or contract. Actin micro-
filaments in nonmuscle cells provide mechanical support for
various cellular structures and help form contractile systems
responsible for some cellular movements (e.g., amoeboid
movement in some protozoa).
Cilia and Flagella: Movement

Cilia (sing., cilium; L. eyelashes) and flagella (sing., flagellum;
FIGURE 2.19 L. small whips) are elongated appendages on the surface of
The Cytoskeleton. Model of the cytoskeleton, showing the some cells by which the cells, including many unicellular
three-dimensional arrangement of the microtubules, intermediate organisms, propel themselves. In stationary cells, cilia or fla-
filaments, and microfilaments. gella move material over the cell’s surface.
24 CHAPTER TWO
It has been recently discovered that a cilium may also microtubules and structurally identical to the centriole. The
act as a signal-receiving “antenna” for the cell. In vertebrates, basal body controls the growth of microtubules in cilia or
almost all cells seem to have one per cell. It is the membrane flagella. The microtubules in the basal body form a 9 1 0 pat-
proteins on this single cilium (a primary cilium) that transmit tern: nine sets of three with none in the middle.
molecular signals from the cell’s external environment to its
internal environment (the cytoplasm). When the molecular
signal gets to the cytoplasm, it leads to changes in the cell’s Centrioles and Microtubule-
activities. Cilia-based signaling appears to be a necessity for Organizing Centers
brain functioning and embryonic development.
The specialized nonmembranous regions of cytoplasm near
Although flagella are 5 to 20 times as long as cilia and
the nucleus are the microtubule-organizing centers. These
move somewhat differently, cilia and flagella have a similar
centers of dense material give rise to a large number of
structure. Both are membrane-bound cylinders that enclose
microtubules with different functions in the cytoskeleton. For
a matrix. In this matrix is an axoneme or axial filament,
example, one type of center gives rise to the centrioles (see
which consists of nine pairs of microtubules arranged in a
figure 2.2) that lie at right angles to each other. Each centriole
circle around two central tubules ( figure 2.21). This is called
is composed of nine triplet microtubules that radiate from the
a 9 1 2 pattern of microtubules. Each microtubule pair (a
center like the spokes of a wheel. The centrioles are dupli-
doublet) also has pairs of dynein (protein) arms projecting
cated before cell division, are involved with chromosome
toward a neighboring doublet and spokes extending toward
movement, and help organize the cytoskeleton.
the central pair of microtubules. Cilia and flagella move as a
result of the microtubule doublets sliding along one another.
In the cytoplasm at the base of each cilium or flagel- Vacuoles: Cell Maintenance
lum lies a short, cylindrical basal body, also made up of
Vacuoles (L. vaccus, empty space) are membranous sacs
that are part of the cytomembrane system. Vacuoles occur
in different shapes and sizes and have various functions.
For example, some freshwater single-celled organisms (e.g.,
protozoa) and sponges have contractile vacuoles that col-
lect water and pump it to the outside to maintain the organ-
ism’s internal environment. Other vacuoles store food.
Vaults: Mysterious Symmetrical Shells

Vaults are cytoplasmic ribonucleoproteins shaped like octag-
onal barrels (figure 2.22). Their name is derived from their
multiple arches that look like vaulted cathedral ceilings.
One cell may contain thousands of vaults. The function of
vaults may be related to their octagonal shape. Similarly, the
nuclear pores (see figure 2.23) are also octagonally shaped
and the same size as vaults, leading to speculation that vaults
may be cellular “trucks.” Vaults can dock at nuclear pores,
(a) (b)
FIGURE 2.21
Internal Structure of Cilia and Flagella. In cross section, the FIGURE 2.22
arms extend from each microtubule doublet toward a neighboring Vaults. (a) A three-dimensional drawing of the octagonal
doublet, and spokes extend toward the central paired microtubules. barrel-shaped organelle believed to transport mRNA from the
The dynein arms push against the adjacent microtubule doublet to nucleus to the ribosomes. (b) A vault opened to show its octagonal
bring about movement. structure.
pick up molecules synthesized in the nucleus, and deliver

their load to various places within the cell. Because many
vaults are always located near the nucleus, it is thought that
they are picking up mRNA from the nucleus and transporting
it to the ribosomes for protein synthesis.
Exosome: Signaling Structures

Secreted vesicles known as exosomes were first discovered
nearly 30 years ago. However, they were considered little
more than garbage cans whose job was to discard unwanted
cellular components. Over the past few years, however,
evidence has accumulated that these dumpsters also act as
messengers conveying information to other cells and tissues.
Exosomes contain cell-specific payloads of proteins, lipids,
and genetic material that are transported to other cells and
tissues, where they alter many cellular functions. They are
small in size and are secreted by most cell types and contrib-
ute to functions including tissue repair, neural communica-
tion, and the transfer of pathogenic proteins.
SECTION REVIEW 2.5

Lysosomes function in the digestion of material within the FIGURE 2.23
cell; mitochondria convert energy into a form (ATP) the cell can
The Nuclear Envelope. A color-enhanced electron micrograph
use; ribosomes are the sites for protein synthesis; and vesicles of a section through the nuclear envelope showing nuclear pores
are the site of intracellular digestion, storage, and secretion. (arrows).
The microtubules, intermediate filaments, and microfilaments
make up the “cell skeleton” and function in connecting the
various organelles and cellular components, and also trans- endoplasmic reticulum at a number of points. More than 3,000
port through microtubules. nuclear pores penetrate the surface of the nuclear envelope
(figure 2.23). These pores allow materials to enter and leave
“What is the relationship between nuclear pores,
the nucleus, and they give the nucleus direct contact with the
vaults, ribosomes, and the endoplasmic reticulum?”
endoplasmic reticulum (see figure 2.2). Nuclear pores are not
simply holes in the nuclear envelope; each is composed of
2.6 T HE N UCLEUS: an ordered array of globular and filamentous granules, prob-
ably proteins. The size of the pores prevents DNA from leav-
I NFORMATION C ENTER ing the nucleus but permits RNA to be moved out—possibly
aided by vaults (see figure 2.22).
LEARNING OUTCOME
1. Categorize the functions of the nucleus in terms of the Chromosomes: Genetic Containers
structure of the nucleus.
The nucleoplasm is the inner mass of the nucleus. In a non-
The nucleus (L. kernel or nut) contains the DNA and is the dividing cell, it contains genetic material called chromatin.
control and information center for the eukaryotic cell. It has Chromatin consists of a combination of DNA and protein
two major functions: (1) It directs chemical reactions in cells and is the uncoiled, tangled mass of chromosomes (“col-
by transcribing genetic information from DNA into RNA, ored bodies”) containing the hereditary information in
which then translates this specific information into proteins segments of DNA called genes. During cell division, each
(e.g., enzymes) that determine the cell’s specific activities chromosome coils tightly, which makes the chromosome
(functions). (2) It stores genetic information and transfers this visible when viewed through a light microscope.
information during cell division from one cell to the next, and
from one generation of organisms to the next. Nucleolus: Preassembly Point
for Ribosomes
Nuclear Envelope:
The nucleolus (pl., nucleoli) is a non-membrane-bound struc-
Gateway to the Nucleus ture in the nucleoplasm that is present in nondividing cells
The nuclear envelope is a membrane that separates the (figure 2.24). Two or three nucleoli form in most cells, but
nucleus from the cytoplasm and is continuous with the some cells (e.g., a mphibian eggs) have thousands. Nucleoli
26 CHAPTER TWO
that are functionally differentiated and exhibit a division of

labor. The third level is the tissue level. Jellyfishes have
aggregations of cells organized into definite patterns or lay-
ers, which form a tissue. The fourth level is the organ level.
Organs are composed of one or more tissues and have more
Nucleolus specialized functions than tissues. This level first appears in
the flatworms, where specific structures such as reproduc-
tive organs, eyespots, and feeding structures are present. The
Nuclear fifth and highest level of organization is the system level. At
envelope
this level, organs work together to form systems such as the
Chromatin circulatory, digestive, reproductive, and respiratory systems.
This level first appears in the nemertean worms. Most animal
phyla exhibit this level of organization.
SECTION REVIEW 2.7

FIGURE 2.24 The first level of organization in a higher animal is the protoplas-
Nucleus. The nucleolus, chromatin, and nuclear envelope are mic level, followed by the cellular level, tissue level, and organ
visible in this nucleus (LM 37900). level, and the highest and most complex is the system level.
Do most organs have more than one type of tissue? Explain.
are preassembly points for ribosomes and usually contain

proteins and RNA in many stages of synthesis and assembly. 2.8 T ISSUES
Assembly of ribosomes is completed after they leave the
nucleus through the pores of the nuclear envelope. LEARNING OUTCOMES
1. Explain the structure and function of different epithelia.
2. Identify the different types of connective tissues.
SECTION REVIEW 2.6 3. Identify a unique feature of muscle cells.
The nucleus is surrounded by an envelope of two phos- 4. Describe the basic function of neurons.
pholipid bilayers. The outer layer is continuous with the ER.
Pores allow the passage of small molecules. The nucleolus In an animal, individual cells differentiate during develop-
is where rRNA is transcribed and ribosomes are assembled. ment to perform special functions as aggregates called tis-
Numerous chromosomes are present in eukaryotes. sues. A tissue (Fr. tissu, woven) is a group of similar cells
specialized for the performance of a common function. The
Would you expect the pores in the nuclear envelope to study of tissues is called histology (Gr. histos, tissue 1 logos,
have a function? If so, what is it? discourse). Animal tissues are classified as epithelial, connec-
tive, muscle, or nervous.
2.7 L EVELS OF O RGANIZATION Epithelial Tissue:

IN V ARIOUS A NIMALS Many Forms and Functions
Epithelial tissue exists in many structural forms. In general, it
LEARNING OUTCOME either covers or lines something and typically consists of
1. Describe, from the simplest to the most complex, the renewable sheets of cells that have surface specializations
five levels of organization in a higher animal. adapted for their specific roles. Usually, a basement mem-
brane separates epithelial tissues from underlying, adjacent
Animals exhibit five major levels of organization. Each level tissues. Epithelial tissues absorb (e.g., the lining of the small
is more complex than the one before and builds on it in a intestine), transport (e.g., kidney tubules), excrete (e.g.,
hierarchical manner (see figure 2.1). sweat and endocrine glands), protect (e.g., the skin), and
The first level exhibits protoplasmic organization. contain nerve cells for sensory reception (e.g., the taste buds
This level is found in unicellular organisms such as the pro- in the tongue). The size, shape, and arrangement of epithe-
tozoa where all life functions occur within the boundaries of lial cells are directly related to these specific functions.
a single cell. The second level exhibits cellular organiza- Epithelial tissues are classified on the basis of shape
tion. Flagellates such as Volvox and some sponges can be and the number of layers present. Epithelium can be simple,
placed at this level, where there is an aggregation of cells consisting of only one layer of cells, or stratified, consisting
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were comparatively free, though the difference in the amount of
shelling of the two places was not noticeable.
Of great importance with all gases is the posting of a sufficient
number of sentries around men sleeping within the range of gas
shell. The worst projector gas attack against the Americans was one
where the projectors were landed among a group of dugouts
containing men asleep without sentries. The result was a very heavy
casualty list, coupled with a high death rate, the men being gassed in
their sleep before they were awakened.
Destruction of Mustard Gas

Prior to the introduction of mustard gas all that was necessary to
get rid of gas was to thoroughly ventilate the spot. Thus in trenches
and dugouts, fires were found to be very efficient, simply because
they produced a circulation of air. In the early days, among the
British, the Ayrton fan, a sort of canvas scoop, was used to throw the
gas out of the trenches. While this was taken up in the American
Service, it did not become very important, since it was found that,
under ordinary atmospheric conditions, natural ventilation soon
carried the gas out of the trench proper, while fires in dugouts were
far more efficient than the fans. Likewise the Ayrton fan smacked too
much of trench warfare which had reached a condition of
“stalemate”—a condition that never appealed to the Americans and a
condition that it is hoped never will.
With mustard gas, however, conditions were entirely changed.
This liquid having a very high boiling point and evaporating very
slowly, remains for days in the earth and on vegetation and other
material sprinkled with it. This was particularly true in shell holes
where the force of the explosion drove the gas into the earth around
the broken edges of the hole. While many substances were
experimented with, that which proved best and most practical under
all conditions, was chloride of lime. This was used to sprinkle in shell
holes, on floors of dugouts and any other places where the liquid
might be splashed from bursting shells. It was also found very
desirable to have a small box of this at the entrance to each dugout,
so that a person who had been exposed to mustard gas could
thoroughly coat his shoes with it and thus kill the mustard gas that
collected in the mud on the bottom and sides of his shoes.
Carrying Mustard Gas on Clothing

There are many instances where the occupants of dugouts were
gassed from the gas on the shoes and clothing of men entering the
dugout. Not only were occupants of dugouts thus gassed but a
number of nurses and doctors were gassed while working in closed
rooms over patients suffering from mustard gas poisoning. Even
under the conditions of warfare existing where the Americans were
generally in action, the quantity of chloride of lime required
amounted to several hundred tons per month which had to be
shipped from the United States. Chloride of lime was also very
convenient to have at hand around shell dumps for the purpose of
covering up leaky shells, though rules for handling mustard gas
shells usually prescribed that they be fired and where that was not
practicable to bury them at least five feet under the surface of the
ground. This depth was not so much for the purpose of getting rid of
the gas as it was to get the shell so deep into the ground that it
would not be a danger in any cultivation that might later take place.
Mustard Gas in Cold Weather

Much was learned toward the end of the war about ways of
getting through or around areas infected with mustard gas. For
instance, if mustard gas be fired when the weather is in the
neighborhood of freezing or somewhat below, it will remain on the
ground at night with so little evaporation as not to be dangerous. The
same will be true during the day time if the weather is cloudy as well
as cold. If, however, the days are bright and the nights cold, mustard
gassed areas can be safely crossed by troops at night provided care
is taken in brush and bushes to protect the feet and clothing from the
liquid splashed on bushes. If the sun comes out warm in the morning
such areas may be quite dangerous for three to four hours following
sun-up and indeed for the greater part of the day. Quite a large
number of casualties were ascribed to this fact in the heavy attack
on the British front west of Cambrai just prior to the great German
drive against Amiens, March 21, 1918.
Degassing Units
Since mustard gas has a greatly delayed action it was found that
if men who had been exposed to it could be given a thorough bath
with soap and water within a half hour or even a full hour, the
mustard gas burns would be prevented or very greatly reduced in
severity. Accordingly degassing units were developed consisting
essentially of a 5 ton truck with a 1200 gallon water tank, fitted with
an instantaneous heater and piping to connect it to portable shower
baths. Another truck was kept loaded with extra suits of
underclothing and uniforms. These degassing units were to be
provided at the rate of two per division. Then, in the event of a
mustard gas attack anywhere in the division, one of these units
would be rushed to that vicinity and the men brought out of the line
and given a bath and change of clothing as soon as possible. At the
same time they were given a drink of bicarbonate of soda water and
their eyes, ears, mouth and nasal passages washed with the same.
Protecting Food from Mustard Gas

It was very early learned that mustard gas, or minute particles of
the liquid gas settling on food, caused the stomach to be burned if
the food were eaten, just as the eyes, lungs and skin of the body are
burned from gas in the air. This made it necessary then to see that
all food liable to exposure to mustard gas attacks was protected, and
tarred paper for box linings or tops was found by the Gas Service to
furnish one of the cheapest and most available means of doing this.
Alarm Signals
Numerous, indeed, were the devices invented at one time or
another with which to sound gas alarms. The English early devised
the Strombos horn, a sort of trumpet operated by compressed air
contained in cylinders carried for that purpose. Its note is penetrating
and can be heard, under good conditions, for three or four miles.
When cloud gas attacks, which occurred only at intervals of two to
four months, were the only gas attacks to be feared, it was easy
enough to provide for alarm signals by methods as cumbersome and
as technically delicate as the Strombos horn.
With the advent of shell gas in general, and mustard gas in
particular, the number of gas attacks increased enormously. This
made it not only impossible, but inadvisable also, to furnish sufficient
Strombos horns for all gas alarms, as gas shell attacks are
comparatively local. In such cases, if the Strombos horn is used to
give warning, it causes troops who are long distances out of the area
attacked to take precautions against gas with consequent
interference with their work or fighting.
To meet these local conditions metal shell cases were first hung
up and the alarm sounded on them. Later steel triangles were used
in the same way. At a still later date the large policeman’s rattle, well
known in Europe, was adopted and following that the Klaxon horn.
As the warfare of movement developed the portability of alarm
apparatus became of prime importance. For those reasons the
Klaxon horn and the police rattle were having a race for popularity
when the Armistice was signed.
A recent gas alarm invention that gives promise is a small siren-
like whistle fired into the air like a bomb. It is fitted with a parachute
which keeps it from falling too rapidly when the bomb explodes and
sets it free. Its tone is said to be very penetrating and to be quite
effective over an ample area. Since future gas alarm signals must be
efficient and must be portable, the lighter and more compact they
can be made the better; hence the desirability of parachute whistles
or similar small handy alarms.
Issuing New Masks

One of the problems that remained unsolved at the end of the
war was how to determine when to issue new boxes, or canisters, for
masks. One of the first questions asked by the soldier is how long
his mask is good in gas, and how long when worn in drill where there
is no gas. This information is of course decidedly important.
Obviously, however, it is impossible to tell how long a canister will
last in a gas attack, unless the concentration of gas is known—that
is, the life of the box is longer or shorter as the concentration of gas
is weak or heavy.
A realization of this need led mask designers to work very hard,
long before the necessity for comfort in a mask was as fully realized
as it was at the end of the war, to increase the length of life of the
canister. To get longer life they increased the chemicals and this in
turn increased the breathing resistance, thereby adding to the
discomfort of the soldier when wearing the mask. Finally, however, it
was found that in the concentration of gas encountered on an
average in the field, the life of the comparatively small American
boxes was sufficient to last from fifty to one hundred hours, which is
longer than any gas attack or at least gives time to get out of the
gassed area.
The British early appreciated the necessity of knowing when
boxes should be replaced. They accordingly devised the scheme of
furnishing with each mask a very small booklet tied to the carrying
case in which the soldier could not only enter a complete statement
of the time he had worn the mask but also the statement as to
whether it was in gas or for drill purposes only. The soldier was then
taught that if he had worn the mask, say for forty hours, he should
get a new box. But the scheme didn’t work. In fact, it was one of
those things which foresight might have shown wouldn’t work.
Indeed, any man who in the hell of battle can keep such a record
completely, should be at once awarded a Distinguished Service
Medal.
As gas warfare developed not only were all kinds of gas shells
sent over in a bunch but they were accompanied by high explosive,
shrapnel and anything else in the way of trouble that the enemy
possessed. A man near the front line, under those conditions, had all
he could do and frequently more than he could do, to get his mask
on and keep it on while doing his bit. Consequently he had no time,
even if he had the inclination, to record how long he had the mask in
the various gases.
In this connection, after the Armistice was signed we in the field
were requested to obtain for experimental purposes 10,000 canisters
that had been used in battle. Each was to be labeled with the length
of time it had been worn in or out of gas, and if in gas, the name of
each gas and the time the mask was worn in it. This request is just a
sample of what is asked by those who do not realize field conditions.
One trip to the front would have convinced the one making the
request of the utter impossibility of complying with it, for really no
man knows how long he wears a mask in gas. With gas as common
and as difficult to detect (when intermingled with high explosive
gases and other smells of the battle field) as it was at the end of the
war, each man wore the mask just as long as he could, simply as a
matter of precaution.
Before hostilities ceased we were trying out a method of calling in
say fifty canisters per division once a week for test in the laboratory.
If the tests showed the life of the canisters to be short new canisters
would be issued. While we did not have opportunity to try out this
plan, it gave promise of being the best that could be done. With gas
becoming an every day affair, the only other alternative would seem
to be to make issues of new boxes at stated intervals. On the other
hand there are no definite records of casualties occurring from the
exhaustion of the chemicals in the box. Undoubtedly some did occur,
but they were very, very few. In nearly all cases the masks got
injured, or the box became rusted through before the chemicals gave
out.
Tonnage and Number of Masks Required

It will probably be a shock to most people to learn that with more
than two million men in France we required nearly 1500 tons of gas
material per month. This tonnage was increasing, rather than
decreasing, to cover protective suits, gloves, pastes, and chloride of
lime, as well as masks. The British type respirator was estimated to
last from four to six months. The active part of the war, in which the
Americans took part, was too short to determine whether this was
correct or not. The indications were, however, that it was about right,
considering rest periods and fighting periods.
With the new American mask, with its much stronger and stiffer
face material, the chances are that the life will be considerably
increased although the more constant use of the mask will probably
offset its greater durability. A longer life of mask would of course be a
decided advantage as it would not only reduce tonnage, but would
reduce manufacturing and distribution as well. The estimates on
which we were working at the end looked forward to requiring from
the United States about one-third pound per man per day for all
troops in France, in order to keep them supplied with gas defense
material and with the gases used offensively by gas troops. All gas
shell, hand grenades, etc., used by other than gas troops required
tonnage in addition to the above.
Summing Up
In summing up then, it is noted that there are several important
things in defense against gas. First, the mask which protects the
eyes and the lungs. Second, the training that teaches the man how
to utilize to best advantage the means of protection at his disposal,
whether he be alone or among others. Third, protective clothing that
protects hands and feet and the skin in general. Fourth, a knowledge
of gases and their tactical use that will enable commanders,
whenever possible, to move men out of gas infected areas. Fifth,
training in the offensive use of gas, as well as in defensive methods,
to teach the man that gas has no uncanny power and that it is simply
one element of war that must be reckoned with, thus preventing
stampedes when there is really no danger.
While these are the salient points in defense against gas, above
them and beyond them lies the vigorous offensive use of gas. This
involves not only the research, development and manufacture of
necessary gases in peace time, but also the necessary training to
enable our nation to hurl upon the enemy on the field of battle
chemical warfare materials in quantities he cannot hope to attain.
CHAPTER XXV
PEACE TIME USES OF GAS
“Peace hath her victories no less renowned than war.” Thus runs
the old proverb. In ancient times war profited by peace far more than
peace profited by war if indeed the latter ever actually occurred. The
implements developed for the chase in peace became the weapons
of war. This was true of David’s sling-shot, of the spear and of the
bow. Even powder itself was probably intended and used for scores
of years for celebrations and other peaceful events.
The World War reversed this story, especially in its later phases.
The greater part of the war was fought with implements and
machines prepared in peace either for war or for peaceful purposes.
Such implements were the aeroplane, submarine, truck, automobile
and gasoline motors in general. The first gas attack, which was
simply an adaptation of the peacetime use of the chemical chlorine,
inaugurated the change. Gas was so new and instantly recognized
as so powerful that the best brains in research among all the first
class powers were put to work to develop other gases and other
means of projecting them upon the enemy. The result was that in the
short space of three and one-half years a number of substances
were discovered, or experimented with anew, that are aiding today
and will continue to aid in the future in the peaceful life of every
nation.
Chlorine is even more valuable than ever as a disinfectant and
water purifier. It is the greatest bleaching material in the world, and
has innumerable other uses in the laboratory. Chloropicrin, cyanogen
chloride and cyanogen bromide are found to be very well adapted to
the killing of weevil and other similar insect destroyers of grain.
Hydrocyanic acid gas is the greatest destroyer today of insect pests
that otherwise would ruin the beautiful orange and lemon groves of
California and the South.
Fig. 120.
Phosgene, so extensively used in the war both in cloud gas and

in shell, is finding an ever increasing use in the making of brilliant
dyes—pinks, greens, blues and violets. On account of its cheapness
and simplicity of manufacture, it has great possibilities in the
destruction of rodents such as rats around wharves, warehouses
and similar places that are inaccessible to any other means of
reaching those pests. Since phosgene is highly corrosive of steel,
iron, copper and brass, it cannot be used successfully in places
where those metals are present.
Instead of phosgene for killing rodents and the like in
storehouses and warehouses, cyanogen bromide has been
developed. This is a solid and can be burned like an ordinary sulphur
candle. It is much safer for the purpose of fumigating rooms and
buildings than is hydrocyanic acid gas when so used. This is for the
reason that cyanogen bromide is an excellent lachrymator in
quantities too minute to cause any injury to the lungs. It will thus give
warning to anyone attempting to enter a place where some of the
gas may still linger.
Among tear gases, the new chloracetophenone, a solid, is
perhaps the greatest of all. When driven off by heat it first appears
as a light bluish colored cloud. This cloud is instantly so irritating to
the eyes that within a second anyone in the path of the cloud is
temporarily blinded. It causes considerable smarting and very
profuse tears which even in the smallest amount continue for two to
five minutes. In greater quantities it would continue longer. So far as
can be ascertained, it is absolutely harmless so far as any
permanent injuries are concerned.
Considering that it is instantly effective, that minute quantities are
unbearable to the eyes, that it can be put in hand grenades or other
small containers and driven off by a heating mixture which will not
ignite even a pile of papers, and that it needs no explosion to burst
the grenade (all that is used is a light cap, set off by the action of the
spring, sufficient to ignite the burning charge), the future will see
every police department in the land outfitted with chloracetophenone
or other similar grenades. Every sheriff’s office, every jail and every
penitentiary will have a supply of them. No jail breaking, no lynching,
no rioting can succeed where these grenades are available. Huge
crowds can be set to weeping instantly so that no man can see and
no mob will continue once it is blinded with irritating tears. More than
that, it is an extremely difficult gas to keep out of masks, ordinary
masks of the World War being entirely useless against it.
The same is true of diphenylaminechlorarsine. This is not a tear
gas but it is extraordinarily irritating to the lungs, throat and nose,
where it causes pains and burning sensations, and in higher
concentrations vomiting. It is hardly poisonous at all so that it is
extremely difficult to get enough to cause danger to life. This is
mentioned because of its possible use for the protection of bank
vaults, safes, and strong rooms generally.
There are many other gases that can be used for this same
purpose. It is presumed that gases that are not powerful enough to
kill are the ones desired, and there are half a dozen at least that can
be so used. If desired deadly gases can just as readily be used.
Already a number of inventors are at work on the problem, with
some plans practically completely worked out and models made.
It has been suggested that one of these gases could be used by
trappers in trapping wild animals. Hydrocyanic acid gas may be so
used. It acts quickly and is very rapidly dissipated. An animal
exposed to the fumes would die quickly and the trap be safe to
approach within two minutes after it was sprung. It is said that the
loss from animals working their way out of traps by one means or
another is nearly 20 per cent. More than this, it would meet the
objections of the S. P. C. A. in that the animal would not suffer from
having its limbs torn and lacerated by the trap.
Attempts are being made to attack the locust of the Philippines
and the far west and the boll weevil of the cotton states of the South.
So far these tests have not proven more successful than other
methods, but inasmuch as the number of gases available for trial are
so great and the value of success of so much importance, this
research should be continued on a large scale to definitely determine
whether poisonous gas can be used to eradicate these pests—
especially the boll weevil.
As an interesting application of war materials to peaceful uses,
we may consider the case of cellulose-acetate, known during the war
as “aeroplane dope,” the material used to coat the linen covering
aeroplane wings. With a little further manipulation, this cellulose-
acetate, or aeroplane dope, becomes artificial silk—a silk that today
is generally equal to the best natural silk—and which promises in the
future to become a standard product better in every way than that
from the silk worm.
Fig. 121.
These few examples of the peacetime value of gas are worthy of

thought from another standpoint. Being so valuable, their use in
peace will not be stopped. If they are thus manufactured and used in
peace, they will always be available for use in war, and as the
experience of the World War proved, they certainly will be so used
even should anybody be foolish enough to try to abolish their use. As
for this latter idea, the world might as well recognize at once that
half-way measures in war simply invite disaster.
This chapter would not be complete without a brief statement of
the necessity of a thoroughly developed chemical industry in the
United States as a vital national necessity if the United States is to
have real preparedness for a future struggle. As will be indicated a
little later, no one branch of the chemical industry can be allowed to
go out of existence without endangering some part of the scheme of
preparedness.
Let us consider first the coal tar industry. Coal tar is a by-product
of coke ovens or the manufacture of artificial gas from coal. The coal
tar industry is of the utmost importance because in the coal tars are
the bases of nearly all of the modern dyes, a large percentage of the
modern medicines, most of the modern high explosives, a large
proportion of poisonous gases, modern perfumes, and photographic
materials.
A consideration of these titles alone shows how vital the coal tar
industry is. The coal tar as it comes to us as a by-product is distilled,
giving off at different temperatures a series of compounds called
crudes. Ten of these are of very great importance. The first five are
benzene, toluene, naphthalene, anthracene and phenol (carbolic
acid). The second group comprises xylene, methylanthracene,
cresol, carbazol and phenanthrene.
These, when treated with other chemicals, produce a series of
compounds called intermediates, of which there are some 300 now
known. From these intermediates by different steps are produced
either dyes, high explosives, poisonous gases, pharmaceuticals,
perfumes or photographic materials.
We have all heard that Germany controlled the dye industry of
the world prior to the World War. A little study of the above brief
statement of what is contained in the coal tar industry along with
dyes will show in a measure one of the reasons why Germany felt
that she could win a war against the world. That she came so
desperately close to winning that war is proof of the soundness of
her view.
In many of the processes are needed the heavy chemicals such
as chlorine, sulfuric acid, nitric acid, hydrochloric acid and the like.
The alcohol industry is also of very great importance. Grain alcohol
is used extensively in nearly all research problems and in very great
quantities in many commercial processes such as the manufacture
of artificial silk and for gasoline engines in addition to its use in
compounding medicines. It is of very great importance to the
Chemical Warfare Service in that from grain alcohol is obtained
ethylene gas, one of the three essentials in the manufacture of
mustard gas. While this ethylene may be obtained from many
sources, the most available source, considering ease of
transportation and keeping qualities, is in the form of grain alcohol.
Allied to the chemical industries just mentioned is the nitrate
industry for making nitric acid from the nitrogen of the air. Nitrates
are used in many processes of chemical manufacture and
particularly in those for the production of smokeless powders. The
fertilizer industry is of large importance because it deals with
phosphorus, white phosphorus being not only one of the best smoke
producing materials but a material that is, as stated elsewhere, of
great use against men through its powerful burning qualities.
Another point not mentioned above in connection with these
industries is the training of chemists, chemical engineers and the
building up of plants for the manufacture of chemicals, all of which
are necessary sources of supply for wartime needs. Chemists are
needed in the field, in the laboratory and in manufacturing plants.
The greater their number, the more efficiently can these materials be
handled, and since chemicals as such will probably cause more than
50 per cent of all casualties in future wars, their value is almost
unlimited.
Instead of trying to ameliorate the ravages of war, let us turn
every endeavor towards abolishing all war, remembering that the
most scientific nations should be the most highly civilized, and the
ones most desirous of abolishing war. If those nations will push
every scientific development to the point where by the aid of their
scientific achievements they can overcome any lesser scientific
peoples, the end of war should be in sight.
However, we can never be certain that war is abolished until we
convince at least a majority of the world that war is disastrous to the
conqueror as well as to the conquered, and that any dispute can be
settled peacefully if both parties will meet on the common ground of
justice and a square deal.
CHAPTER XXVI
THE FUTURE OF CHEMICAL WARFARE
The pioneer, no matter what the line of endeavor, encounters

difficulties caused by his fellow-men just in proportion as the thing
pioneered promises results. If the promise be small, the difficulties
usually encountered are only those necessary to make the venture a
success. If, however, the results promise to be great, and especially
if the rewards to the inventor and those working with him promise to
be considerable, the difficulties thrown in the way of the venture
become greater and greater. Indeed whenever great results are
promised, envy is engendered in those in other lines whose
importance may be diminished, or who are so short-sighted as to be
always opposed to progress.
Chemical warfare has had, and is still having, its full share of
these difficulties. From the very day when chlorine, known to the
world as a benign substance highly useful in sanitation, water
purification, gold mining and bleaching was put into use as a
poisonous gas, chemical warfare has loomed larger and larger as a
factor to be considered in all future wars. Chlorine was first used in
the cylinders designed for shipping it. These cylinders were poorly
adapted for warfare, and made methods of preparing gas attacks
extremely laborious, cumbersome and time-consuming.
It was not many months, however, until different gases began to
appear in large quantities in shells and bombs, while the close of the
war, 3½ years later, saw the development of gas in solid form
whereby it could be carried with the utmost safety under all
conditions—a solid which could become dangerous only when the
heating mixture, that freed the gas, was properly ignited.
While some of the chemicals developed for use in war prior to the
Armistice have been made known to the world, a number of others
have not. More than this, every nation of first class importance has
continued to pursue more or less energetically studies into chemical
warfare. These studies will continue, and we must expect that new
gases, new methods of turning them loose, and new tactical uses
will be developed.
Already it is clearly foreseen that these gases will be used by
every branch of the Army and the Navy. While chemicals were not
used by the Air Service in the last war, it was even then realized that
there was no material reason why they should not have been so
used. That they will be used in the future by the Air Service, and
probably on a large scale, is certain. The Navy, too, will use gases,
and probably on a considerable scale. Thus chemical materials as
such become the most universal of all weapons of war.
Some of the poisonous gases are so powerful in minute
quantities and evaporate so slowly that their liberation does not
produce sufficient condensation to cause a cloud. Consequently, we
have gases that cannot be seen. Others form clouds by themselves,
such, for instance, as the toxic smoke candle, where the solid is
driven off by heating, while still others cause clouds of condensed
vapor. This brings the discussion into the realm of ordinary smokes
that have no irritating and no poisonous effects.
These smokes are extremely valuable where the purpose is to
form a screen, whether it be to hide the advance of troops or to cut
off the view of observers. These smokes are equally useful on land
and on sea. So great is the decrease in efficiency of the rifle or
machine gun, and of artillery even when firing at troops that cannot
be seen, that smoke for screening purposes will be used on every
future field of battle. When firing through a screen of smoke, a man
has certainly less than one-quarter the chance to hit his target that
he would have were the target in plain view. Since smoke clouds
may or may not be poisonous and since smoke will be used in every
battle, there is opened up an unlimited field for the exercise of
ingenuity in making these smoke clouds poisonous or non-poisonous
at will. It also opens up an unlimited field for the well-trained
chemical warfare officer who can tell in any smoke cloud whether
gas be present and whether, if present, it is in sufficient
concentration to be dangerous.
At the risk of repetition, it is again stated that there is no gas that
will kill or even permanently injure in any quantity that cannot be
detected. For every gas, there is a certain minimum amount in each
cubic foot of air that is necessary to cause any injury. In nearly all
gases, this minimum amount is sufficient to be readily noticeable by
a trained chemical warfare officer through the sense of smell.
It would be idle to attempt to enumerate the ways and means by
which chemicals will be used in the future. In fact, one can hardly
conceive of a situation where gas or smoke will not be employed, for
these materials may be liquids or solids that either automatically,
upon exposure to the air, turn into gas, or which are pulverized by
high explosive, or driven off by heat. This varied character of the
materials enables them to be used in every sort of artillery shell,
bomb or other container carried to the field of battle.
Some of the gases are extremely powerful as irritants to the nose
and throat in very minute quantities, while at the same time being
highly poisonous in high concentrations. Diphenylchloroarsine, used
extensively by the Germans in high explosive shell, is more
poisonous than phosgene, the most deadly gas in general use in the
past war. In addition, it has the quality of causing an intolerable
burning sensation in the nose, throat, and lungs, in extremely minute
quantities. This material can be kept out of masks only by filters,
whereas true gases are taken out by charcoal and chemical
granules.
There is still another quality which helps make chemical warfare
the most powerful weapon of war. Gas is the only substance used in
war which can be counted on to do its work as efficiently at night as
in the daytime. Indeed, it is often more effective at night than in the
daytime, because the man who goes to sleep without his mask on,
who is careless, who loses his mask, or who becomes excited in the
darkness of night, becomes a casualty, and the past war showed
that these casualties were decidedly numerous even when the
troops knew almost to the minute the time the gas would arrive.
Accordingly, chemical warfare is an agency that must not only be
reckoned with by every civilized nation in the future, but is one which
civilized nations should not hesitate to use. When properly safe-
guarded with masks and other safety devices, it gives to the most
scientific and most ingenious people a great advantage over the less
scientific and less ingenious. Then why should the United States or
any other highly civilized country consider giving up chemical
warfare? To say that its use against savages is not a fair method of
fighting, because the savages are not equipped with it, is arrant
nonsense. No nation considers such things today. If they had, our
American troops, when fighting the Moros in the Philippine Islands,
would have had to wear the breechclout and use only swords and
spears.
Notwithstanding the opposition of certain people who, through
ignorance or for other reasons, have fought it, chemical warfare has
come to stay, and just in proportion as the United States gives
chemical warfare its proper place in its military establishment, just in
that proportion will the United States be ready to meet any or all
comers in the future, for the United States has incomparable
resources in the shape of the crude materials—power, salt, sulfur
and the like—that are necessary in the manufacture of gases.
If, then, there be developed industries for manufacturing these
gases in time of war, and if the training of the army in chemical
warfare be thorough and extensive, the United States will have more
than an equal chance with any other nation or combination of nations
in any future war.
It is just as sportsman-like to fight with chemical warfare materials
as it is to fight with machine guns. The enemy will know more or less
accurately our chemical warfare materials and our methods, and we
will have the same information about the enemy. It is thus a
matching of wits just as much as in the days when the Knights of the
Round Table fought with swords or with spears on horseback. The
American is a pure sportsman and asks odds of no man. He does
ask, though, that he be given a square deal. He is unwilling to agree
not to use a powerful weapon of war when he knows that an outlaw
nation would use it against him if that outlaw nation could achieve

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Zoology 10th Edition Stephen A.

This book is printed on acid-free paper.

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Library of Congress Cataloging-in-Publication Data

Preface ix 16 Ambulacraria: Echinoderms and Hemichordates 301

The Exoskeleton 257 CHAPTER 18

CHAPTER 22 Endocrine Systems of Vertebrates Other Than

Control of Water and Solutes (Osmoregulation Sexual Reproduction in Vertebrates 580

REPRODUCTION AND DEVELOPMENT 576 Glossary Online

CONTENT AND ORGANIZATION PEDAGOGY

An Evolutionary The Process of Science

TEACHING AND LEARNING RESOURCES

McGraw-Hill Connect® provides online presentation, assignment, and assessment solutions. It

Connect® Insight is a powerful data

Help Your Students by Making Assignments—

LearnSmart Advantage® is a new series of adaptive learning products including

Powered by a diagnostic and adaptive engine, SmartBook® facilitates the

The reports in SmartBook help

MH Campus® integrates all of your digital products

Tegrity Campus is a fully automated lecture cap-

GENERAL ZOOLOGY updates in animal taxonomy and evolution. The pedagogy

Anatomy Study of the structure of entire organisms

Lake Kivu Kenya

How Do We Know about Genetic

Muscidae Musca Musca

Tabanidae Tabanus Tabanus

Osteichthyes Perciformes Cichlidae Perissodus Perissodus

Mammalia Primates Hominidae Homo Homo

expected to continue in the twenty-first century. Virtu- T a b l e 1 . 3

S UMMARY 2. A change in the genetic makeup of populations of organisms

A NALYSIS AND A PPLICATION

This photomicrograph is a longitudinal section through skeletal muscle tissue,

Animal TABLE 2.1

Polyribosome FIGURE 2.3

The following are important points of the fluid-mosaic Hydrophilic (polar)

How Do Zoologists Investigate the Inner

T he small size of cells is the

SECTION REVIEW 2.3

Functions of Cell Membranes 2.4 M OVEMENT ACROSS

Cell membranes (1) regulate material moving into and out of

Simple Diffusion Facilitated Diffusion

TYPE OF MOVEMENT DESCRIPTION EXAMPLE IN THE BODY OF A FROG

(a) (b) (c) (d)

FIGURE 2.8 FIGURE 2.9

Selectively permeable Water molecule Isotonic Hypertonic Hypotonic

Normal cells Shriveled cells Cells swell and

solution, the solute concentration is the same inside and outside

called organelles. Organelles effectively compartmentalize

Ribosomes: Protein Workbenches

COMPONENT STRUCTURE/DESCRIPTION FUNCTION

Lysosome engulfing Lysosomes digesting

Rough ER Golgi apparatus Smooth ER Food particle

hydrogen peroxide. (If these oxidative enzymes were not

Cilia and Flagella: Movement

Vaults: Mysterious Symmetrical Shells

pick up molecules synthesized in the nucleus, and deliver

Exosome: Signaling Structures

SECTION REVIEW 2.5

that are functionally differentiated and exhibit a division of

SECTION REVIEW 2.7

are preassembly points for ribosomes and usually contain

2.7 L EVELS OF O RGANIZATION Epithelial Tissue:

Destruction of Mustard Gas

Preface ix 16 Ambulacraria: Echinoderms and Hemichordates 301