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Sylvia S. Mader
Michael Windelspecht

mad87328_fm_i-xvi.indd 1 13/09/19 10:32 PM


Acknowledgments

Dr. Sylvia Mader is one of the icons of science education. Her dedication to her students, coupled to her clear, concise writing
style, has benefited the education of thousands of students over the past four decades. As an educator, it is an honor to continue
her legacy and to bring her message to the next generation of students.
As always, I had the privilege to work with a phenomenal group of people on this edition. I would especially like to thank you,
the numerous instructors who have shared emails with me or have invited me into your classrooms, both physically and virtually,
to discuss your needs as instructors and the needs of your students. You are all dedicated and talented teachers, and your energy
and devotion to quality teaching is what drives a textbook revision.
Many dedicated and talented individuals assisted in the development of this edition of Essentials of Biology. I am very grateful
for the help of so many professionals at McGraw-Hill who were involved in bringing this book to fruition. Therefore, I would like
to thank the following:
∙ My product developer, Anne Winch, for her incredible ability to manage all aspects of this project simultaneously.
∙ My executive portfolio manager, Michelle Vogler, for her guidance and for reminding me why what we do is important.
∙ My marketing manager, Britney Ross, and market development manager, Beth Bettcher, for placing me in contact with great
instructors, on campus and virtually, throughout this process.
∙ My digital expert, Eric Weber, for helping me envision the possibilities in our new digital world.
∙ My content project manager, Kelly Hart, and program manager, Angie FitzPatrick, for guiding this project throughout the
publication process.
∙ Lori Hancock and David Tietz for the photos within this text. Biology is a visual science, and their contributions are evident
on every page.
∙ Michael McGee and Sharon O’Donnell who acted as my proofreaders and copyeditors for this edition.
∙ Jane Peden for her behind-the-scenes work that keeps us all functioning.
As both an educator and an author, communicating the importance of science represents one of my greatest passions. Our modern
society is based largely on advances in science and technology over the past few decades. As I present in this text, there are many
challenges facing humans, and an understanding of how science can help analyze, and offer solutions to, these problems is critical
to our species’ health and survival.
I also want to acknowledge my family and friends for all of their support. My wife, Sandy, who has never wavered in her support of my
many projects. Over the course of my work with McGraw-Hill, I have watched the natural curiosity of my children, Devin and Kayla,
develop them into the phenomenal individuals that they are today. Thank you both for your motivation in making our world a better place.
Michael Windelspecht, PhD
Blowing Rock, NC

Reviewers for This Edition


I would like to thank the individuals below for taking the time to review the content of the previous edition. Your comments and
suggestions played an important part of this revision.

Reviewers of the Sixth Edition


Jennifer Bogdanoff, Coastal Carolina Community College Michele Engel, California State University Bakersfield
Lauri Carey, Illinois Valley Community College Carole S. Griffiths, LIU Brooklyn
Thomas R. Cholmondeley, Cincinnati State Technical and Diane Lewis, Ivy Tech Community College
Community College Craighton S. Mauk, Madisonville Community College (Kentucky
Gregory A. Dahlem, Northern Kentucky University Community and Technical College System)
Joy O. Davis, Baton Rouge Community College Murali Panen, Luzerne County Community College
Dale Emeagwali, Excelsior College Amanda Rodrigues, Johnston Community College
Ray Emmett, Daytona State College Clement Yedjou, Jackson State University

viii

mad87328_fm_i-xvi.indd 8 13/09/19 10:33 PM


Contents
CHAPTER
1 Types of Chemical Bonds 26
Chemical Formulas and Reactions 28

Biology: The Science of Life 1 2.2 Water’s Importance to Life 29


The Structure of Water 29
1.1 The Characteristics of Life 2 Properties of Water 29
Life Requires Materials and Energy 2
Living Organisms Maintain an Internal Environment 4 2.3 Acids and Bases 32
Living Organisms Respond and Develop 5 Acidic Solutions 33
Living Organisms Have Adaptations 5 Basic Solutions 33
pH and the pH Scale 34
1.2 Evolution: The Core Concept of Biology 6 Buffers and pH 35
Natural Selection and Evolutionary Processes 7
Organizing the Diversity of Life 9

1.3 Science: A Way of Knowing 11


CHAPTER
3
Start with an Observation 12 The Organic Molecules of Life 38
Develop a Hypothesis 12
3.1 Organic Molecules 38
Make a Prediction and Perform Experiments 12
The Carbon Atom 38
Develop a Conclusion 13
The Carbon Skeleton and Functional Groups 39
Scientific Theory 14
An Example of a Controlled Study 14 3.2 The Biological Molecules of Cells 40
Publishing the Results 15 Carbohydrates 40
Lipids 43
1.4 Challenges Facing Science 16
Proteins 48
Climate Change 16
Nucleic Acids 52
Biodiversity and Habitat Loss 17

4
Emerging and Reemerging Diseases 18
CHAPTER

Inside the Cell 57


UNIT I The Cell
4.1 Cells Under the Microscope 58

2
The Limit to Cell Size 60
CHAPTER
4.2 The Plasma Membrane 60
The Chemical Basis of Life 21 Functions of Membrane Proteins 62

2.1 Atoms and Atomic Bonds 22 4.3 The Two Main Types of Cells 63
Prokaryotic Cells 63
Atomic Structure 23
The Periodic Table 23 4.4 A Tour of the Eukaryotic Cell 65
Isotopes 24 Nucleus and Ribosomes 67
Arrangement of Electrons in an Atom 25 Endomembrane System 70
Vesicles and Vacuoles 71

ix

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x CONTENTS

Energy-Related Organelles 71 6.4 Variations in Photosynthesis 108


The Cytoskeleton and Motor Proteins 74 C3 Photosynthesis 109
Centrioles 75 C4 Photosynthesis 109
Cilia and Flagella 75 CAM Photosynthesis 110
4.5 Outside the Eukaryotic Cell 76 Evolutionary Trends 110
Cell Walls 76
Extracellular Matrix 76
Junctions Between Cells 76
CHAPTER
7
Energy for Cells 113
CHAPTER
5 7.1 Cellular Respiration 114
Phases of Complete Glucose Breakdown 114

The Dynamic Cell 81 7.2 Outside the Mitochondria: Glycolysis 116


Energy-Investment Step 116
5.1 What Is Energy? 82
Energy-Harvesting Steps 117
Measuring Energy 82
Energy Laws 83 7.3 Outside the Mitochondria: Fermentation 117
Lactic Acid Fermentation 118
5.2 ATP: Energy for Cells 84 Alcohol Fermentation 119
Structure of ATP 84
Use and Production of ATP 85 7.4 Inside the Mitochondria 119
The Flow of Energy 87 Preparatory Reaction 119
The Citric Acid Cycle 120
5.3 Metabolic Pathways and Enzymes 88 The Electron Transport Chain 120
An Enzyme’s Active Site 89
Energy of Activation 90 7.5 Metabolic Fate of Food 123
Energy Yield from Glucose Metabolism 123
5.4 Cell Transport 90 Alternative Metabolic Pathways 124
Passive Transport: No Energy Required 91
Active Transport: Energy Required 94
Bulk Transport 94 UNIT II Genetics

CHAPTER
6
CHAPTER
8
Cellular Reproduction 128
Energy for Life 98
8.1 An Overview of Cellular Reproduction 129
6.1 Overview of Photosynthesis 99 Chromosomes 129
Plants as Photosynthesizers 100 Chromatin to Chromosomes 130
The Photosynthetic Process 101
8.2 The Cell Cycle: Interphase, Mitosis, and
6.2 The Light Reactions—Harvesting Energy 102 Cytokinesis 131
Photosynthetic Pigments 102 Interphase 131
The Light Reactions: Capturing Solar Energy 103 M (Mitotic) Phase 132

6.3 The Calvin Cycle Reactions—Making 8.3 The Cell Cycle Control System 137
Sugars 106 Cell Cycle Checkpoints 137
Overview of the Calvin Cycle 106 Internal and External Signals 138
The Many Uses of G3P 107 Apoptosis 138

mad87328_fm_i-xvi.indd 10 13/09/19 10:33 PM


CONTENTS xi

8.4 The Cell Cycle and Cancer 139 10.3 Beyond Mendel’s Laws 176
Proto-oncogenes and Tumor Incomplete Dominance 176
Suppressor Genes 140 Multiple-Allele Traits 177
Other Genetic Changes and Cancer 141 Polygenic Inheritance 177
8.5 Characteristics of Cancer 143 Gene Interactions 179
Characteristics of Cancer Cells 143 Pleiotropy 180
Cancer Treatment 144 Linkage 180
Prevention of Cancer 145 10.4 Sex-Linked Inheritance 181
Sex-Linked Alleles 182

9
Pedigrees for Sex-Linked Disorders 182
CHAPTER X-Linked Recessive Disorders 183

Meiosis and the Genetic Basis of


Sexual Reproduction 149
9.1 An Overview of Meiosis 150
CHAPTER
11
Homologous Chromosomes 150
DNA Biology 187
The Human Life Cycle 151 11.1 DNA and RNA Structure
Overview of Meiosis 152 and Function 188
The Structure of DNA 188
9.2 The Phases of Meiosis 154
Replication of DNA 192
The First Division—Meiosis I 155
RNA Structure and Function 193
The Second Division—Meiosis II 155
11.2 Gene Expression 194
9.3 Meiosis Compared with Mitosis 156
From DNA to RNA to Protein 195
Meiosis I Compared with Mitosis 157
Review of Gene Expression 199
Meiosis II Compared with Mitosis 158
Mitosis and Meiosis Occur at 11.3 Gene Regulation 201
Different Times 158 Levels of Gene Expression Control 201
9.4 Changes in Chromosome Number 158

12
Down Syndrome 160
Abnormal Sex Chromosome Number 160 CHAPTER

Biotechnology and Genomics 210


CHAPTER
10 12.1 Biotechnology 211
Recombinant DNA Technology 211
Patterns of Inheritance 164 DNA Sequencing 212
Polymerase Chain Reaction 212
10.1 Mendel’s Laws 165
DNA Analysis 213
Mendel’s Experimental Procedure 165
Genome Editing 214
One-Trait Inheritance 166
Two-Trait Inheritance 169 12.2 Biotechnology Products 215
Mendel’s Laws and Probability 170 Genetically Modified Bacteria 215
Mendel’s Laws and Meiosis 171 Genetically Modified Plants 215
Genetically Modified Animals 216
10.2 Mendel’s Laws Apply to Humans 172
Family Pedigrees 172 12.3 Stem Cells and Cloning 217
Genetic Disorders of Interest 174 Reproductive and Therapeutic Cloning 218

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xii CONTENTS

12.4 Genomics and Proteomics 219


Sequencing the Bases of the Human Genome
Proteomics and Bioinformatics 221
220
CHAPTER
15
Evolution on a Small Scale 255
CHAPTER
13 15.1 Natural Selection 256
Types of Selection 257
Sexual Selection 258
Mutations and Genetic Testing 224 Adaptations Are Not Perfect 259
13.1 Gene Mutations 225 Maintenance of Variations 259
Causes of Gene Mutations 225
15.2 Microevolution 261
Types and Effects of Mutations 226
Evolution in a Genetic Context 261
13.2 Chromosomal Mutations 227 Causes of Microevolution 264
Deletions and Duplications 227
Translocation 228
Inversion 229
CHAPTER
16
13.3 Genetic Testing 229 Evolution on a Large Scale 269
Obtaining Chromosomes for Analysis 230
16.1 Speciation and Macroevolution 270
Karyotype Analysis 230
Defining Species 270
Testing for a Protein 231
Models of Speciation 273
Testing the DNA 231
Testing the Fetus 233 16.2 The Fossil Record 276
Testing the Embryo and Egg 234 The Geologic Time Scale 276
The Pace of Speciation 278
13.4 Gene Therapy 235
Causes of Mass Extinctions 279
Ex Vivo Gene Therapy 235
In Vivo Gene Therapy 236 16.3 Systematics 281
Linnaean Classification 281
Phylogenetic Trees 282
Cladistics and Cladograms 284
The Three-Domain System 285
UNIT III Evolution

CHAPTER
14 UNIT IV Diversity of Life
Darwin and Evolution 239
14.1 Darwin’s Theory of Evolution 240
Before Darwin 241
Darwin’s Conclusions 242
CHAPTER
17
Natural Selection and Adaptation 244
Viruses, Bacteria, and Protists 290
Darwin and Wallace 247 17.1 Viruses 291
Structure of a Virus 291
14.2 Evidence of Evolutionary Change 248
Viral Reproduction 292
Fossil Evidence 248
Plant Viruses 293
Biogeographical Evidence 250
Animal Viruses 293
Anatomical Evidence 250
Molecular Evidence 252 17.2 Viroids and Prions 296

mad87328_fm_i-xvi.indd 12 13/09/19 10:33 PM


CONTENTS xiii

17.3 Prokaryotes 298 Molluscs 349


The Origin of the First Cells 298 Annelids: Segmented Worms 350
Bacteria 299 19.4 Roundworms and Arthropods:
The Archaea 304 The Ecdysozoans 352
17.4 Protists 306 Roundworms: Pseudocoelomates 352
Evolution of Protists 306 Arthropods: Jointed Appendages 353
Classification of Protists 306 19.5 Echinoderms and Chordates:
The Deuterostomes 358
CHAPTER
18 Echinoderms 358
Chordates 359
Fishes: First Jaws and Lungs 361
Plants and Fungi 316
Amphibians: Jointed Vertebrate Limbs 363
18.1 An Overview of Plants 317 Reptiles: Amniotic Egg 364
An Overview of Plant Evolution 317 Mammals: Hair and Mammary Glands 366
Alternation of Generations 319
19.6 Human Evolution 369
18.2 The Diversity of Plants 320 Evolution of Humanlike Hominins 370
Nonvascular Plants 320 Evolution of Modern Humans 373
Vascular Plants 321
Gymnosperms 324
Angiosperms 326
UNIT V Plant Structure and Function
Economic Benefits of Plants 329
Ecological Benefits of Plants 329

18.3 Fungi 330


CHAPTER
20
General Biology of a Fungus 331 Plant Anatomy and Growth 378
Fungal Diversity 331
Ecological Benefits of Fungi 334 20.1 Plant Cells and Tissues 379
Economic Benefits of Fungi 335 Epidermal Tissue 379
Fungi as Disease-Causing Organisms 336 Ground Tissue 380
Vascular Tissue 381

CHAPTER
19 20.2 Plant Organs 381
Monocots Versus Eudicots 382

Animals 341 20.3 Organization of Leaves, Stems,


and Roots 383
19.1 The Evolution of Animals 342
Leaves 384
Ancestry of Animals 343
Stems 386
The Evolutionary Tree of Animals 343
Roots 388
Evolutionary Trends 344
20.4 Plant Nutrition 391
19.2 Sponges and Cnidarians: The Early
Adaptations of Roots for Mineral Uptake 392
Animals 346
Sponges: Multicellularity 346 20.5 Transport of Nutrients 393
Cnidarians: True Tissues 347 Water Transport in Xylem 393
Sugar Transport in Phloem 394
19.3 Flatworms, Molluscs, and Annelids:
The Lophotrochozoans 348
Flatworms: Bilateral Symmetry 348

mad87328_fm_i-xvi.indd 13 13/09/19 10:33 PM


xiv CONTENTS

21
Sensory Input and Motor Output 431
CHAPTER Reproduction 431

22.3 Homeostasis 432


Plant Responses and Organ Systems and Homeostasis 432
Reproduction 398 Negative Feedback 433
21.1 Plant Hormones 399
Auxins 399
Gibberellins 400
Cytokinins 401
CHAPTER
23
Abscisic Acid 401
The Transport Systems 437
Ethylene 402 23.1 Open and Closed Circulatory Systems 438
Open Circulatory Systems 439
21.2 Plant Responses 403
Closed Circulatory Systems 440
Tropisms 403
Comparison of Vertebrate Circulatory Pathways 440
Photoperiodism 404
23.2 Circulation in Humans 441
21.3 Sexual Reproduction in Flowering Plants 405
The Human Heart 442
Overview of the Plant Life Cycle 405
Blood Vessels 443
Flowers 406
Lymphatic System 446
From Spores to Fertilization 407
Capillary Exchange in the Tissues 447
Development of the Seed in a Eudicot 409
Monocots Versus Eudicots 409 23.3 Blood: A Transport Medium 448
Fruit Types and Seed Dispersal 410 Plasma 448
Germination of Seeds 411 Formed Elements 449
Cardiovascular Disorders 451
21.4 Asexual Reproduction and Genetic
Engineering in Plants 413
Propagation of Plants in a Garden 413
Propagation of Plants in Tissue Culture 413
Genetic Engineering of Plants 414
CHAPTER
24
The Maintenance Systems 456
24.1 Respiratory System 457
The Human Respiratory Tract 458
UNIT VI Animal Structure and Function
Breathing 459

22
Lungs and External Exchange of Gases 460
CHAPTER Respiration in Other Animals 461
Transport and Internal Exchange of Gases 461
Being Organized and Steady 420
24.2 Urinary System 463
22.1 The Body’s Organization 421 Human Kidney 464
Epithelial Tissue Protects 423 Problems with Kidney Function 467
Connective Tissue Connects and Supports 425

25
Muscular Tissue Moves the Body 427
Nervous Tissue Communicates 428 CHAPTER

22.2 Organs and Organ Systems 429


Transport and Protection 430
Digestion and Human Nutrition 471
Maintenance of the Body 430 25.1 Digestive System 472
Control 430 Complete and Incomplete Digestive Systems 472

mad87328_fm_i-xvi.indd 14 13/09/19 10:33 PM


CONTENTS xv

The Digestive Tract 472


Accessory Organs 478
Digestive Enzymes 479
CHAPTER
27
The Control Systems 519
25.2 Nutrition 481
Introducing the Nutrients 481 27.1 Nervous System 520
Examples of Nervous Systems 521
25.3 The Classes of Nutrients 482 The Human Nervous System 521
Carbohydrates 483 Neurons 522
Lipids 484 The Nerve Impulse 522
Proteins 485 The Synapse 524
Minerals 486 Drug Abuse 524
Vitamins 488 The Central Nervous System 526
Water 489 The Peripheral Nervous System 529
25.4 Understanding Nutrition Guidelines 490 27.2 Endocrine System 532
Updating Dietary Guidelines 490 The Action of Hormones 532
Visualizing Dietary Guidelines 490 Hypothalamus and Pituitary Gland 533
The Bottom Line 492 Thyroid and Parathyroid Glands 536
25.5 Nutrition and Health 493 Adrenal Glands 537
Body Mass Index 493 Pancreas 538
Disorders Associated with Obesity 496
Eating Disorders 498
CHAPTER
28
CHAPTER
26 Sensory Input and Motor Output 542
28.1 The Senses 543
Defenses Against Disease 502 Chemical Senses 543
26.1 Overview of the Immune System 503 Hearing and Balance 544
Lymphatic Organs 503 Vision 548
Cells of the Immune System 505 Cutaneous Receptors and Proprioceptors 550

26.2 Nonspecific Defenses and 28.2 The Motor Systems 552


Innate Immunity 505 Types of Skeletons 552
Barriers to Entry 505 The Human Skeleton 553
The Inflammatory Response 506 Skeletal Muscle Structure and Physiology 554
The Complement System 507
Natural Killer Cells 507

26.3 Specific Defenses and Adaptive


CHAPTER
29
Immunity 508 Reproduction and Embryonic
B Cells and the Antibody Response 508
Development 562
T Cells and the Cellular Response 509
29.1 How Animals Reproduce 563
26.4 Immunizations 512 Asexual Versus Sexual Reproduction 563
26.5 Disorders of the Immune System 514 Sexual Reproduction 563
Allergies 514
29.2 Human Reproduction 565
Autoimmune Diseases 515
Male Reproductive System 565
AIDS 515

mad87328_fm_i-xvi.indd 15 13/09/19 10:33 PM


xvi CONTENTS

Female Reproductive System 568 31.2 Ecology of Ecosystems 615


Control of Reproduction 571 Autotrophs 615
Infertility 573 Heterotrophs 615
Sexually Transmitted Diseases 574 Energy Flow and Chemical Cycling 616
29.3 Human Embryonic Development 577 Chemical Cycling 619
Fertilization 578 31.3 Ecology of Major Ecosystems 625
Early Embryonic Development 578 Primary Productivity 628
Later Embryonic Development 579
Placenta 581
Fetal Development and Birth 582
CHAPTER
32
Human Impact on the Biosphere 632
32.1 Conservation Biology 633

UNIT VII Ecology 32.2 Biodiversity 634


Direct Values of Biodiversity 635

CHAPTER
30 Indirect Values of Biodiversity 637

32.3 Resources and Environmental Impact 639


Ecology and Populations 586 Land 639
Water 641
30.1 The Science of Ecology 587
Food 643
Ecology: A Biological Science 588
Energy 645
30.2 The Human Population 589 Minerals 647
Present Population Growth 589 Other Sources of Pollution 647
Future Population Growth 590
32.4 Sustainable Societies 649
More-Developed Versus Less-Developed Countries 591
Today’s Society 649
Comparing Age Structures 592
Characteristics of a Sustainable Society 650
Population Growth and Environmental Impact 593

30.3 Characteristics of Populations 594


Distribution and Density 594 Appendix A Periodic Table of Elements
Population Growth 594 & The Metric System A-1
Patterns of Population Growth 596 Appendix B Answer Key B-1
Factors that Regulate Population Growth 597
Glossary G-1
30.4 Life History Patterns and Extinction 601
Extinction 602 Index I-1

CHAPTER
31
Communities and Ecosystems 605
31.1 Ecology of Communities 606
Community Composition and Diversity 607
Ecological Succession 608
Interactions in Communities 609
Community Stability 613

mad87328_fm_i-xvi.indd 16 04/10/19 12:39 PM


C H A P T E R

1
Biology: The
Science of Life

©Thomas Deerinck/Science Source

Artificial Life OUTLINE


What are the minimal requirements for life? That question has occupied the 1. 1 The Characteristics of Life
minds of philosophers and scientists for thousands of years. However, in just
the past decade, answers to this question have begun to emerge from a devel-
1.2 Evolution: The Core Concept of
oping field of scientific study called artificial life. Biology
One of the first of these studies occurred in 2010, when a research team 1.3 Science: A Way of Knowing
led by Craig Venter (a pioneer in genetic research) was successful in removing
the genetic information contained within the DNA of a bacterium and replacing
1.4 Challenges Facing Science
it with a synthetic form of DNA.
In 2016, the same group of researchers took their research one step fur-
ther. This time, they asked what minimal instructions were needed by a cell for
it to be considered alive. They constructed a cell that functioned on just 473
genes (humans have around 19,000). In the process, they not only narrowed in
on what the minimal requirements for life are, but also created the first example
of an artificial species.
The development of artificial life opens up the opportunity for humans to
construct cells that perform specific tasks, such as producing insulin, cleaning
toxic waste, or producing fuel more efficiently. However, there are concerns
about these new endeavors, and some scientists are urging constraint until the
risks have been determined.
In this chapter, we will explore the concept of life by examining the general
characteristics shared by all living organisms on our planet.

As you read through this chapter, think about the following questions:
1. What characteristics do you share with the diversity of life on the
planet?
2. How does adaptation and the process of evolution relate to living
organisms?
3. What are some of the challenges facing science and society today?

mad87328_ch01_001-020.indd 1 19/08/19 1:50 PM


2 CHAPTER 1 Biology: The Science of Life

1.1 The Characteristics of Life


Learning Outcomes

Upon completion of this section, you should be able to


1. Explain the basic characteristics common to all living organisms.
2. Distinguish between the levels of biological organization.
3. Summarize how the terms homeostasis, metabolism, and adaptation
relate to all living organisms.
4. Contrast chemical cycling and energy flow within an ecosystem.

bacteria human
As we observed in the chapter opener, life is diverse (Fig. 1.1). Life may be
found everywhere on the planet, from thermal vents at the bottom of the ocean
to the coldest reaches of Antarctica. Biology is the scientific study of life.
Biologists study not only life’s diversity but also the characteristics shared by
all living organisms. These characteristics include levels of organization, the
ability to acquire materials and energy, the ability to maintain an internal envi-
ronment, the ability to respond to stimuli, the ability to reproduce and develop,
and the ability to adapt and evolve to changing conditions. By studying these
characteristics, we gain insight into the complex nature of life, which helps us
distinguish between living organisms from nonliving things. In the next sec-
tions, we will explore these characteristics in more detail.
plant fungi The complex organization of life begins with atoms, the basic units of
matter. Atoms combine to form small molecules, which then join to form
Figure 1.1 Diversity of life. larger molecules within a cell, the smallest, most basic unit of life. Although a
Biology is the study of life in all of its diverse forms. cell is alive, it is made from nonliving molecules (Fig. 1.2).
(bacteria): ©Science Photo Library/Getty Images; (human): ©Purestock/
Superstock; (plant): ©Zeljko Radojko/Shutterstock; (fungi): ©Jorgen Bausager/
The majority of life on the planet, such as bacteria, are single-celled.
Getty Images Plants, fungi, and animals are multicellular organisms and are therefore com-
posed of many types of cells, which often combine to form tissues. Tissues
make up organs, as when various tissues combine to form the heart of an ani-
mal or the leaf of a plant. Organs work together in organ systems; for example,
the heart and blood vessels form the cardiovascular system. Various organ
systems often work together within complex organisms.
The organization of life extends beyond the individual organism. A
­species is a group of similar organisms that are capable of interbreeding. All of
the members of a species within a particular area belong to a population.
When populations interact, such as the humans, zebras, and trees in Figure 1.2,
they form a community. At the ecosystem level, communities interact with the
physical environment (soil, atmosphere, etc.). Collectively, the ecosystems on
the planet are called the biosphere, the zone of air, land, and water at the sur-
face of the Earth where living organisms are found.

Life Requires Materials and Energy


All life, from single cells to complex organisms, is not capable of maintaining
organization or carrying on its necessary activities without an outside source of
materials and energy. Food provides nutrient molecules, which are used as
building blocks or energy sources. Energy is the capacity to do work, and it
takes work to maintain the organization of the cell and the organism. When
cells use nutrient molecules to make their parts and products, they carry out a
sequence of chemical reactions. The term metabolism encompasses all the
chemical reactions that occur in a cell.

mad87328_ch01_001-020.indd 2 19/08/19 1:54 PM


CHAPTER 1 Biology: The Science of Life 3

Figure 1.2 Levels of biological organization.


Biosphere All life is connected by levels of biological organization that extend from
Regions of the Earth’s crust,
atoms to the biosphere.
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

Species
A group of similar, interbreeding organisms

human tree
Organism
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

nerve cell plant cell


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

mad87328_ch01_001-020.indd 3 19/08/19 1:56 PM


4 CHAPTER 1 Biology: The Science of Life

The ultimate source of energy for nearly all life on Earth is the sun.
Plants and certain other organisms are able to capture solar energy and carry
on photosynthesis, a process that transforms solar energy into the chemical
energy of nutrient molecules (see Section 6.1). For this reason, these organ-
isms are commonly called producers. Animals and plants get energy by metab-
olizing (Fig. 1.3), or breaking down, the nutrient molecules made by the
producers (see Section 7.1).
The energy and chemical flow between organisms also defines how an
ecosystem functions (Fig. 1.4). Within an ecosystem, 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) occurs as chemicals move from one popula-
tion to another in a food chain, until death and decomposition allow inorganic
nutrients to be returned to the producers once again. Energy (red arrows), on
Figure 1.3 Acquiring nutrient materials and energy. the other hand, flows from the sun through plants and the other members of the
All organisms, including this otter eating shellfish, require nutrients food chain as they feed on one another. The energy gradually dissipates and
and energy. returns to the atmosphere as heat. Because energy does not cycle, ecosystems
©Kirsten Wahlquist/Shutterstock could not stay in existence without solar energy and the ability of photosyn-
thetic organisms to absorb it.
Energy flow and nutrient cycling in an ecosystem largely determine
where different ecosystems are found in the biosphere. The two most biologi-
Solar cally diverse ecosystems—tropical rain forests and coral reefs—occur where
energy
solar energy is very abundant and nutrient cycling is continuous.
The availability of energy and nutrients also determines the type of bio-
logical communities that occur within an ecosystem. One example of an eco-
Heat
system in North America is the grasslands, which are inhabited by populations
of rabbits, hawks, and various types of grasses, among many others. The energy
input and nutrient cycling of a grassland are less than those of a rain forest,
Producers
which means that the community structure and food chains of these ecosys-
tems also differ.
Heat

Living Organisms Maintain an


Consumers
Internal Environment
For metabolic processes to continue, living organisms need to keep themselves
Chemicals

stable with regard to temperature, moisture level, acidity, and other factors
critical to maintaining life. Many of the metabolic activities of an organism are
Chemicals

involved in maintaining homeostasis, or an internal environment that acts


within a set of physiological boundaries.
Animals often vary their activity to regulate their internal environment.
A chilly lizard may raise its internal temperature by basking in the sun on a hot
rock. When it starts to overheat, it scurries for cool shade. Other organisms
have control mechanisms that do not require any conscious activity. When you
Decomposers Heat are studying and forget to eat lunch, your liver releases stored sugar to keep
your blood sugar level within normal limits. Many of the organ systems of our
bodies are involved in maintaining homeostasis.
Figure 1.4 Chemical cycling and energy flow in an ecosystem.
In an ecosystem, chemical cycling (aqua arrows) and energy flow
(red arrows) begin when plants use solar energy and inorganic
nutrients to produce their own food. Chemicals and energy are
Living Organisms Respond
passed from one population to another in a food chain. Eventually, Living organisms find energy and/or nutrients by interacting with their
energy dissipates as heat. With the death and decomposition of ­surroundings. Even single-celled organisms, such as bacteria, can respond to
organisms, chemicals are returned to living plants once more. their environment. The beating of microscopic hairs or the snapping of ­whiplike

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CHAPTER 1 Biology: The Science of Life 5

tails moves them toward or away from light or chemicals. Multi-


cellular organisms can manage more complex responses. A mon-
arch butterfly can sense the approach of fall and begin its flight
south, where resources are still abundant. A vulture can smell
meat a mile away and soar toward dinner.
The ability to respond often results in movement: The
leaves of a plant turn toward the sun, and animals dart toward
safety. Appropriate responses help ensure survival of the organ-
ism and allow it to carry on its daily activities. Altogether, we
call these activities the behavior of the organism.

Living Organisms Reproduce and Develop


Life comes only from life. Every living organism has the ability
to reproduce, or make another organism like itself. Bacteria and
other types of single-celled organisms simply split in two. In
­multicellular organisms, the reproductive process usually 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
divisions, results in an immature individual, which grows and
develops through various stages to become an adult.
An embryo develops into a whale or a yellow daffodil or a
human because of the specific set of genes, or genetic instructions,
inherited from its parents (Fig. 1.5). In all organisms, the genes are
located on long molecules of DNA (deoxyribonucleic acid), the
genetic blueprint of life. Variations in genes account for the differ-
ences between species and individuals. These differences are the
result of mutations, or inheritable changes in the genetic informa-
tion. Mutation provides an important source of variation in the
genetic information. However, not all mutations are bad—the
observable differences in eye and hair color are examples of DNA
NA
mutations.
By studying DNA, scientists are able to understand not
only the basis for specific traits, like susceptibility for certain
types of cancer, but also the evolutionary history of the species.
Reproduction involves the passing of genetic information from a
parent to its offspring. Therefore, the information found within
Figure 1.5 Reproduction is a characteristic of life.
the DNA represents a record of our molecular heritage. This includes not
Whether they are single-celled or multicellular, all organisms
only a record of the individual’s lineage, but also how the species is related
reproduce. Offspring receive a copy of their parents’ DNA and
to other species. therefore a copy of the parents’ genes.
DNA provides the blueprint or instructions for the organization and (photo): ©Purestock/Superstock; (DNA): ©Molekuul/SPL/AGE Fotostock
metabolism of the particular organism. All cells in a multicellular organism
contain the same set of genes, but only certain ones are turned on in each type
of specialized cell. Through the process of development, cells express spe-
cific genes to distinguish themselves from other cells, thus forming tissues
and organs.

Living Organisms Have Adaptations


Adaptations are modifications that make a species suited to their way of life.
Some hawks have the ability to catch fish; others are best at catching rabbits.
Hawks can fly, in part, because they have hollow bones to reduce their weight

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6 CHAPTER 1 Biology: The Science of Life

and flight muscles to depress and elevate their wings. When a hawk dives, its
strong feet take the first shock of the landing, and its long, sharp claws reach
out and hold onto the prey. Hawks have exceptionally keen vision, which
enables them not only to spot prey from great heights but also to estimate
­distance and speed.
Humans also have adaptations that allow them to live in specific envi-
ronments. Humans who live at extreme elevations in the Himalayas (over
13,000 feet, or 4,000 meters) have an adaptation that reduces the amount of
hemoglobin produced in the blood (see Section 19.6). Hemoglobin is impor-
tant for the transport of oxygen. Normally, as elevation increases, the amount
CONNECTING THE CONCEPTS of hemoglobin increases, but too much hemoglobin makes the blood thick,
1.1 All living organisms, from bacteria which can cause health problems. In some high-elevation populations, a
to humans, share the same basic mutation in a single gene reduces this risk.
characteristics of life. Evolution, or the manner in which species become adapted to their envi-
ronment, is discussed in the next section of this chapter.

Check Your Progress 1.1


1. List the basic characteristics common to all life.
2. List in order, starting with the least organized, the levels of biological
organization.
3. Explain how chemical cycling and energy flow occur at both the
organism and the ecosystem levels of organization.

1.2 Evolution: The Core Concept


of Biology
Learning Outcomes

Upon completion of this section, you should be able to


1. Define the term evolution.
2. Explain the process of natural selection and its relationship to
evolutionary processes.
3. Summarize the general characteristics of the domains and major
kingdoms of life.

Despite diversity in form, function, and lifestyle, organisms share the same
basic characteristics. As mentioned, they are all composed of cells organized in
a similar manner. Their genes are composed of DNA, and they carry out the
same metabolic reactions to acquire energy and maintain their organization.
The unity of living organisms suggests that they are descended from a common
ancestor—the first cell or cells.
An evolutionary tree is like a family tree (Fig. 1.6). Just as a family
tree shows how a group of people have descended from one couple, an
evolutionary tree traces the ancestry of life on Earth to a common ancestor.
One couple can have diverse children, and likewise a population can be a
common ancestor to several other groups, each adapted to a particular set of

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CHAPTER 1 Biology: The Science of Life 7

Bacteria Archaea Protists Plants Fungi Animals


0

0.5
Billions of Years Ago (BYA)

1.0

1.5

2.0

2.5
EUKARYA Figure 1.6 An evolutionary tree.
ARCHAEA Organisms grouped on the same branch of the tree have a common
3.0 BACTERIA ancestor located at the base of the branch. Organisms grouped on the
3.5 same branch (such as fungi and animals) are more closely related to one
another, meaning they have a more recent common ancestor than
4.0
organisms on different branches (such as animals and plants). The base of
First ancestral cell the tree itself represents the common ancestor of all living organisms.

environmental conditions. Evolution is the process in which populations


change over time to adapt to their environment, and pass on these changes
to the next generation. Evolution is considered the unifying concept of biol-
ogy because it explains so many aspects of biology, including the tremen-
dous diversity of life on the planet and how living organisms arose from a
single ancestor.

Natural Selection and Evolutionary


Processes
In the nineteenth century, two naturalists—Charles Darwin and Alfred ­Russel
Wallace—came independently to the conclusion that evolution occurs by
means of a process called natural selection (see Section 14.1). Charles
­Darwin is the more famous of the two because he wrote a book called On the
Origin of Species, which presented his observations on how the process of
evolution worked using natural selection. Since that time, evolution has
become the core concept of biology, not only because the theory explains so
many different scientific observations, but because the wealth of data col-
lected since Darwin’s era supports the theory’s importance in every aspect of
the biological sciences.
The process of natural selection is the mechanism by which evolution-
ary change occurs. It is based on how a population changes in response to its
environment. Environments may change due to the influence of living factors
(such as a new predator) or nonliving factors (such as temperature). As the
environment changes over time, some individuals of a species may possess
certain adaptations that make them better suited to the new environment.
Individuals of a species that are better adapted to their environment tend to
live longer and produce more offspring than other individuals. This differen-
tial reproductive success, called natural selection, results in changes in the

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8 CHAPTER 1 Biology: The Science of Life

characteristics of a population over time. That is, adaptations that result in


higher reproductive success tend to increase in frequency in a population
from one generation to the next. This change in the frequency of traits in
populations is called evolution.
The phrase “common descent with modification” sums up the pro-
cess of evolution because it means that, as descent occurs from common
ancestors, modifications occur that cause the organisms to be adapted
(suited) to the environment. As a result, one spe-
cies can be a common ancestor to several
species, each adapted to a particular set of
environmental conditions. Specific adapta-
Kauai tions allow species to play particular roles in
‘Akialoa their environment.
The Hawaiian honeycreepers are
Laysan finch a remarkable example of this process
Kona finch
(Fig. 1.7). The more than 50 known
‘Akepa Maui parrotbill
species of honeycreepers (of which
Nukupu’u
only 17 species remain today) all
evolved from one species of finch, which
likely originated in North America and
Alauwahio ‘Anianiau arrived in the Hawaiian islands between 3
and 5 million years ago. Modern hon-
Palila
eycreepers have an assortment of bill
Amakihi shapes adapted to different types of
`Ō'ō food. Some honeycreeper species
have curved, elongated bills used for
‘Akiapola’au
drinking flower nectar. Others have
Crested honeycreeper strong, hooked bills suited to digging in
tree bark and seizing wood-boring insects or short, straight, finchlike bills
for feeding on small seeds and fruits. Even with such dramatic differences
‘Ula-‘ai-Hawane in feeding habits and bill shapes, honeycreepers still share certain charac-
‘Apapane
teristics, which stem from their common finch ancestor. The various hon-
eycreeper species are similar in body shape and size, as well as mating and
nesting behavior.
The study of evolution encompasses all levels of biological organiza-
tion. Indeed, much of today’s evolution research is carried out at the molecu-
lar level, comparing the DNA of different groups of organisms to determine
how they are related. Looking at how life has changed over time, from its
origin to the current day, helps us understand why there are so many different
Mamos
kinds of organisms and why they have the characteristics they do. An under-
standing of evolution by natural selection also has practical applications,
including the prevention and treatment of disease.
Today, we know that, because of natural selection, resistance to anti-
liwi
biotic drugs has become increasingly common in a number of bacterial
species, including those that cause tuberculosis, gonorrhea, and staph
infections. Antibiotic drugs, such as penicillin, kill susceptible bacteria.
However, some bacteria in the body of a patient undergoing antibiotic
treatment may be unharmed by the drug. Bacteria can survive antibiotic
drugs in many different ways. For example, certain bacteria can endure
treatment with penicillin because they break down the drug, rendering it
Figure 1.7 Evolution of Hawaiian honeycreepers. harmless. If even one bacterial cell lives because it is antibiotic-resistant,
Hawaiian honeycreepers, descendants of a single ancestral species, then its descendants will inherit this drug-defeating ability. The wide-
display an amazing diversity of bill shapes and sizes. spread use of antibiotics has favored the evolution of resistant bacterial

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CHAPTER 1 Biology: The Science of Life 9

strains, which in turn is limiting the effectiveness of many classes of


antibiotics.

Connections
How does evolution affect me personally?
Each year, starting around September, health agencies and pharmacies
begin alerting us to get our annual flu vaccine. Often, people question
the need to get an annual vaccine, and in the process, place themselves
at risk of infection.

What we call the flu is actually a response by our bodies to an infection


by influenza virus. Influenza viruses are relatively simple (see Section
17.1), but they have the ability to mutate each year, creating new strains
that have not been seen before by our immune systems. Some of these
changes have been responsible for strains that have created significant
outbreaks in the human population.

The Centers for Disease Control and Prevention (CDC) and other agen-
cies are responsible for monitoring the evolution of the influenza virus,
and for developing annual vaccines. However, at times, the virus may
evolve during the influenza season (as happened in 2018), thus reducing
the effectiveness of that year’s vaccine.

Organizing the Diversity of Life


Think of an enormous department store, offering thousands of different items
for sale. The various items are grouped in departments—electronics, apparel,
furniture, and so on—to make them easy for customers to find. Because life is
so diverse, it is helpful to have a system that groups organisms into categories.
Two areas of biology help us group organisms into categories: T ­ axonomy is
the discipline of identifying and naming organisms according to certain rules,
and systematics makes sense out of the bewildering variety of life on Earth by
classifying organisms according to their presumed evolutionary relationships.
As systematists learn more about evolutionary relationships between species,
the taxonomy of a given organism may change. Systematists are even now
making observations and performing experiments that will one day bring about
changes in the classification system adopted by this text.
Table 1.1 Levels of Biological Organization
Categories of Classification Category Human Corn
The classification categories, from least inclusive to most inclusive, are spe- Domain Eukarya Eukarya
cies, genus, family, order, class, phylum, kingdom, and domain (Table 1.1).
Kingdom Animalia Plantae
Each successive category above species contains more types of organisms than
the preceding one. Species placed within one genus share many specific char- Phylum Chordata Anthophyta
acteristics and are the most closely related, while species placed in the same Class Mammalia Liliopsida
domain share only general characteristics. For example, all species in the genus
Order Primates Commelinales
Pisum look pretty much the same—that is, like pea plants—but species in the
plant kingdom can be quite varied, as is evident when we compare grasses with Family Hominidae Poaceae
trees. By the same token, only modern humans are in the genus Homo, but Genus Homo Zea
many types of species, from tiny hydras to huge whales, are members of the
Species* H. sapiens Z. mays
animal kingdom. Species placed in different domains are the most distantly
related. For now, we will focus on the general characteristics of the domains *To specify an organism, you must use the full binomial name, such as
Homo sapiens.
and kingdoms of life.

mad87328_ch01_001-020.indd 9 04/10/19 12:55 PM


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SYMPTOMS.—An attack of chorea is usually preceded by more or less
failure of the general health and evidences of some mental
disturbance. It is quite common to be told by the parents of a child
suffering from chorea that the little patient had seemed unwell for
some time previous to the attack; that the appetite had failed, and
that the child had looked pale; that he had been irritable or excitable,
and at school the teacher had complained of restlessness or
inattention in the pupil. In a little girl who was brought to me recently
with her second attack of chorea her mother stated that for several
days before the outbreak the child had been in excessive spirits, and
that she had been singing loudly and in a peculiar manner. The
same symptoms had preceded the first attack. Sometimes nothing is
observed until it is found by the parents or teacher that there are
abnormal twitchings and movements of the limbs.

At first there is a general restlessness and fidgetiness. The child may


be punished at school for not keeping still or for dropping things.
Soon irregular movements of groups of muscles are seen. The
shoulder is shrugged or the fingers move spasmodically. At first the
patient is aware of the movements and tries to control them, but
before long the twitching and jerking are constant, and extend to
most of the voluntary muscles of the body. He is then unable to
control them for any length of time.

The sudden jerk of a limb followed by an odd grimace, the quick


protrusion of the tongue, and the rolling of the eyes or snapping of
the lids give a characteristic picture which can hardly be mistaken.

The extent of the movements varies in different cases. In some they


are slight and affect only certain muscles. Often the disorder is
confined to one lateral half of the body. In other cases the
movements involve all the limbs and the trunk, and are so violent
and constant that the patient does not seem to have a moment's
rest. The trunk may be suddenly drawn backward, then the arms are
extended or thrown up, and the legs flexed and tossed about with
great quickness. Sometimes the patient is thrown off the bed or from
the chair on which he may be to the floor.
The speech is often affected. The patient speaks in a thick or jerky
manner, as if the tongue were too large for the mouth, and saliva
usually flows in great quantities. Sometimes in bad cases there are
involuntary utterances made at frequent intervals.

The features undergo contortions continually, and when at rest


relapse into a condition of vacancy which makes the patient look
almost idiotic. The expression of a child with chorea is so peculiar
that the disease may almost be diagnosticated by this.

During sleep the movements usually cease, but generally the patient
is restless while asleep, and in some instances the irregular
movements continue even at this time.

The mental condition commonly shows some change. The child is


irritable and peevish, cries and laughs readily, or is sullen and
morose. Sometimes he is violent to those about him, but this is rare.
Intellectually the patient suffers somewhat. He is not able to study as
before, and the memory may be impaired. Sometimes there is a mild
form of dementia.

During the course of the disease there may be exacerbations, and


sometimes after convalescence has seemed established there are
relapses.

Recovery is gradual, and as the abnormal movements cease the


mental condition improves, and the patient regains his health without
any traces of the disease remaining.

We will now consider some of the symptoms separately. First, as to


disturbances of motion. As before remarked, the disorderly
movements occur soon after the general restlessness is seen. They
most commonly begin in one upper extremity. The hand is thrown
into various positions, the fingers are flexed and extended or
separated, and all of the movements occur with great rapidity. In a
day or two the whole arm is affected, and then the leg of the same
side is involved in the jerkings and twitchings. In many cases the
facial muscles are contorted, the mouth is pursed up or opened
wide, and then quickly twisted into some other shape. If the patient is
told to put out his tongue, it is protruded after a moment's hesitation,
and then suddenly retracted, the jaws coming together with a snap.
A smacking sound is made with the lips quite often, and words are
uttered involuntarily. The movements may remain confined to one
side of the body, constituting what is called hemichorea. This is quite
common, and the right side is rather more frequently involved than
the left. There is so great difference of opinion among authors on this
point that it is probable that one side is affected about as often as the
other. Of 252 cases which I have examined, 69 were right and 43 left
hemichoreas. Gerhard20 found in 80 cases of chorea that 32 were
unilateral; of these 20 were right and 12 were left. Sée, however,
found that in 97 of 154 cases the movements were either confined to
the left side or were more marked on that side. He states that in his
experience the proportion between left and right hemichorea is as 37
to 27. Pye-Smith in 33 cases of unilateral chorea found 15 on the
right and 18 on the left side. Many cases which begin as hemichorea
soon become general.
20 American Journal of Med. Sci.

The disease reaches its greatest severity in about two weeks, and if
the case is a bad one we find by this time all of the voluntary
muscles are in constant movement. At this time the French name for
chorea, folie musculaire, is most appropriate. Patients are often
unable to walk or to sit up, and sometimes they may be thrown from
the bed by violent spasmodic movements of the trunk. Strange as it
seems, patients rarely complain of fatigue, notwithstanding the
violent muscular exercise. This is probably because each set of
movements is of short duration and is constantly changing its seat.

As a rule, the movements cease completely during sleep or under an


anæsthetic. Sometimes occasional twitchings of muscles are seen in
sleep, and in rare instances we are told by the parents of a child with
chorea that the movements are as active in sleep as in the waking
hours.
The movements of chorea occur either while the limbs are at rest or
under the influence of voluntary effort. This fact has been pointed out
by Mitchell and by Gowers. In some cases the movements are most
marked when the patient is at rest. If a directed effort is made to use
the member for a time, the choreic movements are suspended. For
example, a patient may be able to carry a glass of water to the
mouth without spilling a drop, while a moment before the hand may
have been performing a continual dance. I have often observed that
while the limb to which the whole attention has been directed in
performing some movement has been steadied, the other limbs
become violently agitated.

In another class of cases the movements are comparatively slight


when the part is at rest, but when a motion is attempted the disorder
of the muscles is so much increased that it is almost impossible for
the act to be completed. The patient is told to pick up some small
object: he throws the hand out toward it, and it is jerked away before
he can grasp it. He again puts the hand forward, reaches the object,
and the fingers open and shut and sprawl over the article before it is
taken up. Sometimes it cannot be grasped at all. This has been
called choreic ataxia, but it is only one type of the cases commonly
seen.

This brings us to the influence of the will on the movements in


chorea. There are some cases, as mentioned above, in which the
movements may be controlled by the will for a brief period, but they
will sooner or later return. In other cases it is quite impossible for the
patient to check the movements at all, and one frequently sees in a
case of hemichorea the sound hand used to grasp the other, so as to
control the movements. We have referred to this because of
Niemeyer's opinion that corporeal punishment would shorten an
attack of chorea.

Chorea is sometimes confined to a single muscle or group of


muscles. When limited in this way it is generally in the head, face, or
perhaps in the shoulder. These cases of localized chorea have been
spoken of by Mitchell as habit chorea.21 They are often very
obstinate in resisting treatment, and sometimes last during life.
21 Lectures on Nervous Diseases, p. 146.

PARALYSIS.—Not infrequently in chorea there is paralysis to a greater


or less extent. It is generally one-sided, and most often involves the
upper extremity. The limb affected is the one in which the
movements were most violent. The arm may hang entirely powerless
or it may be only enfeebled, and feeling to the patient like a dead
weight. The paralysis always recovers with the chorea or soon after.

POST-PARALYTIC CHOREA.—Under this term Mitchell and Charcot have


described a variety of chorea which is seen in patients after an
attack of hemiplegia. The movements are chiefly on voluntary effort,
and are those of inco-ordination. They come on from one to several
months after an attack of unilateral paralysis, and are sometimes
seen in cases in which almost complete recovery has taken place.
Mitchell has reported22 a case which was under my care for several
years, and which he saw in consultation with me. This patient had
two attacks of left hemiplegia, the last being fatal. After the first
attack there was great gain of power to use the arm and leg, but the
movements were performed awkwardly and with an irregular jerking
movement. A post-mortem examination revealed a spot of softening
the size of a filbert in the left corpus striatum, which was apparently
recent, and a point of red degeneration in the right crus cerebri. The
vessels at the base of the brain were extensively atheromatous.
22 American Journal of the Med. Sci.

Of the electrical condition of the muscles in chorea but little is known.


Rosenthal23 found increase of faradic contractility in three cases of
hemichorea, and the galvanic test showed a high degree of
excitability, demonstrated by the fact that weak currents gave
contractions at cathodic closure, or even tetanic contractions, and
also contractions were produced at cathodic opening.
23 Ziemssen's Cyclopædia, loc. cit., p. 434.
The affection of speech which is so common in chorea is due to
disordered action of the laryngeal muscles, or it may be from choreic
action of the abdominal muscles. Sometimes it is chiefly from the
awkwardness of the tongue. The usual form of trouble is that the
patient speaks in a staccato manner and the syllables seem as if
they were driven out. When the chorea is in the laryngeal muscles,
the tone and pitch of the voice are altered.

Chorea of the heart is sometimes spoken of, but it has never been
satisfactorily demonstrated that there is any real disorder of cardiac
rhythm in chorea. It is not unusual in chorea to meet with over-action
or palpitation of the heart, but these conditions do not necessarily
depend on the disease.

Valvular murmurs are often met with from the beginning of an attack.
In some instances they are the result of an endocarditis, but
frequently they are functional or anæmic. They are usually heard at
the apex. Sometimes there is a reduplication of the first sound,
giving the idea of a want of synchronism in action of the two sides of
the heart; but this is probably not the result of chorea of the heart. I
recall one patient, a child of seven or eight years, in whom the
reduplication of the first sound was very distinct during an attack of
St. Vitus's dance. She was brought to me at the beginning of a
second attack a year later, and the reduplication of the cardiac
sounds was heard again, so it is likely that it had continued during
the interval, and was probably a congenital condition.

The pupils are commonly dilated in chorea and respond sluggishly to


light.

REFLEXES.—I have examined the condition of the patellar reflex in 50


cases. In 26 of these it was present in normal degree, in 15 it was
diminished, and in 9 it could not be excited. In one patient it was
absent during the height of the choreic movements, but could be
readily produced after the patient had recovered. The condition of
the reflexes has also been examined by Joffroy and Saric,24 and they
found that of 16 cases of chorea the reflexes were abolished or
diminished in 12.
24 L'Union médicale, Sept. 22, 1885.

SENSIBILITY.—Authors state that disorders of sensation are met with


in chorea, such as localized anæsthesia or a general hyperæsthesia:
I have never met with any such instances. Patients often complain of
pain in the joints or in the limbs, and this may be unaccompanied
with swelling or tenderness on pressure. Tenderness on pressure
over the vertebræ is rare in my experience, although others speak of
its being of frequent occurrence. Mental disorders are generally
present, but only to a slight extent. There is almost always irritability
of temper and peevishness. The most sweet-tempered children
become cross and perverse, laugh immoderately at trifling things, or
cry as readily if they are annoyed. There is generally failure of
memory and incapacity for study or thought. In most cases, however,
this exists to so slight an extent as not to be noticed except on very
close observation. Sometimes there is marked mental disorder
amounting almost to imbecility, and occasionally the mental
weakness remains for some time after the motor disorders have
recovered.

The condition of the pulse is generally unchanged, but sometimes it


is abnormally frequent. The temperature, according to Von
Ziemssen, is unchanged.

The nutrition generally suffers. The patient rapidly loses flesh, and
becomes anæmic; the skin grows dry, and the hair gets harsh. The
digestion is apt to be disordered. The tongue is large, pallid, and
coated thickly, and there is sometimes nausea or vomiting. The
appetite is not good. The bowels are often constipated. The urine
has been examined by several observers. Bence Jones found an
excess of urea at the height of the disease. Albumen is not present
except accidentally, but there is usually an excess of phosphates. In
several cases in which we have examined the urine at the Infirmary
for Nervous Diseases we found that the specific gravity was high
while the chorea was at its height, but fell to normal as the patient
recovered.
Chorea is spoken of as acute and chronic, but all cases are more or
less chronic. Those cases which last eight or ten weeks may be
considered acute, while those running on for months or years are
properly called chronic.

DURATION.—Considerable difference of opinion exists as to the


duration of chorea. Some writers speak of three or four weeks as an
average attack. Gray and Tuckwell, in a series of cases treated by
the expectant plan,25 found an average duration of ten weeks.
Occasionally a patient is seen with an attack of chorea which lasts
only a few days. The parents of a little patient whom I saw a few
days ago assured me that her second attack lasted only a week.
They are educated and intelligent persons whose statement can be
relied upon.
25 Lancet, Nov. 28, 1876.

The course is not always regular. In some cases the disease


gradually reaches a crisis, remains stationary for a few days, and
then by degrees declines; in others there are exacerbations. The
patient will seem to be almost well, and then become very much
worse for a time. Relapses are not infrequent, and are generally
caused by fright or excitement.

The recurrence of attacks of chorea is well known. A child who has


had the disease one year may have it a second or third year. It is
most likely to recur in the spring. Some cases have as many as five
attacks, but as puberty approaches the attacks are lighter, and finally
cease. Of 282 cases to which I have referred, 198 were first attacks,
47 had had chorea twice; 23 were in their third attack, 8 in the fourth,
and 3 in the fifth attack.

TERMINATION.—The disease in most instances terminates in complete


cure, but sometimes there is nervousness or want of co-ordination
remaining for a time. Rarely the inco-ordination or a certain
quickness in movement becomes permanent.
Death is a rare termination of chorea except in pregnancy. If it does
occur, it is usually from some complication. In pregnancy the
mortality is great. Of 64 cases collected by Wenzel, 18 died. In
Philadelphia, in seventy-four years from 1807 to 1881, there have
been but 64 deaths from chorea; of these, 38 were under twenty
years, and 26 over that age.

Hutchinson reports a fatal case in a boy of twelve years.26 After


complaining of headache and rheumatic pains for several days,
choreic movements began. They soon became general and very
violent. At the end of two weeks he was admitted to the
Pennsylvania Hospital. At this time the patient was so extremely
convulsed that it was impossible to keep him in bed without tying
him. The movements continued but little abated, and the child died in
two days.
26 Philadelphia Med. Times, vol. vi. p. 535.

Another case of unusual interest is reported by Hunt.27 The patient, a


man of twenty-nine years, had suffered from chorea of the face and
arms for years. In consequence of a fall on the pavement he
fractured the left humerus. The movements were immediately
exaggerated, and in spite of a carefully adjusted splint it was
impossible to keep the arm at rest. The fragments were in a state of
constant movement, and the points of bone threatened to penetrate
the skin. The skin was so much excoriated that it was determined to
dispense with the splint and attempt to keep the limb at rest by the
administration of morphia hypodermically in half-grain doses three
times daily. This failed to keep the arm quiet, and the seat of fracture
became greatly inflamed. No form of appliance or medication
succeeded in keeping the arm at rest, and the patient finally sank
and died from exhaustion on the tenth day after admission to the
hospital. The post-mortem examination revealed no gross lesion of
the brain or cord. No microscopic examination was made of the
brain.
27 Pennsylvania Hospital Reports, vol. ii.
MORBID ANATOMY AND PATHOLOGY.—In a disease so seldom fatal as
chorea it is not surprising that there have been but few post-mortem
examinations made. In the earlier autopsies, before the microscope
was extensively used, but little of value was recorded. Sée, who
collected 84 cases in which post-mortem examinations were made,
reported that in 16 no changes were found in the nervous system. In
32 there were lesions in the brain and nervous centres, usually
softening and tuberculosis, and in the remainder inflammatory
changes in the serous membranes. In 29 there were evidences of
heart disease. Sée considered that but few cases of death in chorea
were caused by inflammatory diseases of the heart, but that the
majority should be referred to nervous excitement and anæmia.

Ogle28 in a report of 96 cases of chorea mentions 16 which were


fatal. Post-mortem examinations were made in all of these. Cardiac
lesions were found in 13. In 10 of these deposits were found upon
the valves, and in 3 there was some change in the pericardium. He
speaks of having noted congestion of the nervous centres six times,
and softening of the cord once.
28 Brit. and For. Med.-Chir. Review, Jan., 1868.

In all of 11 autopsies reported by Pye-Smith29 there were cardiac


lesions found. In every case old or recent deposits were observed
upon the valves. In two instances the heart was hypertrophied, and
in one there was pericarditis. Changes in the nervous system were
less often found by this writer. In 1 case there was hyperæmia of the
cord, and in 3 cerebral hyperæmia.
29 Guy's Hospital Reports, 1874.

Dickinson found in 22 fatal cases of chorea 17 in which the heart


was diseased. “In every instance making up the large tale of cardiac
disease there were recent vegetations on the mitral valve, and often
also elsewhere.”

In the fatal case of Hutchinson referred to above the heart was found
diseased, the aortic valves were incompetent, the leaflets being
swollen and softened, and the aorta was atheromatous above the
sinus of Valsalva.

Of late years a number of careful autopsies have been made in


cases of chorea. The brain and spinal cord have been closely
examined, and in almost every instance some lesion has been found
in both of these organs.

Steiner reported in 1868 the results of post-mortem examinations in


3 fatal cases of chorea. In 1 case he found cerebro-spinal anæmia,
serous effusion into the spinal canal, and proliferation of the
connective tissue in the upper part of the cord; and in another
hyperæmia of the brain and cord.

Elischer,30 who reports a fatal case in a parturient woman who had


an attack of chorea in her eighth year, two in her sixteenth year, and
another in a previous pregnancy, found at the autopsy hyperæmia
and œdema of the brain and gray substance of the cord.
Microscopically, the brain showed fatty, amyloid, and pigmentary
changes in the nerve-elements and vessels of the large central
ganglia, small secondary extravasations of blood in the connective
tissue, and numerous emboli in the smallest vessels, especially in
the cortex. In the spinal cord there was seen abundant proliferation
of nuclei in the adventitia of the vessels. In the central canal serum
was found, and the surrounding connective tissue was harder than
usual.
30 Cyclopædia of the Practice of Medicine, Von Ziemssen, vol. xiv. p. 450.

Dickinson has contributed an excellent paper on the pathology of


chorea.31 He relates the particulars of the autopsies in 7 fatal cases
in which he personally made microscopical examination of the brain
and spinal cord. He also adds the results of post-mortem
examinations in 17 other cases at St. George's Hospital and at the
Hospital for Sick Children. In all of the 7 cases in which microscopic
examinations of the brain and cord were made there were found
hyperæmia of both of these structures, in many instances
hemorrhages into the substance of the nervous tissues, dilatation of
the smaller vessels, and in chronic cases sclerotic changes in the
course of the vessels. “The first visible change,” he remarks, “would
seem to be the injection or distension of the arteries, succeeded by
extrusion of their contents, to the irritation and injury of the
surrounding tissue.” The changes seemed to affect both brain and
cord in all cases. The parts of the brain most constantly affected lay
between the base and the floor of the lateral ventricles in the track of
the middle cerebral arteries, the substantia perforata, the corpora
striata, and the beginning of the Sylvian fissures. “Of the cord no
region was exempt, but perhaps the cervical and dorsal regions were
usually more affected than the lumbar. With regard to the vertical or
physiological divisions of the cord, these all, whether white or gray,
shared in the vascular destruction; this condition, however, was
usually most marked in the vessels belonging to or in connection
with the lateral part of the gray matter about the root of each
posterior horn. And it is to be observed that this was also the chosen
situation of the more definite and special changes, whether
hemorrhagic (as in two instances), sclerose, or exudatory. Speaking
generally, the chosen seats of the choreic changes are the parts of
the brain which lie between the beginning of the middle cerebral
arteries and the corpora striata—the parta perforata; and in the cord
the central portion of each lateral mass of gray matter comprising the
root of each posterior horn.”
31 Medico-Chirurgical Transactions, vol. xli. p. 1, 1876.

The embolic theory of chorea has been held by several investigators,


among them Hughlings-Jackson. It is undoubtedly an attractive and
reasonable view, especially when we consider the large proportion of
cases in which there is valvular disease of the heart. Dickinson,
however, does not consider this hypothesis tenable. In none of the
cases in which he made post-mortem examinations did he find
evidences of embolism. “In none of the instances described were
decolorized fibrin, detached clots, or signs of impaction detected,
and the erraticism of embolic accident was wanting: the constancy
indeed with which the changes repeated themselves in certain
positions, and the equality with which they affected both sides of the
body, are conclusive objections to this hypothesis. The corpora
striata, for example, were affected with almost absolute symmetry,
notwithstanding that these bodies receive their blood respectively
from the right and left carotids and different parts of the aortic arch.”

Rheumatism is associated with or precedes chorea in a large


proportion of cases, and this was pointed out by Kirkes in 1850 and
again in 1863. This connection between rheumatism and chorea,
and the frequent occurrence of endocarditis in chorea, has led some
authors to believe that the endocarditis is always rheumatic, and that
the chorea is the result of the endocarditis. Dickinson, however,
points out that in cases in which there is a distinct history of the
chorea beginning suddenly from fright there are often well-marked
cardiac murmurs heard. He believes that in all cases of chorea in
which there are cardiac murmurs they are due to endocarditis, and
suggests that in these cases from fright the endocarditis is due to
irregularity of cardiac action. This, of course, is mere hypothesis, and
we must bear in mind that in all cases of chorea there is anæmia,
and that the murmur may be purely functional.

H. C. Wood, in a communication read before the College of


Physicians of Philadelphia,32 gives his views of the pathology of
chorea, based upon the results of post-mortem examinations made
in a number of dogs who had the disease. He believes the history of
chorea to be this: “Owing to emotional disturbance, sometimes
stopping of various vessels of the brain, or sometimes the presence
of organic disease, there is an altered condition of the ganglionic
cells throughout the nerve-centres. If the cause is removed and the
altered condition of the nerve-cells goes only so far, it remains what
we call a functional disease. If it goes so far that the cells show
alteration, we have an organic disease of the nervous system.”
32 Philada. Med. News, May 30, 1885.

In two dogs which were choreic the movements continued after


section of the cord. This shows that in dogs, at any rate, the
movements originate in the cord. In four instances of canine chorea
in which Wool made autopsies there were found in the cords of three
mild grades of infiltration of leucocytes in the gray matter. In the
fourth, in which the dog had died of the disease, the ganglion-cells
were degenerated, and in some places had disappeared. He
concludes, therefore, that choreic movements may depend upon a
diseased condition of the motor cells of the cord.

Although there are several recorded cases of human chorea in which


lesions of the spinal ganglionic cells have been found, we cannot
believe that this can be a constant lesion in chorea. The disease is
too transient in many cases, and presents too many variations and
anomalies, for the cord to be always the seat of the diseased
condition.

In an interesting paper read by Angel Money before the London


Medical and Chirurgical Society in 1885 he detailed some
experiments in which, by injecting a fluid containing arrowroot,
starch-granules, or carmine into the carotids of animals, he produced
movements closely resembling chorea; and this was found to be
associated always with embolism of the capillaries of the cord. In the
discussion which followed Broadbent and Sturges expressed their
disbelief in the embolic origin of chorea in man. Hughlings-Jackson
said that he held the view of the cerebral origin of chorea, one of his
reasons being the frequency with which the face-muscles are
affected in this disease.

The probabilities are that in chorea there is a disordered condition of


the brain and cord more or less general. The lesions are no doubt
slight in mild cases of short duration, but in severe cases of long
standing there occur well-marked changes in portions of both brain
and cord. We cannot do better than to sum up the pathology of
chorea in the words of Dickinson: “A widely-distributed hyperæmia of
the nervous centres, not due to any mechanical mischance, but
produced by causes mainly of two kinds—one a morbid, probably a
humoral, influence which may affect the nervous centres as it affects
other organs and tissues; the other, irritation in some mode, usually
mental, but sometimes what is called reflex, which especially
belongs to and disturbs the nervous system, and affects persons
differently according to the inherent mobility of their nature.”

DIAGNOSIS.—The only diseases for which chorea may be mistaken


are paralysis agitans and disseminated sclerosis. The former occurs
only in adult life, and the tremor is of a regular rhythmical character.
In the latter the tremor occurs only on voluntary effort, and is also
more regular than the movements of chorea. There are forms of
congenital sclerosis seen in children which closely resemble chorea.
Here the duration of the disease and the association of contractures
with it distinguish it from chorea.

Hysterical subjects have a form of chorea which can only be


differentiated from the true disorder by noting the general hysterical
character of the case and the result of treatment, which strongly
influences the will-power of the patient.

PROGNOSIS.—In the great majority of cases this is favorable. If the


disease occurs in childhood and is without complications, recovery
generally takes place spontaneously after a few weeks. Should the
movements be violent and continuous, so as to interfere with sleep
and the taking of food, or should there be any complication, such as
acute rheumatism or cardiac disorder, then the prospect of recovery
is not so good.

The prognosis as to relapses should be given with caution. If in a


child, it is possible that there will be a return of the disease after a
longer or shorter interval. It is not likely to recur until after several
months, usually at about the same season the following year. As the
child grows older the intervals become longer, and it may safely be
asserted that after puberty is passed and bodily development
completed there will be no more returns of the affection.

The cure is usually complete. It must be remembered, however, that


for some time after an apparently complete cure there may be slight
inco-ordination of movements, particularly in the arms and the face.
These are shown in the unnecessary haste in making uncertain
motions or in slight grimaces, or if excited an awkwardness in the
use of the fingers.

Death is a rare termination in uncomplicated cases, especially in


children. The fatal cases are generally when acute rheumatism has
been associated with the chorea or when there has been a fracture
or an injury as a complication.

In Sée's statistics there is a mortality of 5.7 per cent. in 158 cases in


the Children's Hospital. In adults, and more particularly in pregnant
women, death is more common. Wenzel's cases referred to above
gave a mortality of 27.3 per cent.

The cause of death in chorea may be from the intensity of the


disease, and in this case the symptoms are generally violent from
the outset, increase to an extreme extent, and then collapse and
coma come on. The movements may cease when the collapse
occurs, but they may continue to the last, growing gradually less until
death.

TREATMENT.—A vast number of remedies have been popular in this


disease from all ages. The medicine which is most generally
depended upon at the present day is arsenic. It is advised by most
writers, and in my own experience is decidedly the most reliable
remedy for chorea which we know. The best way to administer it is in
the form of Fowler's solution, and it should be given in large doses. I
have given the bromide of arsenic, but did not find it superior to
Fowler's solution. The amount of arsenic which can be safely borne
by children with chorea is surprising to those who have not had
experience in its administration. The medicine should be given in
gradually increasing doses until the toxic effects are well marked or
until the patient is convalescing. In a child of six years three drops
may be given to begin with, three times a day. One drop additional
should be added to the dose each day, and the child soon acquires a
remarkable tolerance of the drug. As much as twelve or fifteen drops
at a dose is borne by a child of eight years. If vomiting or much
œdema of the face occurs, the medicine should be stopped for a day
or two, and then the original dose should be taken, to be again
increased as before.

Seguin recommends that the patient should begin again with the
dose at which tolerance ceased. For instance, if vomiting occurred
after a dose of nine drops, he stops the medicine for a day, and
begins again with eight drops. I have found that sometimes this
causes vomiting again, and I think it preferable to resume the
medicine with a small dose.

It is often seen that a patient becomes worse during the first few
days that the arsenic is taken, but improvement generally begins
after a week of the arsenical treatment, and is well marked after two
weeks.

In obstinate cases it is of marked advantage to give the arsenic


hypodermically. Cases which do not yield to the drug when given by
the mouth often improve at once when it is given hypodermically.
Chronic cases which have resisted all forms of medication
sometimes are cured by hypodermic injections of arsenic. For giving
arsenic in this way it is best to use Fowler's solution, made without
the compound spirit of lavender. It is less likely to cause abscess to
form at the point of puncture.

Other remedies enjoy a reputation in the treatment of chorea.


Sulphate of zinc is relied upon by many, and it is the means which
Ross recommends. It should be given in increasing doses like
arsenic, and very large doses may often be taken without disturbing
the stomach. Trousseau, Hammond, and Hamilton favor strychnia,
but I have had no experience in its use.

Cimicifuga and conium are both often beneficial in their effects. I


have seen the former do good when arsenic had failed. Conium to
be efficacious must be given in large doses. Eserine and
hyoscyamine have both been successfully employed, the former by
Bouchut, and the latter by Oulmont and Laurent. Recently,
DaCosta33 has reported, in a clinical lecture at the Pennsylvania
Hospital, a case of very severe chorea successfully treated with
hyoscyamine. The patient was a boy of eleven years, and the
disorder had followed an attack of acute rheumatism. He was given
1/100 gr. of hyoscyamine three times a day.

33 Philada. Med. Times, Jan. 23, 1886.

Ziegler34 has recorded several cases which recovered under the use
of nitrite of amyl. The bromides and chloral are useful adjuncts to
treatment in case of sleeplessness or mental irritability. Cases of
cure by the use of chloral alone have been reported. Bouchut gave a
girl of fourteen and a half years, with chorea and dementia, 45 grains
of chloral a day for twenty-seven days. She slept most of the time,
but improvement was seen on the fifth day, and cure was completed
on the twenty-eighth day of the use of the chloral. Electricity has
been efficient in the hands of many writers. I have found
galvanization of the spine to produce a quieting effect in some
cases.
34 Ibid., vol. vi. p. 486.

Iron is always of use in chorea; it may be given during the course of


the disease, and is generally necessary in convalescence. Cod-liver
oil or malt extract should be given in feeble persons.

It is scarcely necessary to mention the other remedies which have


been recommended. DaCosta has used the bromide of iron. H. C.
Wood has used a preparation of skunk cabbage, and there are a
great number of other remedies which have been found of value.

Next to the internal means come external applications. Baths and


frictions are useful in their effect on the general health. The ether
spray to the spine or the application of an ice-bag for ten minutes
once or twice daily is sometimes found to assist the other means.
Cold douches have been advised by some, but they may do harm.
The care of the general health of the patient is of first importance,
and his surroundings should be as quiet as possible.
It is of the greatest value in bad cases to place the patient in bed and
keep him there until the symptoms improve. John Van Bibber of
Baltimore has treated a number of cases of chorea successfully by
keeping them secluded in a darkened room. Such an extreme
degree of isolation is not often necessary, and it might make a child
more nervous.

In children the patient should always be taken from school and kept
from exciting play. Plenty of fresh air and wholesome food should be
insisted upon. Change of air to the mountains or to the seashore
often effects a cure in a short time.

Some cases do not appear to be benefited by any treatment. These


are the hereditary form of chorea and some of the localized choreas.
The latter are often helped or cured by the hypodermic use of
arsenic even in long-standing cases.

ATHETOSIS.
BY WHARTON SINKLER, M.D.

This disease was first described by Hammond in his work on Diseases of the Nervous System in 1871,
and cases have since been reported by many observers, among them Clifford Allbutt, Claye Shaw,
Eulenburg, Oulmont, and Gowers. The disease is named by Hammond from the word ἀθετος, without
fixed position.1 The principal features are an inability to retain the fingers and toes in any position in
which they may be placed, and the continual movements which persist in the parts—a condition called
by Gowers mobile spasm.
1 Diseases of the Nervous System, p. 722.

Athetosis is often connected with impaired mental powers; many of Shaw's cases were in imbecile
children.

The movements of athetosis are not confined to the hand in all cases, but they are sometimes met with
in the foot, and even in the muscles of the face and back.

The following is Hammond's original case:2 “J. P. R——, aged thirty-three, a native of Holland, consulted
Hammond Sept. 13, 1869. His occupation was bookbinding, and he had the reputation, previous to his
present illness, of being a first-class workman. He was of intemperate habits. In 1860 he had an
epileptic paroxysm, and since that time, to the date of his first visit to me, had a fit about once in six
weeks. In 1865 he had an attack of delirium tremens, and for six weeks thereafter was unconscious,
being more or less delirious during the whole period. Soon after recovering his intelligence he noticed a
slight sensation of numbness in the whole of the right upper extremity and in the toes of the same side.
At the same time severe pain appeared in these parts, and complex involuntary movements ensued in
the fingers and toes of the same side.

“At first the movements of the fingers were to some extent under the control of his will, especially when
this was strongly exerted and assisted by his eyesight, and he could, by placing his hand behind him,
restrain them to a still greater degree. He soon, however, found that his labor was very much impeded,
and he had gradually been reduced from time to time to work requiring less care than the finishing, at
which he had been very expert.

“The right forearm, from the continual action of the muscles, was much larger than the other, and the
muscles were hard and developed like those of a gymnast. When told to close his hand he held it out at
arm's length, clasped the wrist with the other hand, and then, exerting all his power, succeeded, after at
least half a minute, in flexing the fingers, but instantaneously they opened again and resumed their
movements.

“In this patient there was impairment of intellect, his memory was enfeebled, and his ideas were dull.
There was no paralysis of any part of the body, but there was slight tremor of both upper extremities.
The involuntary movements were of the right arm, and continued during sleep. Sensation was normal.
The spasm of the muscles causes severe pain in the arm, and keeps him from sleeping at night.”
Hammond used various remedies without relief, and had the patient under his charge for many years.
Finally, he showed the patient to the American Neurological Society at the annual meeting in 1883, with
almost complete relief to the movements as a result of nerve-stretching.
2 Ibid.

Athetosis is found in two forms—the hemiplegic and the bilateral varieties. In the former there has
usually been an attack of hemiplegia more or less marked, or there has been an epileptic fit or
unconsciousness from alcohol, as in Case I. There is often hemianæsthesia or some disorder of

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