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Mader) Sylvia S. Mader

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 Sylvia S. Mader
Michael Windelspecht
Appalachian State University

With contributions by
Jason Carlson
St. Cloud Technical and Community College
David Cox
Lincoln Land Community College
Gretel Guest
Durham Technical Community College
Jeffrey Isaacson
Nebraska Wesleyan University
BIOLOGY, TWELFTH EDITION

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Library of Congress Cataloging-in-Publication Data

Mader, Sylvia S.
Biology / Sylvia S. Mader, Michael Windelspecht, Appalachian State University ; with contributions by April Cognato. — Twelfth edition.
pages cm
I. Windelspecht, Michael, 1963- II. Cognato, April. III. Title.
QH308.2.M23 2016
570--dc23
2014022212

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.

www.mhhe.com
About the Authors

Sylvia S. Mader Sylvia S. Mader has authored several nationally recognized biology texts published by McGraw-Hill.
Educated at Bryn Mawr College, Harvard University, Tufts University, and Nova Southeastern University, she holds degrees in both
Biology and Education. Over the years she has taught at University of Massachusetts, Lowell; Massachusetts Bay Community Col-
lege; Suffolk University; and Nathan Mayhew Seminars. Her ability to reach out to science-shy students led to the writing of her first
text, Inquiry into Life, that is now in its thirteenth edition. Highly acclaimed for her crisp and entertaining writing style, her books
have become models for others who write in the field of biology.
Although her writing schedule is always quite demanding, Dr. Mader enjoys taking time to visit and explore the various
ecosystems of the biosphere. Her several trips to the Florida Everglades and Caribbean coral reefs resulted in talks she has given
to various groups around the country. She has visited the tundra in Alaska, the taiga in the Canadian Rockies, the Sonoran Desert in Arizona, and tropical
rain forests in South America and Australia. A photo safari to the Serengeti in Kenya resulted in a number of photographs for her texts. She was thrilled
to think of walking in Darwin’s steps when she journeyed to the Galápagos Islands with a group of biology educators. Dr. Mader was also a member of a
group of biology educators who traveled to China to meet with their Chinese counterparts and exchange ideas about the teaching of modern-day biology.

Michael Windelspecht As an educator, Dr. Windelspecht has taught introductory biology, genetics, and human
genetics in the online, traditional, and hybrid environments at community colleges, comprehensive universities, and military insti-
tutions. For over a decade he served as the Introductory Biology Coordinator at Appalachian State University where he directed a
program that enrolled over 4,500 students annually. He currently serves as an adjunct professor of biology at ASU where he teaches
nonmajors biology and human genetics in the online and hybrid formats. He was educated at Michigan State University and the
University of South Florida. Dr. Windelspecht is also active in promoting the scientific literacy of secondary school educators.
He has led multiple workshops on integrating water quality research into the science curriculum, and has spent several summers
teaching Pakistani middle school teachers.
As an author, Dr. Windelspecht has published five reference textbooks, and multiple print and online lab manuals. He served as the series editor for
a ten-volume work on the human body. For years Dr. Windelspecht has been active in the development of multimedia resources for the online and hybrid
science classrooms. Along with his wife, Sandra, he owns a multimedia production company, Ricochet Creative Productions, which actively develops and
assesses new technologies for the science classroom.

Contributors
Jason Carlson is a Biology Instructor at St. Cloud Technical and Community College in Minnesota where he teaches introductory biology,
microbiology, nutrition, and human biology. Before entering higher education, he was a middle and high school science teacher with education
from the University of Idaho, Bemidji State University, and St. Cloud State University. In the classroom, he supports a student-driven applied
curriculum with relevant and hands-on research and investigation.

Dave Cox serves as Associate Professor of Biology at Lincoln Land Community College, in Springfield, Illinois. He was educated at Illinois
College and Western Illinois University. As an educator, Professor Cox teaches introductory biology for nonmajors in the traditional classroom
format as well as in a hybrid format. He also teaches biology for majors, and marine biology and biological field studies as study-abroad
courses in Belize. He is the co-owner of Howler Publications, a company that specializes in scientific study abroad courses. Professor Cox
served as a contributor to the fourteenth edition of Inquiry and the thirteenth edition of Human Biology.

Gretel Guest is a Professor of Biology at Durham Technical Community College, in Durham, North Carolina. She has been teaching non-
majors and majors Biology, Microbiology, and Genetics for more than 15 years. Dr. Guest was educated in the field of botany at the University
of Florida, and received her Ph.D. in Plant Sciences from the University of Georgia. She is also a Visiting Scholar at Duke University’s Gradu-
ate School. There she serves the Preparing Future Faculty program by mentoring post-doctoral and graduate students interested in teaching
careers. Dr. Guest was a contributor to the fourth edition of Essentials of Biology.

Jeffrey Isaacson is an Associate Professor of Biology at Nebraska Wesleyan University, where he teaches courses in microbiology, im-
munology, pathophysiology, infectious disease, and senior research. He also serves as the Assistant Provost for Integrative and Experiential
Learning. Dr. Isaacson was educated at Nebraska Wesleyan, Kansas State College of Veterinary Medicine, and Iowa State University. He
worked as a small-animal veterinarian in Nevada and California, and completed a post-doctoral fellowship in the Department of Immunology
at the Mayo Clinic in Minnesota. Dr. Isaacson has been a significant contributor and coauthor for three editions of Inquiry Into Life, for the eleventh edition
of Biology, and is a frequent contributor to McGraw-Hill’s LearnSmart adaptive learning program for several textbooks.

iii
Preface

Goals of the Twelfth Edition

The mission of Dr. Sylvia Mader’s text, Biology, has always been to give students an understanding of biological concepts and a working
knowledge of the scientific process. However, like the world around us, the process of teaching science is changing rapidly. Increasingly,
instructors are being asked to engage their students by making content more relevant, while still providing students with a firm foundation
in those core principles on which biology is founded. These changes are clearly outlined in the AAAS/NSF report, Vision and Change in
Undergraduate Biology Education (2009). The eleventh edition of Biology was one of the first texts to address the principles of Vision
and Change by integrating themes within the text. In this edition we expand on that effort with the development of a number of new
resources and processes.
In addition to the evolution of the introductory biology curriculum, students and instructors are increasingly requesting digital
resources to utilize as learning resources. McGraw-Hill Education has long been an innovator in the development of digital resources,
and the Biology text, and its authors, are at the forefront of the integration of these technologies into the science classroom.
In this edition, the authors focused on the following areas:
1. utilization of the data from the LearnSmart adaptive learning platforms to identify content areas within the text that students
demonstrated difficulty in mastering,
2. further development of the themes that connect the content of the text across multiple chapters,
3. development of a new series of videos and websites to introduce relevancy and engage students in the content,
4. refinement of digital assets to provide a more effective assessment of learning outcomes to enable instructors in the flipped,
online, and hybrid teaching environments.

Relevancy
The use of real world examples to demonstrate the importance • A new website, RicochetScience.com that provides updates
of biology in the lives of students is a key component of Vision on news and stories that are interesting to nonscience majors.
and Change and an effective teaching strategy for introductory The Biology101 project links these resources to the major
biology. The development of relevancy-based resources is a major topics of the text. The site also features videos to assist
focus for the authors of the Mader series of texts. Some examples the students in recognizing the relevancy of what they are
include: learning in the classroom.
• A series of new chapter openers to introduce relevancy to the
chapter. The authors chose topics that would be of interest to
a nonscience major, and represent what would typically be
found on a major news source.
• The development of new relevancy-based videos, BioNow
Sessions, that offer relevant, applied classroom resources
to allow students to feel that they can actually do and learn
biology themselves. For more on these, see page ix.
The Vision and Change document clearly identifies the
need to integrate core concepts throughout the curriculum.
We recognize that scientific literacy is not based upon the
memorization of a series of facts. Instead, learning is based on
establishing associations and links between what, at first glance,
appear to be diverse topics. The main themes we have chosen to
emphasize include:
• Evolution
• Nature of Science
• Biological Systems

iv
 These themes are integrated into all aspects
of the textbook, from the unit learning outcomes
to the theme-based feature readings in the text.
At the start of each chapter, “Following the
Themes” introduces the relationship of the chapter’s content to
each of the themes. At the end of each chapter, “Connecting the
Concepts with the Themes” not only reminds the student of the
relationships between chapter content and the three core themes,
but also acts as a prelude to topics in the next few chapters of
the text. In essence, the themes act as the threads that unite
the concepts throughout the text, enabling the student to see
relationships from the molecular to ecosystem levels of biology.

Evolution Theme
Evolutionary change, along with the mechanism of natural
selection, represents the unifying concept of the biological
sciences. In essence, biological evolution is the thread that
links all life together. Throughout this textbook, feature
readings on this theme both demonstrate the process of
evolution and illustrate how scientists study and measure
evolutionary change. By following this theme through the
book, students develop a better understanding of why
evolution is a dynamic process, and one that has shaped,
and will continue to influence, life on this planet.

Nature of Science Theme

Through the processes of observation, the application of
the scientific method, and the use of both inductive and
deductive reasoning, scientists study life. To develop a
deeper understanding of the biological sciences, students
must appreciate that the study of life is a process, and that
this process has application in their everyday lives. This
theme focuses not only on how biologists do science, but
also on the influences scientific inquiry has on our
understanding of our world.

Biological Systems Theme

From cells to ecosystems, all life is interconnected. Increasingly, scientists
are becoming aware that small changes in the chemical composition of an
ecosystem can have a tremendous influence on the life in that ecosystem.
This theme was chosen to provide a holistic approach to the study of the
life sciences, by demonstrating not only that all life is interconnected, but
also that the principles regulating life at the cellular level play a role in
physiology and ecosystem biology as well.

Assessment To help instructors assess their student’s understanding

of these core concepts, we have designed a series of Connect questions for
each theme in each unit of the text .

v
 Author’s Guide to Using the Textbook
I use LearnSmart Labs to encourage critical thinking, teach
scientific processes, and to integrate lab activities into the
classroom environment.
I encourage my students
to use the Before You
Begin feature to identify
concepts they need to
review before beginning to
read the chapter content.

Pre-Class During Class

Built-in Preview Discussion

and Review Tools Questions
and Activities
Students come to
class prepared Flip class and
engage students

Learning Outcomes at the start of each section provide a

preview of the content to come, while the Check Your Progress
feature at the end of the section helps my students assess how
well they understood the material. The learning outcomes are
the same ones used in Connect, so I can easily assign a quiz to
assess which topics I need to clarify during class.

I use the Questions to Consider at the end of each reading and

Thinking Scientifically questions at the end of the chapter as
the basis for class discussions and active learning exercises.

vi
 Media Study Tools includes a table
that shows students the animations,
videos, and multimedia assets
that are available to further explain
difficult topics. These may be used
as tutorials for the students, and
I may assign the accompanying
Connect activities to gauge whether
my students understand the content.

Post-Class

Assessment and
Integration

Apply content and

build critical thinking

Traditional end-of-chapter summaries and review questions provide students

with an opportunity for low-stakes assessment of their comprehension of the
chapter’s topics.

Features like Following the Themes and Connecting the Concepts with
the Themes help them understand how the main concepts of the chapter
relate to each other, building a deeper understanding of the content.

vii
 Author’s Guide to the Digital Classroom

Using reports from within the LearnSmart

system, especially the Most Missed
Questions report, I am able to identify areas
of content that my students are struggling
with before they enter the lecture.

Pre-Class During Class

LearnSmart & Instructor

SmartBook Resources

Students come to Flip class and

class prepared engage students

I generally assign 20-30 minutes of SmartBook 3-5 days before class. The assignments cover only the
core topics for the upcoming lesson.

viii
 During class I can focus on engaging the students with the
relevancy of the content using the BioNow Sessions videos,
active learning exercises, and animations. Tegrity lecture capture
lets my students review these concepts later.

Using feedback from the

LearnSmart reports, I am
Post-Class able to design Connect
assignments that act as
Connect tutorials that target the
concepts my students are
Apply content and struggling with.
build critical thinking

The Connect reports allow me to assess whether my students The quizzing option within Connect allows me to develop
have met the learning objectives. assessments for any classroom environment.

ix
Engaging Your Students
Today’s science classroom relies heavily on the use of digital assets, including animations
and videos, to engage students and reinforce difficult concepts. Biology 12e includes two
resources specifically designed for the introductory science class to help you achieve
these goals.

BioNow Sessions Videos

A relevant, applied approach
allows your students to feel they
can actually do and learn biology
themselves. While tying directly
to the content of your course, the
videos help students relate their
daily lives to the biology you teach
and then connect what they learn
back to their lives.

Each video provides an engaging

and entertaining story about
applying the science of biology
to a real situation or problem.
Attention is taken to use tools and
techniques that any regular person
could perform, so your students
see the science as something they
could do and understand.
A 27-video series narrated and produced by author Jason Carlson

Our new tutorials were prepared to assist

students in understanding some of the more
difficult topics in biology. Each of the videos
explores a specific figure in the text.

For students, these act as informal office

hours, where they can review the most difficult
concepts in the chapter at a pace which helps
them learn.

Instructors of hybrid and flipped courses will

find these useful as online supplements.

A 36-animation series narrated by Michael Windelspecht and produced by

Ricochet Creative Productions, LLC

x

Theme Evolution
Metagenomics 256
The Anatomy of Speciation 299
Carboniferous Forests 421
Evolutionary History of Maize 427
Survival Mechanisms of Plants 438
Theme Biological Systems
The Impact of Acid Deposition 31
How Cells Talk to One Another 87
Tropical Rain Forest Destruction and Climate Change 122
Moving Genes Between Species: Green Fluorescent
Protein and Cells 219
Same but Not the Same—the Role of Epigenetics 237
African Sleeping Sickness 388
Pathogenic Protists and Climate Change 386
Deadly Fungi 404
Theme Nature of Science
Saturated and Trans Fats in Foods 44
Microscopy Today 60
Enzyme Inhibitors Can Spell Death 107
Fermentation and Food Production 135
The G1 Checkpoint 150
Reproductive and Therapeutic Cloning 157
Meiosis and the Parthenogenic Lizards 171
Hemophilia and the Royal Families of Europe 203
Testing for Genetic Disorders 254
The Tree of Life: 150 Years of Support for the Theory of
Evolution by Natural Selection 272
Inbreeding in Populations 290
Genetic Basis of Beak Shape in Darwin’s Finches 308
DNA Barcoding of Life 342
Flu Viruses 359
DIY Bio 362
Bryophytes—Frozen in Time 418
The Many Uses of Bamboo 446
Plants Can Be Used for Cleaning and Discovery of
Minerals 462
Why So Many Scientists Work with Arabidopsis 488
Vertebrates and Human Medicine 555
A Genomic Comparison of Homo sapiens
and Chimpanzees 568
Readings
The Chemical Ecology of Plants 484
Plants and Their Pollinators 500
Evolution of the Animal Body Plan 518
Sexual Selection in Male Bowerbirds 830
Interactions and Coevolution 865
The Concept of Water Potential 467
Would You Eat Insects? 538
UV Rays: Too Much Exposure or Too Little? 593
AIDS and Opportunistic Infections 633
Drugs of Abuse 700
Preventing Transmission of STDs 787
Preventing and Testing for Birth Defects 805
When a Population Grows Too Large 848
Biomagnification of Mercury 901
Some Major Questions Remaining to Be Answered About
Human Evolution 576
Regenerative Medicine 588
Recent Findings About Preventing Cardiovascular
Disease 612
How Horseshoe Crabs Save Human Lives 617
Cancer Vaccines: Becoming a Reality 635
Should You Go Gluten-Free? 650
New Approaches to Treating Obesity 655
Artificial Lung Technology 671
Is “Vaping” Safer Than Smoking? 673
The Misuse of Erythropoietin in Sports 686
An Accidental Experimental Model for Parkinson
Disease 706
Artificial Retinas Come into Focus 723
The Accidental Discovery of Botox 744
Identifying Insulin as a Chemical Messenger 764
Preimplantation Genetic Diagnosis 784
Do Animals Have Emotions? 825
Island Biogeography Pertains to Biodiversity 860
Global Climate Change 877
Wildlife Conservation and DNA 887
Overexploitation of Asian Turtles 915
Emiquon Floodplain Restoration 919
xi
Overview of Content Changes
to Biology, Twelfth Edition
Chapter 1: A View of Life has been reorganized to provide a
briefer overview of biology as a science. The content on the scien-
tific process (section 1.3) has been reworked with new examples
and a new section (1.4) has been added that explores some of the
major challenges facing science.
Unit 1: The Cell
Chapter 2: Basic Chemistry starts with new content on the search
for life on Mars. Chapter 3: The Chemistry of Organic ­Molecules
opens with a look at trans fats in common foods. ­Chapter 5:
­Membrane Structure and Function begins with a new open-
ing article on chili peppers and calcium channels. C ­ hapter 6:
Metabolism: Energy and Enzymes includes new material on
the function of ATP in cells. The content on redox reactions now
focuses more on the processes of photosynthesis and cellular res-
piration. Chapter 7: Photosynthesis begins with new content on
biofuels.
Unit 2: Genetic Basis of Life
Chapter 9: The Cell Cycle and Cellular Reproduction now
contains information on the structure of a eukaryotic chromosome
(section 9.2). Chapter 10: Meiosis and Sexual Reproduction
starts with a new essay on the importance of meiosis and a new
featured reading, “Meiosis and the Parthenogenic Lizards.” The
figure comparing meiosis I and II (Fig. 10.5) has been reworked to
provide an easier comparison of the two processes. Chapter 11:
Mendelian Patterns of Inheritance begins with a new essay on
PKU sensitivity. A new featured reading examines hemophilia
and the royal families of Europe. The content on polygenic in-
heritance now contains references to the genetics of skin coloring
(Fig. 11.17). Chapter 12: Molecular Biology of the Gene has a
new chapter opener that explains the genetic basis of skin, hair,
and eye coloration. The content on the eukaryotic chromosome
has been moved to chapter 9. There is a new illustration on semi-
conservative replication (Fig 12.6) and a new featured reading
that examines GFP protein. Chapter 13: Regulation of Gene
­Expression starts with new content on how gene regulation relates
to the physiology and behavior of primates. A new featured reading,
“Same but Not the Same—the Role of Epigenetics,” has been in-
cluded on epigenetic inheritance. Chapter 14: Biotechnology and
Genomics opens with a new essay on how biotechnology is being
used to treat dental disease. New illustrations on the PCR reaction
(Fig. 14.3) and the nature of transposons (Fig. 14.10) are included.
Unit 3: Evolution
Chapter 16: How Populations Evolve opens with an essay on
MRSA evolution. Chapter 17: Speciation and Macroevolution
contains an updated illustration on allopatric speciation (Fig. 17.8).
xii
Unit 4: Microbiology and Evolution
Chapter 20: Viruses, Bacteria, and Archaea contains a new fea-
tured reading, “DIY Bio,” that examines synthetic biology. A new
illustration (Fig. 20.8) on gram staining is included. Chapter 21:
Protist Evolution and Diversity has been restructured to give
more emphasis on the supergroup classification system. The chap-
ter begins with a new opener on Naegleria fowleri, and contains a
new featured reading on pathogenic protists and climate change.
The evolutionary relationships in Chapter 22: Fungi Evolution
and Diversity now includes the microsporidia (Fig. 22.1).
Unit 5: Plant Evolution and Biology
Chapter 23: Plant Evolution and Diversity contains a new
featured reading, “Bryophytes—Frozen in Time.” Chapter 24:
Flowering Plants: Structure and Organization begins with new
content on the importance of the neem tree. The chapter has been
reorganized to start with content on plant cells and tissues be-
fore exploring organ systems. Chapter 25: Flowering Plants:
­Nutrition and Transport now contains information on hydro­
ponics (Fig. 25.2) and effects of nutrient deficiencies on plants
(Fig. 25.3). Table 26.1 in Chapter 26: Flowering Plants: Control
of Growth Responses now contains the chemical structures of the
plant hormones. A new featured reading explores coevolution and
the chemical ecology of plants. Chapter 27: Flowering Plants:
Reproduction begins with a new essay on the economic impor-
tance of flowers.
Unit 6: Animal Evolution and Diversity
Chapter 28: Invertebrate Evolution begins with new content that
examines the importance of colony-collapse disorder in honeybees.
A new featured reading, “Would You Eat Insects?,” discusses the
potential benefits of using insects as a food source. Chapter 30:
Human Evolution contains a new featured reading that explores
some of the remaining questions on human evolution.
Unit 7: Comparative Animal Biology
Chapter 31: Animal Organization and Homeostasis begins with
an essay that examines the importance of homeostasis for astro-
nauts. Chapter 32: Circulation and Cardiovascular S ­ ystems
opens with material on cardiovascular-related diseases and the
NFL. Chapter 33: The Lymphatic and Immune Systems has
a new opener on foods and anaphylactic shock. Chapter 34:
­Digestive Systems and Nutrition contains a new featured read-
ing on gluten-free diets. Chapter 35: Respiratory Systems has
a new figure (Fig. 35.8) explaining the relationship between air
pressure and volume of a container. The chapter also contains a
new featured reading on the health aspects of using e-cigarettes.
­Chapter 37: Neurons and Nervous Systems begins with new
Overview of Content Changes to Biology, Twelfth Edition xiii

content on Parkinson disease. Chapter 39: Locomotion and Sup-
port Systems starts with a new essay on Olympian Gabby Doug-
las. Chapter 41: Reproductive Systems has a new opener that
explores variations between the sexes in the animal kingdom.
Unit 8: Behavior and Ecology
Chapter 43: Behavioral Ecology starts with content on behav-
ior and communication in honeybees. The featured reading on
animals and emotions has been updated to include recent develop-
ments. Chapter 44: Population Ecology contains a new illustra-
tion on the environmental impact of developed countries. The
predator-prey relationships and content on global climate change
in ­Chapter 45: Community and Ecosystem Ecology has been
updated to include more recent data. Chapter 47: Conservation
of Biodiversity now begins with an essay on the impact of invasive
species.

The Next Generation of Textbook Reviews: Heat Map Technology

The twelfth edition of Biology is the first text-
book in the Mader s­eries which utilized the
data derived from the LearnSmart platform as
a form of review,
The premise is very straightforward. Stu-
dents don’t know what they don’t know—but
LearnSmart does. By compiling data from all
of the probes answered by all of the students,
and then overlaying that data on the text, we
are able to visualize areas of content where
the students are having problems.
The authors were able to use this infor-
mation to not only identify areas of the text
that the students were having problems with,
but also areas that needed additional digital
resources, such as tutorials and new Connect
questions.
Acknowledgments

Dr. Sylvia Mader represents one of the icons of science education. Twelfth Edition Reviewers
Her dedication to her students, coupled to her clear, concise writing
LaQuetta Anderson, Grambling State University
style, has benefited the education of thousands of students over the
Isaac Barjis, City University of New York
past four decades. As an educator, it is an honor to continue her
Gladys Bolding, Georgia Perimeter College
legacy, and to bring her message to the next generation of students.
Bertha M. Byrd, Wayne County Community College District
As always, I had the privilege to work with the phenomenal
Sarah Clark, Howard Community College
team of science educators and coauthors on this edition. They are
Lewis Deaton, University of Louisiana at Lafayette
all dedicated and talented teachers, and their passion is evident in
Angela Edwards, Trident Technical College
the quality of this text. Thank you also to the countless instructors
Salman Elawad, Chattahoochee Valley Community College
who have invited me into their classrooms, both physically and
Victor Fet, Marshall University
virtually, to discuss their needs as instructors and the needs of their
Julie Fischer, Wallace Community College
students. Your energy, and devotion to quality teaching, is what
Monica Frazier, Columbus State University
drives a textbook revision.
Melanie Glasscock, Wallace State Community College
Many dedicated and talented individuals assisted in the de-
George Goff, Wayne County Community College District
velopment of this edition of Biology. I am very grateful for the
Shashuna J. Gray, Germanna Community College
help of so many professionals at McGraw-Hill who were involved
Sylvester Hackworth, Bishop State Community College
in the development of this project. In particular, let me thank my
Cameron Harmon, Fayetteville Technical Community College
product developer, Anne Winch, for not only keeping me on track
Zinat Hassanpour, Cabarrus College of Health Sciences/Rowan
and her valuable advice, but for her endless patience. My editor
Cabarrus Community College
for this text was Chris Loewenberg. From start to finish a project of
Holly Hereau, Macomb Community College
this magnitude can take over 18 months, and Chris has the natural
Dagne Hill, Grambling State University
ability of keeping his authors focused and in reminding me of the
Kimberly Brown, Mississippi Gulf Coast Community College
importance we are making in education. Thanks also to my market-
Ryan Lazik, Pacific College of Oriental Medicine
ing manager, Chris Ho, who offers a unique insight on the needs of
Lynne Lohmeier, Mississippi Gulf Coast Community College
our students. No modern team would be complete without digital
Geralyne Lopez-de-Victoria, Midlands Technical College
support, and for that I thank Eric Weber and Christine Carlson.
Tiffany McFalls-Smith, Elizabethtown Community and
Production of this text was directed by Angela Fitzpatrick
Technical College
and Jayne Klein, who faithfully steered this project through the
Christian Nwamba, Wayne County Community College District
publication process. I was very lucky to have Dawnelle Krouse,
Tom Reeves, Midlands Technical College
Deb Debord, and Rose Kramer as proofreaders and copy editors.
Lyndell Robinson, Lincoln Land Community College
Today’s textbooks are visual productions, and so I need to thank
William Simcik, Lone Star College-Tomball
the creative talents of David Hash. Lori Hancock and Evelyn Jo
Viji Sitther, Morgan State University
Johnson did a superb job of finding just the right photographs
Phillip Snider, Gadsden State Community College
and micrographs. Electronic Publishing Services produced this
Kimberly Sonanstine, Wallace Community College
textbook, emphasizing pedagogy and beauty to arrive at the best
Chris Sorenson, St. Cloud Technical and Community College
presentation on the page.
Salvatore A. Sparace, Clemson University
Who I am, as an educator and an author, is a direct reflection
Marinko Sremac, Mount Wachusett Community College
of what I have learned from my students. Education is a mutualistic
Todd Tolar, Wallace Community College
relationship, and it is my honest opinion that while I am a teacher,
Frances Turner, Howard Community College
both my professional and personal life have been enriched by inter-
Alanna M. Tynes, Lone Star College-Tomball
actions with my students. They have encouraged me to learn more,
Amale Wardani, Lincoln Land Community College
teach better, and never stop questioning the world around me.
Last, but never least, I want to acknowledge my wife, Sandra.
You have never wavered in your support of my projects. Devin and
Kayla, your natural curiosity of the world we live in gives me the
energy to want to make the world a better place.

Michael Windelspecht
Blowing Rock, NC

xiv
 Contents

Preface iv
7 Photosynthesis 114
1 A View of Life 1 7.1
7.2
Photosynthetic Organisms 115
The Process of Photosynthesis 117
1.1 The Characteristics of Life 2
7.3 Plants Convert Solar Energy 119
1.2 Evolution and the Classification of Life 6
7.4 Plants Fix Carbon Dioxide 123
1.3 The Process of Science 9
7.5 Other Types of Photosynthesis 125
1.4 Challenges Facing Science 13
8 Cellular Respiration 129
Unit
1 The Cell 18
8.1
8.2
8.3
Overview of Cellular Respiration 130
Outside the Mitochondria: Glycolysis 132
Outside the Mitochondria: Fermentation 134
2 Basic Chemistry 19 8.4
8.5
Inside the Mitochondria 136
Metabolism 141
2.1 Chemical Elements 20
2.2 Molecules and Compounds 24
2.3
2.4
Chemistry of Water 26
Acids and Bases 30
Unit
2 Genetic Basis of Life 146

3 The Chemistry of Organic Molecules 35

9 The Cell Cycle and Cellular Reproduction 147
3.1 Organic Molecules 36
9.1 The Cell Cycle 148
3.2 Carbohydrates 39
9.2 The Eukaryotic Chromosome 151
3.3 Lipids 42
9.3 Mitosis and Cytokinesis 152
3.4 Proteins 46
9.4 The Cell Cycle and Cancer 158
3.5 Nucleic Acids 50
9.5 Prokaryotic Cell Division 161
4 Cell Structure and Function 57
10 Meiosis and Sexual Reproduction 166
4.1 Cellular Level of Organization 58
10.1 Overview of Meiosis 167
4.2 Prokaryotic Cells 62
10.2 Genetic Variation 169
4.3 Introduction to Eukaryotic Cells 64
10.3 The Phases of Meiosis 172
4.4 The Nucleus and Ribosomes 67
10.4 Meiosis Compared to Mitosis 174
4.5 The Endomembrane System 69
10.5 The Cycle of Life 176
4.6 Microbodies and Vacuoles 72
10.6 Changes in Chromosome Number and Structure 177
4.7 The Energy-Related Organelles 73
4.8 The Cytoskeleton 75
11 Mendelian Patterns of Inheritance 186
5 Membrane Structure and Function 82 11.1 Gregor Mendel 187
11.2 Mendel’s Laws 188
5.1 Plasma Membrane Structure and Function 83
11.3 Mendelian Patterns of Inheritance and Human
5.2 Passive Transport Across a Membrane 88
Disease 194
5.3 Active Transport Across a Membrane 91
11.4 Beyond Mendelian Inheritance 198
5.4 Modification of Cell Surfaces 95

6 Metabolism: Energy and Enzymes 100

12 Molecular Biology of the Gene 207
12.1 The Genetic Material 208
6.1 Cells and the Flow of Energy 101
12.2 Replication of DNA 211
6.2 Metabolic Reactions and Energy Transformations 103
12.3 The Genetic Code of Life 216
6.3 Metabolic Pathways and Enzymes 105
12.4 First Step: Transcription 218
6.4 Oxidation-Reduction Reactions and Metabolism 109
12.5 Second Step: Translation 220

 xv
xviContents

13 Regulation of Gene Expression 228 22 Fungi Evolution and Diversity 395

13.1 Prokaryotic Regulation 229 22.1 Evolution and Characteristics of Fungi 396
13.2 Eukaryotic Regulation 232 22.2 Diversity of Fungi 398
13.3 Gene Mutations 238 22.3 Symbiotic Relationships of Fungi 405

14 Biotechnology and Genomics 244

5 Plant Evolution and Biology

14.1 DNA Cloning 245
Unit
14.2 Biotechnology Products 247 410
14.3 Gene Therapy 250
14.4 Genomics 251
23 Plant Evolution and Diversity 411

3
23.1 Ancestry and Features of Land Plants 412
Unit
Evolution 260 23.2 Evolution of Bryophytes: Colonization of Land 415
23.3 Evolution of Lycophytes: Vascular Tissue 417
23.4 Evolution of Pteridophytes: Megaphylls 419
15 Darwin and Evolution 261 23.5 Evolution of Seed Plants: Full Adaptation
15.1 History of Evolutionary Thought 262 to Land 423
15.2 Darwin’s Theory of Evolution 265
15.3 Evidence for Evolution 270 24 Flowering Plants: Structure
and Organization 435
16 How Populations Evolve 279 24.1 Cells and Tissues of Flowering Plants 436
16.1 Genes, Populations, and Evolution 280 24.2 Organs of Flowering Plants 440
16.2 Natural Selection 286 24.3 Organization and Diversity of Roots 442
16.3 Maintenance of Diversity 290 24.4 Organization and Diversity of Stems 445
17 Speciation and Macroevolution 296 24.5 Organization and Diversity of Leaves 450

17.1 How New Species Evolve 297

17.2 Modes of Speciation 303
25 Flowering Plants: Nutrition and Transport 456
17.3 Principles of Macroevolution 308 25.1 Plant Nutrition and Soil 457
25.2 Water and Mineral Uptake 461
18 Origin and History of Life 317 25.3 Transport Mechanisms in Plants 465
18.1 Origin of Life 318
18.2 History of Life 323 26 Flowering Plants: Control of Growth
18.3 Geological Factors That Influence Evolution 332 Responses 476
26.1 Plant Hormones 477
19 Taxonomy, Systematics, and Phylogeny 337 26.2 Plant Growth and Movement Responses 485
19.1 Systematic Biology 338 26.3 Plant Responses to Phytochrome 489
19.2 The Three-Domain System 341
19.3 Phylogeny 344 27 Flowering Plants: Reproduction 495
27.1 Sexual Reproductive Strategies 496
Unit
4 Microbiology and Evolution 352 27.2
27.3
Seed Development 502
Fruit Types and Seed Dispersal 504
27.4 Asexual Reproductive Strategies 507
20 Viruses, Bacteria, and Archaea 353
20.1 Viruses, Viroids, and Prions 354
20.2
20.3
20.4
The Prokaryotes 360
The Bacteria 363
The Archaea 368
Unit
6 Animal Evolution and Diversity 512

21 Protist Evolution and Diversity 373 28 Invertebrate Evolution 513

21.1 General Biology of Protists 374 28.1 Evolution of Animals 514
21.2 Supergroup Archaeplastida 377 28.2 The Simplest Invertebrates 521
21.3 Supergroup Chromalveolata 380 28.3 Diversity Among the Lophotrochozoans 524
21.4 Supergroup Excavata 385 28.4 Diversity of the Ecdysozoans 532
21.5 Supergroups Amoebozoa, Opisthokonta, and Rhizaria 389 28.5 Invertebrate Deuterostomes 539
Contents xvii

29 Vertebrate Evolution 544 37 Neurons and Nervous Systems 691

29.1 The Chordates 545 37.1 Evolution of the Nervous System 692
29.2 The Vertebrates 547 37.2 Nervous Tissue 695
29.3 The Fishes 548 37.3 The Central Nervous System 699
29.4 The Amphibians 551 37.4 The Peripheral Nervous System 707
29.5 The Reptiles 553
29.6 The Mammals 559 38 Sense Organs 714
38.1 Sensory Receptors 715
30 Human Evolution 564 38.2 Chemical Senses 716
30.1 Evolution of Primates 565 38.3 Sense of Vision 718
30.2 Evolution of Humanlike Hominins 569 38.4 Senses of Hearing and Balance 724
30.3 Evolution of Early Genus Homo 572 38.5 Somatic Senses 729
30.4 Evolution of Later Genus Homo 573
39 Locomotion and Support Systems 733
39.1 Diversity of Skeletons 734
Unit
7 Comparative Animal Biology 580
39.2 The Human Skeletal System 736
39.3 The Muscular System 742

31 Animal Organization and Homeostasis 581 40 Hormones and Endocrine Systems 750
31.1 Types of Tissues 582 40.1 Animal Hormones 751
31.2 Organs, Organ Systems, and Body Cavities 589 40.2 Hypothalamus and Pituitary Gland 758
31.3 The Integumentary System 591 40.3 Other Endocrine Glands and Hormones 758
31.4 Homeostasis 594
41 Reproductive Systems 770
32 Circulation and Cardiovascular Systems 600 41.1 How Animals Reproduce 771
32.1 Transport in Invertebrates 601 41.2 Human Male Reproductive System 773
32.2 Transport in Vertebrates 603 41.3 Human Female Reproductive System 777
32.3 The Human Cardiovascular System 605 41.4 Control of Human Reproduction 781
32.4 Blood  613 41.5 Sexually Transmitted Diseases 785

33 The Lymphatic and Immune Systems 621 42 Animal Development 793

33.1 Evolution of Immune Systems 622 42.1 Early Developmental Stages 794
33.2 The Lymphatic System 623 42.2 Developmental Processes 798
33.3 Innate Immune Defenses 625 42.3 Human Embryonic and Fetal Development 802
33.4 Adaptive Immune Defenses 628 42.4 The Aging Process 809
33.5 Immune System Disorders
and Adverse Reactions 636

34 Digestive Systems and Nutrition 641

Unit
8 Behavior and Ecology 816
34.1 Digestive Tracts 642
34.2 The Human Digestive System 645 43 Behavioral Ecology 817
34.3 Digestive Enzymes 651 43.1 Inheritance Influences Behavior 818
34.4 Nutrition and Human Health 652 43.2 The Environment Influences Behavior 820
43.3 Animal Communication 824
35 Respiratory Systems 659 43.4 Behaviors That Increase Fitness 828
35.1 Gas-Exchange Surfaces 660
35.2 Breathing and Transport of Gases 665 44 Population Ecology 836
35.3 Respiration and Human Health 669 44.1 Scope of Ecology 837
44.2 Demographics of Populations 838
36 Body Fluid Regulation and Excretory 44.3 Population Growth Models 841
Systems 677 44.4 Regulation of Population Size 844
36.1 Animal Excretory Systems 678 44.5 Life History Patterns 846
36.2 The Human Urinary System 681 44.6 Human Population Growth 849
xviiiContents

45 Community and Ecosystem Ecology 855 Appendices

45.1 Ecology of Communities 856
45.2 Community Development 866
A Answer Key A-1
45.3 Dynamics of an Ecosystem 868 B Tree of Life A-16
46 Major Ecosystems of the Biosphere 881 Glossary G-1
46.1 Climate and the Biosphere 882
46.2 Terrestrial Ecosystems 885 Credits C-1
46.3 Aquatic Ecosystems 895 Index I-1
47 Conservation of Biodiversity 905
47.1 Conservation Biology and Biodiversity 906
47.2 Value of Biodiversity 908
47.3 Causes of Extinction 911
47.4 Conservation Techniques 916
 1
A View of Life

The themes of evolution, the nature of science, and biological

systems are important to understanding biology.

O ur planet is home to a staggering diversity of life. It is estimated that there are over
15 million different species, including our species, Homo sapiens, that inhabit the
globe. Furthermore, life may be found everywhere, from the deepest trenches in
Chapter Outline
1.1 The Characteristics of Life 2
1.2 Evolution and the Classification
the oceans to the tops of the highest mountains. Biology is the area of scientific study
of Life 6
that focuses on understanding all aspects of living organisms. To further our under-
1.3 The Process of Science 9
standing of what it means to be alive, biologists explore life from the molecular level of
the information in our genes to the large-scale ecological interactions of multiple spe- 1.4 Challenges Facing Science 13
cies and their environments.
In this text, we are going to focus on three themes that define these explorations.
The first is evolution—the central theme of biology and the explanation for how life
adapts and changes over time. The second theme is the nature of science. Science is
a process that relies on experimentation and hypothesis testing to validate its findings.
The third theme is biological systems. Throughout this text you will discover that life is
interconnected at many levels, from similarities in our genetic information to the cycling
of nutrients in ecosystems.
As we proceed through this chapter, consider how we as humans are intercon-
nected with other species by these three themes.
As you read through this chapter, think about the following questions:
1. Why is evolution a central theme of the biological sciences?
2. In what ways is life interconnected?
3. How do scientists use the scientific method to study life?

Following the Themes

chapter 1 a view of life

Understanding the scientific process, the theory of evolution, and the interaction
Evolution of biological systems is important in the study of biology.

Scientists make observations, form hypotheses, and conduct experiments in an

Nature of Science attempt to understand the principles of life.

From communities of organisms to individual cells, all life is based on atoms and
Biological Systems molecules.

1
2
1.1 The Characteristics of Life
Learning Outcomes
Upon completion of this section, you should be able to
1. Distinguish among the levels of biological organization.
2. Identify the basic characteristics of life.
Biology is the scientific study of life. Life on Earth takes on a stag-
gering variety of forms, often functioning and behaving in ways
strange to humans. For example, gastric-brooding frogs swallow
their embryos and give birth to them later by throwing them up!
Some species of puffballs, a type of fungus, are capable of produc-
ing trillions of spores when they reproduce. Fetal sand sharks kill
and eat their siblings while still inside their mother. Some Ophrys
orchids look so much like female bees that male bees try to mate
with them. Octopuses and squid have remarkable problem-solving
abilities despite a small brain. Some bacteria live their entire life
in 15 minutes, while bristlecone pine trees outlive 10 generations
of humans. Simply put, from the deepest oceanic trenches to the
upper reaches of the atmosphere, life is plentiful and diverse.
Figure 1.1 illustrates the major groups of living organisms.
From left to right, bacteria are widely distributed, microscopic
organisms with a very simple structure. A Paramecium is an ex-
ample of a microscopic protist. Protists are larger in size and more
complex than bacteria. The other organisms in Figure 1.1 are easily
seen with the naked eye. They can be distinguished by how they
get their food. A morel is a fungus that digests its food externally.
A sunflower is a photosynthetic plant that makes its own food,
and an octopus is an aquatic animal that ingests its food.
Although life is tremendously diverse, it may be defined by
several basic characteristics that are shared by all organisms. Like
nonliving things, organisms are composed of chemical elements.
Also, organisms obey the same laws of chemistry and physics that
govern everything within the universe. The characteristics of life,
however, provide insight into the unique nature of life, and help to
distinguish living organisms from nonliving things.
19,000× 100×
Bacteria Paramecium Morel
Figure 1.1 Diversity of life. Biology is the scientific study of life. This is a sample of the many diverse forms of life that are found on planet Earth.
chapter 1 A View of Life
Life Is Organized
The complex organization of life (Fig. 1.2) begins with atoms, the
basic units of matter. Atoms combine to form small molecules, which
join to form larger molecules within a cell, the smallest, most basic
unit of life. Although a cell is alive, it is made from nonliving mol-
ecules. Some cells, such as single-celled Paramecium, live indepen-
dently. In some cases, single-celled organisms clump together to form
colonies, as does the alga Volvox.
Many living organisms are multicellular, meaning they contain
more than one cell. In multicellular organisms, similar cells com-
bine to form a tissue—for example, the nerve and muscle tissues of
animals. Tissues make up organs, such as the brain or a leaf. Organs
work together to form organ systems; for example, the brain works
with the spinal cord and a network of nerves to form the nervous
system. Organ systems are joined together to form an organism,
such as an elephant.
The levels of biological organization extend beyond the in-
dividual organism. All the members of one species (a group of
similar, interbreeding organisms) in a particular area belong to a
­population. A nearby forest may have a population of gray squir-
rels and a population of white oaks, for example. The populations
of various animals and plants in the forest make up a community.
The community of populations interacts with the physical environ-
ment (water, land, climate) to form an ecosystem. Collectively, all
the Earth’s ecosystems make up the biosphere.
You should recognize from Figure 1.2 that each level of bio-
logical organization builds upon the previous level and is more com-
plex. Moving up the hierarchy, each level acquires new e­ mergent
properties, or new, unique characteristics, that are determined by
the interactions between the individual parts. For example, when
cells are broken down into bits of membrane and liquids, these
parts themselves cannot carry out all the basic characteristics of life.
However, all the levels of biological organization are interconnected
and function as biological systems. For example, a change in carbon
dioxide concentrations (a small molecule) may negatively influence
the operation of organs, organisms, and entire ecosystems. In other
words, life is interconnected at a variety of levels.
Sunflower Octopus
 3

Biosphere
Regions of the Earth’s crust,
waters, and atmosphere inhabited
by living organisms

Ecosystem
A community plus the physical environment

Community
Interacting populations in a particular area

Population
Organisms of the same species
in a particular area

Organism elephant tree

An individual; complex individuals
contain organ systems

Organ System nervous shoot

Composed of several organs system system
working together

Organ the brain leaves

Composed of tissues functioning
together for a specific task

Tissue
A group of cells with a common
structure and function

nervous tissue leaf tissue

Cell nerve cell plant cell

The structural and functional
unit of all living organisms

methane
Molecule
Union of two or more atoms
of the same or different elements

oxygen
Atom
Smallest unit of an element composed of
electrons, protons, and neutrons

Figure 1.2 Levels of biological organization. The basic functional unit of life is the cell, which is built from nonliving molecules and atoms.
4 chapter 1 A View of Life

Life Requires Materials and Energy Solar

Living organisms cannot maintain their organization or carry on energy
life’s activities without an outside source of nutrients and energy
(Fig. 1.3). Food provides nutrients, which are used as building
blocks or for energy. Energy is the capacity to do work, and it Heat
takes work to maintain the organization of the cell and the organ-
ism. When cells use nutrient molecules to make their parts and
products, they carry out a sequence of chemical reactions. The term Producers
metabolism (Gk. meta, “change”) encompasses all the chemical
reactions that occur in a cell.
The ultimate source of energy for nearly all life on Earth is the Heat
sun. Plants and certain other organisms are able to capture solar en-
ergy and carry on photosynthesis, a process that transforms solar
energy into the chemical energy of organic nutrient molecules. All Consumers
life on Earth acquires energy by metabolizing nutrient molecules
made by photosynthesizers. This applies even to plants themselves.

Chemicals
The energy and chemical flow between organisms also de-
fines how an ecosystem functions (Fig. 1.4). Within an ecosystem,

Chemicals
chemical cycling and energy flow begin when producers, such as
grasses, take in solar energy and inorganic nutrients to produce
food (organic nutrients) by photosynthesis. Chemical cycling (aqua
arrows in Fig. 1.4) occurs as chemicals move from one population
to another in a food chain, until death and decomposition allow
inorganic nutrients to be returned to the producers once again. En-
ergy (red arrows), on the other hand, flows from the sun through
Decomposers Heat
plants and the other members of the food chain as they feed on
one another. The energy gradually dissipates and returns to the
atmosphere as heat. Because energy does not cycle, ecosystems
could not stay in existence without solar energy and the ability of Figure 1.4­  Chemical cycling and energy flow in an
photosynthetic organisms to absorb it. ecosystem. In an ecosystem, chemical cycling (aqua arrows)
and energy flow (red arrows) begin when plants use solar energy and
Energy flow and nutrient cycling in an ecosystem climate
inorganic nutrients to produce their own food. Chemicals and energy
largely determine not only where different ecosystems are found are passed from one population to another in a food chain. Eventually,
in the biosphere but also what communities are found in the eco- energy dissipates as heat. With the death and decomposition of
system. For example, deserts exist in areas of minimal rain, while organisms, chemicals are returned to living plants once more.
forests require much rain. The two most biologically diverse

ecosystems—tropical rain forests and coral reefs—occur where

solar energy is most abundant. One example of an ecosystem in
North America is the grasslands, which are inhabited by populations
of rabbits, hawks, and various types of grasses, among many others.
These populations interact with each other by forming food chains
in which one population feeds on another. For example, rabbits feed
on grasses, while hawks feed on rabbits and other organisms.

Living Organisms Maintain Homeostasis

Figure 1.3 Acquiring nutrients and energy.
this bear and the fish, need to acquire energy.
To survive, it is imperative that an organism maintain a state of bio-
logical balance, or homeostasis (Gk. homoios, “like”; stasis, “the
same”). For life to continue, temperature, moisture level, acidity, and
other physiological factors must remain within the tolerance range
of the organism. Homeostasis is maintained by systems that monitor
internal conditions and make routine and necessary adjustments.
Organisms have intricate feedback and control mechanisms
All life, including that do not require any conscious activity. These mechanisms may
be controlled by one or more tissues themselves or by the nervous
CHAPTER 1 A View of Life 5

system. When you are studying and forget to eat lunch, your liver
releases stored sugar to keep blood sugar levels within normal
limits. Many organisms depend on behavior to regulate their in-
ternal environment. In animals, these behaviors are controlled by
the nervous system and are usually not consciously controlled. For
example, a lizard may raise its internal temperature by basking in
the sun, or cool down by moving into the shade.
Living Organisms Respond
Living organisms interact with the environment as well as with
other organisms. Even single-celled organisms can respond to their
environment. In some, the beating of microscopic hairs or, in oth-
ers, the snapping of whiplike tails moves them toward or away
from light or chemicals. Multicellular organisms can manage more
complex responses. A vulture can detect a carcass a kilometer away
and soar toward ­dinner. A monarch butterfly can sense the approach
of fall and begin its flight south, where resources are still abundant.
The ability to respond often results in movement: The leaves
of a land plant turn toward the sun, and animals dart toward
safety. Appropriate responses help ensure the survival of the or-
ganism and allow it to carry on its daily activities. All together,
these activities are termed the behavior of the organism. Organ-
isms display a variety of behaviors as they maintain homeostasis
and search and compete for energy, nutrients, shelter, and mates.
Many organisms display complex communication, hunting, and
defense behaviors.
Mutations help create a staggering diversity of life, even
within a group of otherwise identical organisms. Sometimes, or-
ganisms inherit characteristics that allow them to be more suited to
their way of life.
Living Organisms Have Adaptations
Adaptations are modifications that make organisms better able
to function in a particular environment. For example, penguins
are adapted to an aquatic existence in the Antarctic. An extra
layer of downy feathers is covered by short, thick feathers,
which form a waterproof coat. Layers of blubber also keep the
birds warm in cold water. Most birds have forelimbs propor-
tioned for flying, but penguins have stubby, flattened wings
suitable for swimming. Their feet and tails serve as rudders in
the water, but the flat feet also allow them to walk on land. Pen-
guins also have many behavioral adaptations to living in the Ant-
arctic. Penguins often slide on their bellies across the snow in order
to conserve energy when moving quickly (Fig. 1.5). They carry
their eggs—one or at most two—on their feet, where the eggs are
protected by a pouch of skin. This also allows the birds to huddle
together for warmth while standing erect and incubating the eggs.
From penguins to giant sequoia trees, life on Earth is very
diverse, because over long periods of time, organisms respond
to ever-changing environments by developing new adaptations.

Living Organisms Reproduce and Develop

Life comes only from life. All forms of life have the ability to
reproduce, or make another organism like itself. Bacteria, protists,
and other single-celled organisms simply split in two. In most
multicellular organisms, the reproductive process begins with the
pairing of a sperm from one partner and an egg from the other
partner. The union of sperm and egg, followed by many cell divi-
sions, results in an immature stage, which proceeds through stages
of development, or change, to become an adult.
When living organisms reproduce, their genes, or genetic
instructions, are passed on to the next generation. Random com-
binations of sperm and egg, each of which contains a unique
collection of genes, ensure that the offspring has new and differ-
ent characteristics. An embryo develops into a whale, a yellow
daffodil, or a human because of the specific set of genes it inher-
its from its parents. In all organisms, the genes are made of long
DNA (deoxyribonucleic acid) molecules. DNA provides the
blueprint, or instructions, for the organization and metabolism
of the particular organism. All cells in a multicellular organism
contain the same set of genes, but only certain genes are turned
on in each type of specialized cell. You may notice that not all
members of a species are exactly the same, and that there are
obvious differences between species. These differences are the
result of mutations, or inheritable changes in the genetic infor-
mation. Mutation provides an important source of variation in
the genetic information. However, not all mutations are bad— Figure 1.5 Living organisms have adaptations. Penguins
the observable differences in eye and hair color are examples of have evolved complex behaviors, such as sliding across ice to conserve
mutations. energy, to adapt to their environment.
6 chapter 1 A View of Life

These adaptations are unintentional, but they provide the frame-
work for evolutionary change. Evolution (L. evolutio, “an unroll-
ing”) includes the way in which populations of organisms change
over the course of many generations to become more suited to
their environments. All living organisms have the capacity to
evolve, and the process of evolution constantly reshapes every
species on the planet, potentially providing a way for organisms
to persist, despite a changing environment. We
will take a closer look at this process in the MP3
Life
next section. Characteristics
Check Your Progress 1.1
1. Distinguish between an ecosystem and a population in
the levels of biological organization.
2. List the common characteristics of all living organisms.
3. Explain how adaptations relate to evolutionary change.
1.2 Evolution and the Classification
of Life
Learning Outcomes
Upon completion of this section, you should be able to
1. Explain the relationship between the process of natural
selection and evolutionary change.
2. Distinguish among the three domains of life.
plant species generally produces smooth leaves, but a mutation
occurs that causes one plant to have leaves that are covered with
small extensions, or “hairs.” The plant with hairy leaves has an ad-
vantage, because the deer (the selective agent) prefer to eat smooth
leaves, not hairy leaves. Therefore, the plant with hairy leaves
survives best and produces more seeds than most of its neighbors.
As a result, generations later most plants of this species produce
hairy leaves.
As with this example, Darwin realized that although all in-
dividuals within a population have the potential to reproduce, not
all do so with the same success. Prevention of reproduction can be
the result of a number of factors, including an inability to capture
resources, as when long-necked but not short-necked giraffes can
reach their food source, or an inability to escape being eaten be-
cause long legs, but not short legs, can carry an animal to safety.
Whatever the example, it can be seen that organisms with
advantageous traits can produce more offspring than those that
lack them. In this way, living organisms change over time, and
these changes are passed on from one generation to the next. Over
long periods of time, the introduction of newer, more advantageous
traits into a population may drastically reshape a species. Natural

Despite diversity in form, function, and lifestyle, organisms share Some plants within a population exhibit variation in leaf structure.
the same basic characteristics. As mentioned, they are all com-
posed of cells organized in a similar manner. Their genes are com-
posed of DNA, and they carry out the same metabolic reactions to
acquire energy and maintain their organization. The unity of life
suggests that they are descended from a common ancestor—the
first cell or cells.

Evolution—the Core Concept of Biology

The phrase “common descent with modification” sums up the process Deer prefer a diet of smooth leaves over hairy leaves. Plants with
hairy leaves reproduce more than other plants in the population.
of evolution, because it means that as descent occurs from common
ancestors, so do modifications that cause organisms to be adapted
to their environment. Through many observations and experiments,
Charles Darwin came to the conclusion that natural selection is
the process that makes modification—that is, adaptation—possible.

Natural Selection
During the process of natural selection, some aspect of the environ-
ment selects which traits are more apt to be passed on to the next
generation. The selective agent can be an abiotic agent (part of the Generations later, most plants within the population have hairy
leaves, as smooth leaves are selected against.
physical environment, such as altitude), or it can be a biotic agent
(part of the living environment, such as a deer). Figure 1.6 shows
Figure 1.6 Natural selection. Natural selection selects for or
how the dietary habits of deer might eventually affect the charac- against new traits introduced into a population by mutations. Over many
teristics of the leaves of a particular land plant. generations, selective forces such as competition, predation, and the
Mutations fuel natural selection, because mutation introduces physical environment alter the makeup of a population, favoring those
variations among the members of a population. In Figure 1.6, a more suited to the environment and lifestyle.
CHAPTER 1 A View of Life 7

selection tends to sculpt a species to fit its environment and lifestyle Table 1.1 Levels of Classification
and can create new species from existing ones. The end result is the Category Human Corn
diversity of life classified into the three domains of life (Fig. 1.7).
Domain Eukarya Eukarya
Kingdom Animalia Plantae
Organizing Diversity
  Phylum Chordata Anthophyta
An evolutionary tree is like a family tree. Just as a family tree
   Class Mammalia Monocotyledones
shows how a group of people have descended from one couple, an
evolutionary tree traces the ancestry of life on Earth to a common     Order Primates Commelinales
ancestor (Fig. 1.7). One couple can have diverse children, and      Family Hominidae Poaceae
likewise a population can be a common ancestor to several other       Genus Homo Zea
groups, each adapted to a particular set of environmental condi-        Species* H. sapiens Z. mays
tions. In this way, over time, diverse life-forms have arisen. Evolu-
tion may be considered the unifying concept of biology, because *To specify an organism, you must use the full binomial name, such as Homo
sapiens.
it explains so many aspects of it, including how living organisms
arose from a single ancestor.
Because life is so diverse, it is helpful to group organisms into example, all species in the genus Pisum look pretty much the same—
categories. Taxonomy (Gk. tasso, “arrange”; nomos, “usage”) is that is, like pea plants—but species in the plant kingdom can be quite
the discipline of identifying and grouping organisms according to varied, as is evident when we compare grasses to trees. Species placed
certain rules. Taxonomy makes sense out of the bewildering variety in different domains are the most distantly related.
of life on Earth and is meant to provide valuable insight into evo-
lution. Systematics is the study of the evolutionary relationships Domains
between organisms. As systematists learn more about living organ- Current biochemical evidence suggests that there are three do-
isms, the taxonomy often changes. DNA technology is now widely mains: domain Bacteria, domain Archaea, and domain Eukarya.
used by systematists to revise current information and to discover Figure 1.7 shows how the domains are believed to be related. Both
previously unknown relationships between ­organisms. domain Bacteria and domain Archaea may have evolved from the first
Several of the basic classification categories, or taxa, going common ancestor soon after life began. These two domains contain
from least inclusive to most inclusive, are species, genus, family, the prokaryotes, which lack the membrane-bound nucleus found in
order, class, phylum, kingdom,
and ­domain (Table 1.1). The least
inclusive category, species (L. spe-
cies, “model, kind”), is defined as a
group of interbreeding individuals. BACTERIA
Each successive classification cat-
egory above species contains more
types of organisms than the preced-
ing one. Species placed within one common
ancestor ARCHAEA
genus share many specific char- (first cells)
acteristics and are the most closely
related, while species placed in the
same kingdom share only general
Protists
characteristics with one another. For

Plants
cell with nucleus

Figure 1.7 Evolutionary tree

of life. As existing organisms EUKARYA
change over time, they give rise to Fungi
new species. Evolutionary studies
show that all living organisms arose
from a common ancestor about
4 billion years ago. Domain Archaea
and domain Bacteria include the Animals
prokaryotes. Domain Eukarya includes
both single-celled and multicellular
Past Present
organisms that possess a membrane- Time
bound nucleus.
8 chapter 1 A View of Life

the eukaryotes of domain Eukarya. However, archaea organize their Scientific Name
DNA differently than bacteria, and their cell walls and membranes Biologists use binomial nomenclature to assign each living or-
are chemically more similar to eukaryotes than to ganism a two-part name called a scientific name. For example,
bacteria. So, the conclusion is that eukarya split Animation the scientific name for mistletoe is Phoradendron tomentosum.
Three Domains
off from the archaeal line of descent. The first word is the genus, and the second word is the species
Prokaryotes are structurally simple but metabolically complex. designation (specific epithet) of each species within a genus. The
Archaea (Fig. 1.8) can live in aquatic environments that lack oxy- genus may be abbreviated (e.g., P. tomentosum) and, if the species
gen or are too salty, too hot, or too acidic for most other organisms.
Perhaps these environments are similar to those of the primitive
Earth, and archaea (Gk. archae, “ancient”) are the least evolved
forms of life, as their name implies. Bacteria (Fig. 1.9) are variously Domain Eukarya: Protists
adapted to living almost anywhere—in the water, soil, and atmo-
• Algae, protozoans,
sphere, as well as on our skin and in our mouth and large intestine. slime molds, and
Taxonomists are in the process of deciding how to categorize water molds
archaea and bacteria into kingdoms. Domain Eukarya, on the other • Complex single cell
(sometimes filaments,
hand, contains four major groups of organisms (Fig. 1.10). Protists, colonies, or even
which comprise a number of kingdoms, range from single-celled multicellular)
forms to a few multicellular ones. Some are photosynthesizers, and • Absorb, photosynthesize,
160× or ingest food
some must acquire their food. Common protists include algae, the
protozoans, and the water molds. Figure 1.7 shows that plants, fungi, Paramecium, a single-celled protozoan
and animals most likely evolved from protists. Plants (kingdom Plan-
tae) are multicellular photosynthetic organisms. Example plants in-
clude azaleas, zinnias, and pines. Among the fungi (kingdom Fungi) Domain Eukarya: Kingdom Fungi
are the familiar molds and mushrooms that, along with bacteria, help
decompose dead organisms. Animals (kingdom Animalia) are multi- • Molds, mushrooms, yeasts,
and ringworms
cellular organisms that must ingest and process their food. Aardvarks, • Mostly multicellular filaments
jellyfish, and zebras are representative animals. with specialized, complex cells
• Absorb food

Domain Archaea
Amanita, a mushroom
• Prokaryotic cells
of various shapes
• Adaptations to
extreme environments Domain Eukarya: Kingdom Plantae
• Absorb or
chemosynthesize food
• Certain algae, mosses, ferns,
• Unique chemical
conifers, and flowering plants
characteristics
• Multicellular, usually with
33,200× specialized tissues,
containing complex cells
Sulfolobus, an archaean • Photosynthesize food

Figure 1.8 Domain Archaea.

Phalaenopsis, orchid, a flowering plant

Domain Bacteria Domain Eukarya: Kingdom Animalia

• Prokaryotic cells
• Sponges, worms, insects,
of various shapes
fishes, frogs, turtles,
• Adaptations to
birds, and mammals
all environments
• Multicellular with
• Absorb, photosynthesize,
specialized tissues
or chemosynthesize food
containing complex cells
• Unique chemical
• Ingest food
characteristics
6,600×

Escherichia coli, a bacterium Vulpes, a red fox

Figure 1.9 Domain Bacteria. Figure 1.10 Domain Eukarya.

CHAPTER 1 A View of Life 9

has not been determined, it may simply be indicated with a generic
abbreviation (e.g., Phoradendron sp.). Scientific names are univer-
sally used by biologists to avoid confusion. Common names tend
to overlap and often differ depending on locality and the language
of a particular country. But scientific names are based on Latin, a
universally used language that not too long ago was well known
by most scholars.
Check Your Progress 1.2
1. Explain how natural selection results in new adaptations
within a species.
2. List the levels of taxonomic classification from most
inclusive to least inclusive.
3. Describe the differences that might be used to distinguish
among the various kingdoms of domain Eukarya.
1.3 The Process of Science
Learning Outcomes
Upon completion of this section, you should be able to
1. Identify the components of the scientific method.
2. Distinguish between a theory and a hypothesis.
3. Analyze a scientific experiment and identify the
hypothesis, experiment, control groups, and conclusions.
The process of science pertains to the study of biology. As you can
see from Figure 1.2, the multiple stages of biological organization
mean that life can be studied at a variety of levels. Some biological
disciplines are cytology, the study of cells; anatomy, the study of
structure; physiology, the study of function; botany, the study of
plants; zoology, the study of animals; genetics, the study of hered-
ity; and ecology, the study of the interrelationships between organ-
isms and their environment.
Religion, aesthetics, ethics, and science are all ways in which
human beings seek order in the natural world. The nature of scien-
tific inquiry differs from these other ways of knowing and learning,
because the scientific process uses the scientific method, a stan-
dard series of steps used in gaining new knowledge that is widely
accepted among scientists. The scientific method (Fig. 1.11) acts as
a guideline for scientific studies. Scientists often modify or adapt
the process to suit their particular field of study.
Observation
Scientists believe that nature is orderly and measurable—that natu-
ral laws, such as the law of gravity, do not change with time—and
that a natural event, or phenomenon, can be understood more fully
through observation—a formal way of “seeing what happens.”
Scientists use all of their senses in making observations. The
behavior of chimpanzees can be observed through visual means, the
disposition of a skunk can be observed through olfactory means, and
the warning rattles of a rattlesnake provide auditory information of
imminent danger. Scientists also extend the ability of their senses
by using instruments; for example, the microscope enables us to see
objects that could never be seen by the naked eye. Finally, scientists
may expand their understanding even further by taking advantage
of the knowledge and experiences of other scientists. For instance,
they may look up past studies at the library or on the Internet, or they
may write or speak to others who are researching similar topics.

Hypothesis
Observation After making observations and gathering knowledge about a phe-
nomenon, a scientist uses inductive reasoning to formulate a pos-
sible explanation. Inductive reasoning occurs
Hypothesis 1 whenever a person uses creative thinking to
Potential Hypothesis 2
hypotheses Reject combine isolated facts into a cohesive whole. In
Hypothesis 3 Prediction Experiment hypothesis 1
some cases, chance alone may help a scientist
arrive at an idea.
One famous case pertains to the antibiotic
Remaining Reject penicillin, which was discovered in 1928. While
possible Hypothesis 2 Prediction Experiment
Hypothesis 3 hypothesis 2 examining a petri dish of bacteria that had
hypotheses

Last remaining
possible Hypothesis 3 Figure 1.11 Flow diagram for the scientific
hypothesis Modify hypothesis method. On the basis of new and/or previous
observations, a scientist formulates a hypothesis.
The hypothesis is used to develop predictions to be
Predictions tested by further experiments and/or observations,
and new data either support or do not support the
hypothesis. Following an experiment, a scientist often
Experiment 1 Experiment 2 Experiment 3 Experiment 4 chooses to retest the same hypothesis or to test a
related hypothesis. Conclusions from many different
but related experiments may lead to the development
of a scientific theory. For example, studies pertaining to
Predictions development, anatomy, and fossil remains all support
Conclusion
confirmed
the theory of evolution.
10
become contaminated with the mold Penicillium, Alexander Flem-
ming (1881–1955) observed an area that was free of bacteria. Flem-
ming, an early expert on antibacterial substances, reasoned that the
mold might have been producing an antibacterial compound.
We call such a possible explanation for a natural event a
hypothesis. A hypothesis is not merely a guess; rather, it is an
informed statement that can be tested in a manner suited to the pro-
cesses of science.
All of a scientist’s past experiences, no matter what they might
be, have the potential to influence the formation of a hypothesis.
But a scientist considers only hypotheses that can be tested. Moral
and religious beliefs, while very important in the lives of many
people, differ between cultures and through time and may not be
scientifically testable.
Predictions and Experiments
Scientists often perform an experiment, which is a series of proce-
dures, to test a hypothesis. To determine how to test a hypothesis, a
scientist uses deductive reasoning. Deductive reasoning involves
“if, then” logic. In designing the experiment, the scientist may make
a prediction, or an expected outcome, based on knowledge of the
factors in the experiment.
The manner in which a scientist intends to conduct an ex-
periment is called the experimental design. A good experimental
design ensures that scientists are examining the contribution of a
specific variable, called the experimental variable, to the obser-
vation. The result is termed the responding variable, or dependent
variable, because it is due to the experimental ­variable:
Experimental Variable Responding Variable
(Independent Variable) (Dependent Variable)
Factor of the experiment Result or change that occurs
being tested due to the experimental variable
To ensure that the results will be meaningful, an experiment
contains both test groups and a control group. A test group is ex-
posed to the experimental variable, but the control group is not. If
the control group and test groups show the same results, the experi-
menter knows that the hypothesis predicting a difference between
them is not supported.
Scientists often use model organisms and model systems to
test a hypothesis. Model organisms, such as the fruit fly Drosophila
melanogaster or the mouse Mus musculus, are chosen because they
allow the researcher to control aspects of the experiment, such as age
and genetic background. Cell biologists may use mice for modeling
the effects of a new drug. Like model organisms, model systems
allow the scientist to control specific variables and environmental
conditions in a way that may not be possible in the natural environ-
ment. For example, ecologists may use computer programs to model
how human activities will affect the climate of a specific ecosystem.
While models provide useful information, they do not always answer
the original question completely. For example, medicine that is ef-
fective in mice should ideally be tested in humans, and ecological
experiments that are conducted using computer simulations need to
be verified by field experiments. Biologists, and all other scientists,
chapter 1 A View of Life
continuously design and revise their experiments to better understand
how different factors may influence their original observation.
Presenting and Analyzing the Data
The data, or results, from scientific experiments may be presented
in a variety of formats, including tables and graphs. A graph shows
the relationship between two quantities. In many graphs, the ex-
perimental variable is plotted on the x-axis (horizontal), and the
result is plotted along the y-axis (vertical). Graphs are useful tools
to summarize data in a clear and simplified manner. For example,
the line graph in Figure 1.12 shows the variation in the concentra-
tion of blood cholesterol over a four-week study. The bars above
each data point represent the variation, or standard error, in the
results. The title and labels can assist you in reading a graph;
therefore, when looking at a graph, first check the two axes to
determine what the graph pertains to. By looking at this graph, we
know that the blood cholesterol levels were highest during week 2,
and we can see to what degree the values varied over the course
of the study.
Statistical Data
Most authors who publish research articles use statistics to help
them evaluate their experimental data. In statistics, the standard
error, or standard deviation, tells us how uncertain a particular value
is. Suppose you predict how many hurricanes Florida will have next
year by calculating the average number during the past 10 years.
If the number of hurricanes per year varies widely, your standard
error will be larger than if the number per year is usually about the
same. In other words, the standard error tells you how far off the
average could be. If the average number of hurricanes is four and
the standard error is ± 2, then your prediction of four hurricanes is
between two and six hurricanes. In Figure 1.12, the standard error
is represented by the bars above and below each data point. This
provides a visual indication of the statistical analysis of the data.
Variation in Blood Cholesterol Levels
225
Blood Cholesterol (mg/dL)
standard error
200
y-axis
Data
175
150
Week 1 Week 2 Week 3 Week 4
x-axis
Figure 1.12 Presentation of scientific data. This line graph
shows the variation in the concentration of blood cholesterol over a
four-week study. The bars above each data point represent the variation,
or standard error, in the results.
Another random document with
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upon such terms, conditions, and limitations as to its
internal status as may best subserve the interests of the
United States, and it is not necessary to invest such
territory with the full status of an integral part of the
Union.

"That this is one of the ordinary and necessary sovereign

powers of an independent nation, and nothing in the Federal
Constitution or in the fundamental principles that underlie
our Republic denies to the nation a right to the full exercise
of this usual and common sovereign right.

"That the treaty-making power—the President and the Senate—as

evidenced by the language of the treaty of Paris, did not
intend to make Porto Rico and the Philippine Islands integral
parts of the United States, but intended, in several
particulars, to reserve their final status for adjustment by
Congress, at the same time making peculiar and special
differential provisions for variations and exceptions in
customs and port regulations as to Spain and Spanish goods and
subjects, which are inconsistent with the intention that the
ceded countries became upon the ratification of the treaty a
part of the United States in all respects and in the fullest
sense.

"The Government contends that the term 'foreign countries' in

the act of 1897 is to be regarded as having been understood by
Congress to be subject to the rule of interpretation of the
phrase given by the Supreme Court in the case of Fleming v.
Page, where it was held that under our revenue laws every port
is regarded as a foreign one until expressly established as
domestic under the authority and control of the statutes of
the United States.

"That the clause of the Constitution which declares that

duties, imposts, and excises shall be uniform throughout the
United States does not apply to nor govern these cases,
 because the term 'United States,' as there used, means only
the territory comprised within the several States of the
Union, and was intended only for their benefit and protection,
and not for the benefit or protection of outside territory
belonging to the nation; that in the latter sense duties on
imports from these islands are uniform throughout the United
States, because they are uniformly imposed at every port in
the United States, so that there is no preference given to the
ports of one State over those of another, nor is any
inequality between the several States created.

{671}

"That the right to bring merchandise into the United States is

a right entirely within the regulation of Congress; such right
in no wise differs as to either citizens or aliens.
Citizenship carries with it no special or peculiar privileges
at the custom-house. The American, the Spaniard, the Porto
Rican, are treated alike. The basis of the customs laws is not
ownership, but (1) the geographical origin of the shipment,
and (2) the nature of the goods. The duty is imposed against
merchandise, not upon the importer. "The Government contends,
therefore, that in view of the fact that tariff laws are 'in
rem,' there is no principle of justice, much less of
constitutional restriction, which forbids Congress from taxing
in this way the merchandise of outlying possessions of the
United States when brought into the ports of the Union. That
the limitations of the Constitution as to customs, etc., were
intended to secure equality between the States in the
geographical sense, and not to forbid Congress from exercising
the ordinary sovereign power of taxation as to the products of
other sections of country not included within the geographical
boundaries of the States; for which we rely upon the opinion
of this court in Knowlton v. Moore as decisive and conclusive.

"If the foregoing propositions are sound, then it is

established (1) that the tariff act of 1897 was intended by
Congress to classify as foreign all countries not a part of or
belonging to the United States at the time of its passage, and
the subsequent cession of the Spanish islands to the United
States did not operate to admit imports from those islands
free of duty, under that law; (2) that the tariff act so
construed and enforced violates no constitutional rule of
uniformity.

"And the case of the plaintiffs in error would seem on these

grounds to have no legal foundation.

"The Government might well be content to rest its argument

upon these propositions. But counsel for the plaintiffs in
error, in the court below as well us in this court, have gone
far beyond these limits, and have challenged and denied the
constitutionality of certain provisions of the treaty of
Paris, contending that the cession of Porto Rico and the
Philippine Archipelago effected a complete incorporation of
those countries with the United States, so that they have
become a part of the United States in the fullest and largest
sense, not only internationally, but organically, so
completely, indeed, that no difference or distinction can be
made by law between imports from those countries and imports
from one of the States of the Union.

"They insist that there can be no limited or qualified

acquisition of territory by this nation; that when Porto Rico
was ceded to the United States it came at once under the
obligations of the Constitution and became entitled to the
privileges of the Constitution, its inhabitants citizens of
the United States, and its territory a part of the United
States. They argue, therefore, that the clause of the treaty
which says that 'the civil rights and political status of the
inhabitants shall be determined by the Congress,' in so far as
it is intended to defer the full enjoyment of the rights and
privileges of citizenship under the Constitution until
Congress shall bestow them hereafter upon the inhabitants, is
'ultra vires' and void, or at least superfluous and
ineffective, because the Constitution 'ex proprio vigore'
extends at once, as an automatic operation, to all territory
ceded to this Government, and no treaty or treaty-making power
can hinder or even suspend it. …

"Counsel have been at great pains to prove that the Government

of the United States is one of delegated powers, and that its
powers are not absolute and untrammeled, but subject to
certain limits never and nowhere to be transcended; that the
vague political entity known as The People stands behind the
constituted agencies of government, holding in reserve the
sources of supreme power, capable and ready to alter or
destroy at its pleasure the machinery heretofore set up in its
behalf. They call these doctrines truisms, and so they are.
They do not help us in this case.

"The Government of the United States has been vested not with
all powers but only with certain particular powers. These
particular delegated powers are in some respects limited and
confined in scope and operation, but in other respects they
are entirely unlimited. So that the real and practical
question is whether there is any limitation preventing the
particular thing here complained of.

"It is worth while, in passing, to allude to the undeniable

fact that 'The People' referred to are not the people of the
Territories or of the outlying possessions of the United
States, but the people of the several States, who ordained and
established for themselves and their posterity the Federal
Constitution.

"Counsel confuse ideas when they argue that the contention of

the Government in these cases implies the possession by
Congress of all unlimited and despotic powers in the
government of territory. We mean no more than this court meant
when it said:
"'The power of Congress over the Territories is general and
plenary.

"'Its sovereignty over them is complete.

"'It has full and complete legislative authority over the

people of the Territories and all departments of the
Territorial governments.

"'The people of the United States, as sovereign owners of the

National Territories, have supreme power over them and their
inhabitants.

"'In legislating for the Territories Congress would doubtless

be subject to those fundamental limitations in favor of
personal rights which are formulated in the Constitution and
its amendments, but these limitations would exist rather by
inference and the general spirit of the Constitution than by
any express and direct application of its provisions.'"

In the Supreme Court of the United States,

October Term, 1900, John H. Goetze, Appellant, &c.;
Brief for the United States.

On the 8th of January, 1901, four other causes, involving

substantially the same questions, came before the Supreme
Court, and, by order of the Court, were consolidated, to be
dealt with virtually as one case. The titles of the cases were
respectively as follows:

Elias S. A. Dc Lima et al., plaintiffs in error,

agt. George R. Bidwell;

Samuel B. Downes et al., plaintiffs in error,

agt. George R. Bidwell;
 Henry W. Dooley et al., plaintiffs in error,
agt. the United States;

Carlos Armstrong, appellant,

agt. the United States, and George W. Crossmon et al.,
appellants, agt. the United States.

{672}

For the plaintiffs, in the case of Henry W. Dooley et al., the

Honorable John G. Carlisle made an oral argument, in which he
said: "What is the Constitution? In the first place it is not
only the supreme law of the States composing the union, but
the supreme law of the land; supreme over every branch and
department of the Government; supreme over every one
exercising authority under the Government; supreme over every
other law or order or regulation, and supreme over all the
people, wherever they may be, within its jurisdiction, and
what we claim is, that so long as this Constitution exists
absolute and arbitrary power over the lives, liberties, or
property of the people can be exercised nowhere in this
Republic. It is now argued that it is supreme only within the
boundaries of the several States, unless Congress extends it
to the Territories; that it limits the powers of Congress only
when legislating for the geographical area embraced in the
States; that the inhabitants of the States are the only people
who can, as a matter of right, claim the benefit of its
guarantees and prohibitions for the protection even of those
personal and property rights which have for ages been secured
by the common law of England, and that all other people within
the jurisdiction of the United States are dependent for the
protection of their civil rights substantially upon the will
of Congress. The question whether the Constitution should be
declared to be the supreme law of the whole land, or only the
supreme law of the respective States and their inhabitants or
citizens, was presented in the Federal Convention of 1787, and
was finally disposed of by the adoption of the clause as it
now stands in the Constitution, which declares it to be "the
supreme law of the land.

"In the plan proposed by Mr. Charles Pinckney, of South

Carolina, it was provided that 'all acts made by the
legislature of the United States pursuant to this
Constitution, and all treaties made under the authority of the
United States, shall be the supreme law of the land,' etc. (1
Elliot, page 46). Mr. Patterson's plan proposed 'that all acts
of the United States in Congress assembled made by virtue and
in pursuance of the powers hereby vested in them, and by the
Articles of Confederation and all treaties made and ratified
under the authority of the United States, shall be the supreme
law of the respective States, so far as those acts or treaties
shall relate to such States or their citizens,' etc. (pages
71, 72). These plans and others were referred to the Committee
of the Whole House and were reported back without any
provision upon this subject. Afterwards the Convention
unanimously agreed to the following resolution:

'That the Legislative acts made by virtue and in pursuance of

the Articles of Union and all treaties made and ratified under
the authority of the United States, shall be the supreme law
of the respective States, so far as those acts or treaties
shall relate to the said States or their citizens or
inhabitants' (page 100). Thus it stood when referred to the
committee of five, of which Mr. Rutledge was chairman, and on
the 6th of August, 1787, that committee reported back to the
Convention a draft of the proposed Constitution, the eighth
article of which was the same as the resolution last quoted,
except that in the place of the words 'Articles of Union' it
contained the words 'this Constitution' (page 120). This
report was considered in the Committee of the Whole, and on
the 23d of August the eighth article was unanimously amended
so as to read: 'This Constitution and all laws of the United
States made in pursuance thereof, and all treaties made under
the authority of the United States shall be the supreme law of
the several States and of their citizens and inhabitants,'
etc. (page 151).

"This was the form in which the article stood when the whole
draft was referred to the committee of eleven, but when
reported back September 12, it constituted the second clause
of the sixth article and declared that the Constitution and
laws and the treaties made and to be made should be 'the
supreme law of the land,' and so it now stands as part of the
Constitution. If the clause had been adopted in the form
agreed to in the committee and inserted in the first draft,
there would have been at least a certain degree of
plausibility in the argument made here for the Government, but
even in that case we think the powers of Congress would have
been limited whenever and wherever it might attempt to
exercise them. But it is argued here that the history of the
Constitution and the language employed in the preamble, and in
some other places, show that it was intended to establish a
government only for such of the States then existing as might
ratify it, and such other States as might thereafter be
admitted into the Union, and that, therefore, while it confers
power upon Congress to govern Territories, it does not require
that body to govern them in accordance with the supreme law of
the land; that is, in accordance with the instrument from
which the power to govern is derived. Even if the premises
were true, the conclusion would not follow; but is it true
that the Constitution was ordained and established for the
government of the States only? If so, how did it happen that
the great men who framed that instrument made it confer the
power to govern Territories as well as States? It is true that
the Constitution was ordained and established by the people of
the States, but it created a National Government for national
purposes, not a mere league or compact between the States, and
jurisdiction was conferred upon that Government over the whole
national domain, whatever its boundaries might be. It is not
true that the Government was established only for the States,
their inhabitants or citizens, but if it were true, then it
 could exercise no power outside of the States, and this court
would have to put a new construction upon that provision which
authorizes Congress to dispose of and make all needful rules
and regulations respecting the territory, or other property,
belonging to the United States. The necessary construction of
that clause would be that it conferred power only to dispose
of land or other property, and to make necessary rules and
regulations respecting land or other property belonging to the
United States; that is, belonging to the several States
composing the Union. It would confer no power whatever to
govern the people outside of the States."

Supreme Court of the United States,

October Term, 1900,
Henry W. Dooley [et al.] vs. the United States:
Argument of J. G. Carlisle.

{673}

On one point the argument of Mr. Charles H. Aldrich, attorney

for the plaintiff in the case of the "Fourteen Diamond Rings,"
was as follows: In "the relations of the United States to other
nations, our government is a sovereign state, and has the
right, and as such 'free and independent State has full power,
to levy war, conclude peace, contract alliances, establish
commerce, and to do all other acts and things which
independent States may of right do.' In this relation it is
correct, as I conceive, to speak of the United States of
America as a unit and use a singular verb. It is such unit and
has this power because there was created a government upon
which the people conferred these powers. If war is declared it
must be under the constitution; if peace is concluded it is in
the exercise of a constitutional power; if commerce is
established it is because Congress under the constitution was
given power to regulate commerce; if alliances are contracted
it can only be done under the constitution. In short, the
sovereign nation exists through the adoption of the
constitution, and its powers are derived from that instrument
and must be found, as this court has often declared, in the
language thereof or by necessary implication therefrom. We are
in the Philippines and Porto Rico and can be rightfully there
only in the exercise of some of these enumerated powers, as in
the language of the tenth amendment, 'the powers not delegated
to the United States by the Constitution, nor prohibited by it
to the States, are reserved to the States respectively, or to
the people.' This amendment designates the constitution as the
source of the power of the United States and excludes the idea
of power free from constitutional restraint derived by
implication from powers delegated by the constitution.

"Nor is it true that at the time this declaration was made all
independent states or nations claimed and exercised the right
to acquire, hold and govern foreign dependencies, and no state
or nation then recognized its obligation to confer on the
people of such acquired territory the privileges and
immunities enjoyed by the people of the home government,
except at its own will and discretion. It is true that all
independent states claimed and exercised the right to acquire
territory, but if it were important in this case I think the
arguments of Pitt, Camden and Barre could be used to establish
the proposition that under the British Constitution as it
then was, that nation had, from the time of King John and the
Great Charter until King George, recognized that its subjects
had essential rights not dependent upon the 'will and
discretion' of the home government. It is unnecessary to
follow that subject here. It is sufficient that the
Declaration of Independence was brought about by the assertion
on the part of King George and his ministers of precisely the
present doctrine of this administration and its
representatives in this court. If value is to be attached to
contemporary history that fact cannot be lost sight of. The
speeches of Grenville and Townshend in favor of unlimited
power on the part of Parliament over the American colonists
and their affairs have been substantially parodied in Congress
by the advocates of unrestrained power over our 'colonies,' as
it is now unfortunately fashionable to denominate them. The
signers of the Declaration of Independence held that as
subjects of the British Constitution there was no right to
impose taxes upon them without their consent, to deprive them
of trial by jury, to deprive them of their legislatures, and
to declare Parliament in vested with power to legislate for
them 'in all cases whatsoever.' These and other grievances
were held denials of rights belonging to every British subject
as such and to justify rebellion and war. It seems impossible
that a people who rebelled for such reasons established a
State invested with the very power which they had denied to
the British government and the assertions of which made
rebellion necessary.

"This argument that the power to declare war and conclude

peace carries with it, as an auxiliary, power to do whatever
other nations are accustomed to do with the people and
territory acquired through the exercise of these powers, has a
remarkable likeness to the arguments put forward at the
beginning of the century with reference to the Alien and
Sedition Acts. The supporters of the constitutionality of
these acts claimed that the common law had been introduced and
become a part of the constitution of the United States, and
therefore the powers usually exercisable under the common law
could be exercised by the Congress of the United States in the
respects involved in those acts. Mr. Madison's letter
discussing this contention was answered, so far as it asserted
the right of a state to nullify an act of Congress, but was
never answered, so far as it denied the existence of the
common law as a part of the constitution of the United States.
His objections to that contention, succinctly stated, were,
that if the common law was a part of the constitution, then
there were no constitutional limitations. Congress, like
Parliament, could legislate in all cases whatsoever; that the
President would be possessed of the royal prerogatives (as is
now claimed in this case by the Attorney-General); that the
judiciary would have a discretion little short of legislative
power; that these powers in the different branches of the
government would not be alterable, because, being in the
constitution, they could only be repealed by amendment of that
instrument; and, lastly, that the constitution would have a
different meaning in different States, inasmuch as the common
law was different in such States, and that it would lack the
certainty which a constitution should have, as the common law
was an ever-growing or varying body of law, and, therefore,
with reference to the proper action of the government in each
instance, the question would be important as to what portion
of the common law was in the constitution and what not so
embodied.

"Nearly every sentence of Mr. Madison's able argument with

reference to the common law as a part of the constitution is
applicable to the contention that sovereign powers, so-called,
as derived from or defined by international law, became a part
of the constitution of the United States through the
delegation of the powers to make war, conclude peace, and make
all needful rules and regulations respecting the territory and
other property belonging to the United States. This court has
adopted the view of Mr. Madison. It is hoped that the child of
the old error by which again the executive and legislative
power is sought to be enlarged through the incorporation into
the constitution of 'the sovereign power of other nations'
will receive the same answer.

"In fact, we submit that this court has already held that
sovereign power in the sense that the words are used in the
law of nations as prerogative rights of the King or Emperor,
not only is not vested in the United States or in any branch
of its government, but cannot be so vested. The sovereign
power is with the people. In leaving it with the people our
government marked a departure from all that had previously
existed."
 Supreme Court of the United States,
October Term, 1900, Number 419:
C. H. Aldrich, Argument in reply.

{674}

UNITED STATES OF AMERICA: A. D. 1901.

Military and naval expenditure,
compared with that of other Powers.

See (in this volume)

WAR BUDGETS.

UNITED STATES OF AMERICA: A. D. 1901 (January).

Apportionment of Representatives under the Twelfth Census.
The question of obedience to the Fourteenth Amendment.
Restrictions of the elective franchise in the States.

Section 3 of Article 1 of the Constitution requires that

"Representatives … shall be apportioned among the several
States which may be included within this Union according to
their respective numbers. … The actual enumeration shall be
made within three years after the first meeting of the
Congress of the United States, and within every subsequent
term of ten years. … The number of Representatives shall not
exceed one for every 30,000; but each State shall have at
least one." The first meeting of Congress was in 1789; the
required first census of the United States was taken in 1790,
and, in obedience to the constitutional requirement, the
enumeration has been repeated within the closing year of every
decade since, to supply the basis for a new apportionment of
representatives among the States. The twelfth census, taken in
1900, called for such new distribution, and action upon it was
taken in Congress in January, 1901.

As the section quoted above stood in the Constitution until

1868, it contained a further clause, inserted as one of the
original compromises made between the slaveholding and the
free States, requiring that the determination of numbers to be
represented in the several States should be made "by adding to
the whole number of free persons, including those bound to
service for a term of years, and excluding Indians not taxed,
three-fifths of all other persons." This original clause of
the Constitution was superseded by the Fourteenth Amendment,
adopted in 1868, which introduced this new provision, in its
second section: "Representatives shall be apportioned among
the several States according to their respective numbers,
counting the whole number of persons in each State, except
Indians not taxed. But when the right to vote at any election
for the choice of electors for President and Vice President of
the United States, Representatives in Congress, the executive
and judicial officers of a State, or the members of the
Legislature thereof, is denied to any of the male inhabitants
of such State, being twenty-one years of age, and citizens of
the United States, or in any way abridged, except for
participation in rebellion, or other crime, the basis of
representation therein shall be reduced in the proportion
which the number of such male citizens shall bear to the whole
number of male citizens twenty-one years of age in such
State." To many persons it seemed to be very clear that this
provision of the amended Constitution required account to be
taken of the qualifications by which a number of States have
abridged the suffrage, especially where done for the
understood purpose of disfranchising colored citizens and that
Congress was left with no discretion to do otherwise.

See, (in this volume),

LOUISIANA, NORTH CAROLINA, SOUTH CAROLINA,
MISSISSIPPI, and MARYLAND.

Those holding this view in the House of Representatives gave

support to the following resolution, introduced by Mr.
Olmsted, of Pennsylvania:
"Whereas the continued enjoyment of full representation in
this House by any State which has, for reasons other than
participation in rebellion or other crime, denied to any of
the male inhabitants thereof, being 21 years of age and
citizens of the United States, the right to vote for
Representatives in Congress, Presidential electors, and other
specified officers, is in direct violation of the fourteenth
amendment to the Constitution of the United States, which
declares that in such case 'the basis of representation
therein shall be reduced in the proportion which the number of
such male citizens shall bear to the whole number of male
citizens 21 years of age in such State,' and is an invasion of
the rights and dignity of this House and of its members, and
an infringement upon the rights and privileges in this House
of other States and their representatives; and

"Whereas the States of Massachusetts, Maine, Connecticut,

Delaware, California, Louisiana, Mississippi, North Carolina,
South Carolina, Wyoming, Oregon, and other States do, by the
provisions of the constitutions and statutes of said States,
and for reasons other than participation in rebellion or other
crime, deny the right to vote for members of Congress and
Presidential electors, as well as the executive and judicial
officers of such States and members of the legislatures
thereof, to male inhabitants 21 years of age and over and
citizens of the United States; and such denial in certain of
the said States extends to more than one-half of those who
prior to the last apportionment of representation were
entitled to vote in such States; and

"Whereas in order that the apportionment of membership of the

House of Representatives may be determined in a constitutional
manner: Therefore, be it

"Resolved by the House of Representatives, That the Director

of the Census is hereby directed to furnish this House, at the
earliest possible moment, the following information;
 "First. The total number of male citizens of the United States
over 21 years of age in each of the several States of the
Union.

"Second. The total number of male citizens of the United

States over 21 years of age who, by reason of State
constitutional limitations or State legislation, are denied
the right of suffrage, whether such denial exists on account
of illiteracy, on account of pauperism, on account of
polygamy, or on account of property qualifications, or for any
other reason.

"Resolved further, That the Speaker of the House of

Representatives is hereby authorized and directed to appoint a
select committee of five members from the membership of the
Census Committee of the House of Representatives, who shall
investigate the question of the alleged abridgment of the
elective franchise for any of the causes mentioned in all the
States of the Union in which constitutional or legislative
restrictions on the right of suffrage are claimed to exist,
and that such committee report its findings within twenty days
from the date of the adoption of this resolution to the said
Census Committee, and that within one week after the said
report shall have been received by the Census Committee the
Census Committee shall return a bill to the House of
Representatives providing for the apportionment of the
membership of the House of Representatives based on the
provisions of the fourteenth amendment to the Constitution of
the United States."

{675}

Republicans, hardly less than Democrats, in Congress and

outside, were averse to raising what could not fail to be a
burning sectional issue, and grounds for ignoring the
constitutional mandate were sought with considerable eagerness
on both sides. Strict obedience to the requirement of the
Constitutional provision was claimed to be impracticable, at
least within the time available for proceedings connected with
the present apportionment of representatives. Said one
speaker, opposing the resolutions in the House: "There is not
a State in this Union that has not added to or subtracted from
the Federal constitutional requirements—not one. … If there is
any addition, whether as a matter of police regulation or
otherwise, to the constitutional amendments regulating the
franchise and the resultant representation in this House—if
there is addition or subtraction of one iota—then those who
desire to live up to this Constitution, no matter whether they
ruin their neighbors, no matter whether they again kindle the
fires of sectional strife, those who in their love for the
Constitution are so mentally rigid that they would demand its
enforcement though they set the Union aflame, must include
every State in this Union."

Said another: "How would anybody find out how many people in
the State of Mississippi were disfranchised for the reasons
stated in this resolution? There is there an educational
qualification. How are you to determine how many of the men in
the State of Mississippi who did not vote, did not vote
because they were disfranchised under the educational
qualification? Then there is a qualification in extension and
not in limitation of the suffrage, saying that even those who
can not read and write may still vote, provided they can give
an understanding interpretation of the Constitution or any
part of it. How are you going to determine how many are
disqualified by that? And then there is a qualification which
says that those can not vote who shall not by a certain time
have paid their poll tax. Out of the number of people who did
not vote, how are you going to determine which of them have
not voted because of the educational qualification? Which
because of the understanding qualification? Which because of
the poll-tax qualification? Which because of the registration
qualification? How many because of the pure Australian ballot
which exists in the State of Mississippi? … There is not a
State in the Union which has the Australian ballot which by
the very fact and the necessity of voting according to that
Australian ballot does not prevent the citizen who can not
read and write from voting if he votes a split ticket of any
sort."

A third speaker remarked: "To live up to that amendment, 'that

no male inhabitant shall be deprived of suffrage except for
participation in the rebellion or other crimes,' the male
inhabitant, I take it, is he who has acquired domicile in that
State, and the moment that he acquires domicile, and is a
male, he is a 'male inhabitant' of that State, and entitled
at once to suffrage; and yet every State in the Union, I
believe without exception, has requirements as to residence
not only in the State, but in the city, in the county, in the
precinct and ward and the voting place; and everyone of those
requirements, as every gentleman on that side must admit, are
in direct conflict with and contravention of the fourteenth
amendment to the Constitution of the United States literally
construed."

But the advocates of obedience to the Constitution, supporting

the resolutions of Mr. Olmsted, planted their argument on the
very facts brought against it, as demonstrating the need of
measures to check a growing tendency in the country to
restrict the elective franchise. Said Mr. Shattuck, of Ohio:
"We find that in 1870 there were three States that had
abridged their electorates—California, Connecticut, and
Massachusetts. In these three States there was a
constitutional provision for an educational qualification,
which disfranchised a certain percentage of the
electorate—namely, the illiterates. But, in those States, the
percentage of illiteracy is very light, averaging about 6 per
cent. The basis of representation would hardly have been
affected in those States had the fourteenth amendment been
conformed with.
"An examination into the election laws of the various States
reveals an astonishing tendency at this time to abridge their
electorates. When the Congress which adopted the existing
apportionment discussed the matter ten years ago but three
States had abridged their electorate by action of the State,
and in these the percentage of disfranchised males was but 6
per cent. But since that time similar policies have been
adopted by other States, and to-day we face the fact that ten
of the forty-five States of this Union have abridged their
electorates, and that in these the percentage of males 21
years of age and over, disfranchised, averages over 20 per
cent. The constitutions of several other States permit such an
abridgment. Besides, there are other States preparing to adopt
these policies and to disfranchise thousands of men who to-day
hold the right of franchise. In view of this remarkable
tendency it is inconceivable that Congress can longer permit
the fourteenth amendment to remain a dead letter, and to pass
a bill making an apportionment based solely upon the
population and neglecting the proviso which applies to all
States which have abridged their electorate.

"We will not review the past by any discussion of the question
as to whether the provisions of the fourteenth amendment
should have been made effective when the last apportionment
was made ten years ago. We find to-day conditions existing
which make its enforcement imperative. I do not propose to
discuss at this time whether the reasons given for these
abridgments by the people of the various States are valid or
not. … I am simply pointing out the conditions as they exist;
I am simply pointing out that the time has come when the
tendency of the States to abridge their electorates has grown
to such proportions as to demand that this Congress shall
proceed in a constitutional manner in making the new
apportionment. I do not say that States have not the right to
establish educational qualifications for their electors, but I
do maintain that when they have done so they must pay the penalty
 prescribed in the Constitution, and have their representation
abridged proportionately.
{676}
I do not say that we shall punish only Louisiana;
I do not say that we shall punish only Massachusetts;
I do not say that we shall punish only California;
but I do say and insist, as the representative of a State in
which every male member 21 years of age and over is guaranteed
the sacred right of franchise, that there is a constitutional
remedy prescribed for their acts, and I do demand that that
remedy be applied."

The following interesting table, showing the restrictions of

the electorate in the various States of the Union, was
appended to the remarks of Mr. Shattuck:

STATES.
REQUIREMENTS AS TO CITIZENSHIP. [First paragraph]

PERSONS EXCLUDED FROM SUFFRAGE. [Second paragraph]

ALABAMA.
Citizen of United States, or alien who has declared intention.

Convicted of treason or other crime punishable by

imprisonment, idiots, or insane.

ARKANSAS.
Citizen of United States, or alien who has declared intention.

Idiots, insane, convicted of felony until pardoned, failure

to pay poll tax, United States soldiers on duty in State.

CALIFORNIA.
Citizen by nativity, naturalization, or treaty of Queretaro.

Chinese, insane, embezzlers of public moneys, convicted