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Understanding
Biology
Third Edition
Kenneth A. Mason
University of Iowa
Tod Duncan
University of Colorado Denver
Jonathan B. Losos
William H. Danforth Distinguished University Professor and Director,
Living Earth Collaborative, Washington University
mas40908_FM_i-xxii.indd 1 10/15/19 6:21 PM

A Learning Path to Understanding Biology
Understanding Biology and its online assets have been carefully thought out and crafted to help students and professors work efficiently
and effectively through the material in the course, making the most of study time and furthering instructional goals.
The Learning Path
▲
Every concept is broken down into sections that
cover skills or ideas students should master. Learning
objectives at the beginning of each section help identify
important concepts.
Assessing the Learning Path
Understand 10. Coenzymes

a. can be metal ions.
1. A covalent bond between a hydrogen atom and an oxygen b. can bind in active sites and participate directly in a
atom represents what kind of energy? catalytic reaction.
a. Kinetic energy
▲
c. are sometimes vitamins.
b. Potential energy d. All of the above
c. Mechanical energy
d. Solar energy 11. In competitive inhibition,
Every chapter opens with a Learning Path that walks students 2. During a redox reaction, the molecule that gains an electron is
a. reduced and now has a higher energy level.
a. two enzymes compete with each other for a substrate.
b. an inhibitor molecule binds to an allosteric site on an
enzyme, causing a change in the active site.
through the main concepts in the chapter to enable their

b. oxidized and now has a lower energy level. c. an inhibitor molecule binds to the active site of an
c. reduced and now has a lower energy level. enzyme, so the substrate cannot bind.
d. oxidized and now has a higher energy level. d. the products of a reaction both compete for the active site.
understanding of where the material fits in the context of 3. When a bear eats a salmon, some of the energy stored in the
salmon is used by the bear for its activities and growth. Much
12. Anabolism is
a. the gain of a proton.
of the energy originally in the salmon is dissipated as heat. b. metabolism in animals.
other concepts in the chapter. This is an example of
a. the conversion of kinetic energy to potential energy.
c. the buildup of molecules.
d. the breakdown of molecules.
b. the Second Law of Thermodynamics.
c. 100% efficient energy conversion.
d. a conversion of potential energy to kinetic energy. Apply
4. An endergonic reaction has which of the following properties? 1. When a hibernating animal uses its stored fat to power basic
a. +ΔG and the reaction is spontaneous. body functions (for example, breathing), it is
b. +ΔG and the reaction is not spontaneous. a. converting kinetic energy to potential energy.
c. −ΔG and the reaction is spontaneous. b. converting kinetic energy to chemical energy.
d. −ΔG and the reaction is not spontaneous. c. converting potential energy to kinetic energy.
5. Where is the energy stored in a molecule of ATP? d. converting chemical energy to potential energy.
a. Within the bonds between nitrogen and carbon 2. During certain stages of cellular respiration, electrons are
b. In the carbon-to-carbon bonds found in the ribose transferred from glucose molecules to a molecule called
c. In the phosphorus-to-oxygen double bond nicotinamide adenine dinucleotide (NAD+). During this
d. In the bonds connecting the two terminal phosphate groups reaction,
6. The molecule ATP is less stable than ADP + Pi because a. glucose is oxidized and NAD+ is reduced.
a. the negatively charged phosphates repel each other. b. glucose is reduced and NAD+ is oxidized.
b. the positively charged phosphates repel each other. c. both glucose and NAD+ have gained protons.
c. ATP is much larger than ADP and Pi. d. glucose has gained protons and NAD+ has lost protons.
d. the adenine in ATP is charged. 3. Sodium ions (Na+) can move through channel proteins
7. What is activation energy? across some biological membranes. If Na+ is present in a
a. The thermal energy associated with random movements of higher concentration on one side of a membrane, the ions
molecules will tend to move across the membrane until they are equally
b. The energy released through breaking chemical bonds distributed on both sides of the membrane. This process
c. The difference in free energy between reactants and products a. results in a gain of potential energy for the cell.
d. The energy required to initiate a chemical reaction b. results in a decrease in entropy.
c. follows the Second Law of Thermodynamics.
8. Which of the following is NOT a property of an enzyme?
d. All of the above
a. An enzyme reduces the activation energy of a reaction.
4. If the products of a chemical reaction have higher free energy
b. An enzyme lowers the free energy of the reactants. than the reactants, this reaction
c. An enzyme does not change as a result of the reaction. a. will not proceed spontaneously.
d. An enzyme works in both the forward and reverse b. will proceed spontaneously.
directions of a reaction. c. must have increased the total energy in the universe.
9. Which statement about the influence of temperature on d. must have decreased the total energy in the universe.
enzymes is NOT true? 5. ATP can be used to drive an endergonic reaction if
a. All enzymes have the same intrinsic optimal temperature. a. ATP hydrolysis lowers the activation energy for the reaction.
b. Raising the temperature may increase the activity of an b. the sum of ΔG for ATP hydrolysis and ΔG for the reaction
enzyme. is negative.
c. Some enzymes are stable at the boiling point of water. c. the sum of ΔG for ATP hydrolysis and ΔG for the reaction
d. Raising the temperature may decrease the activity of an is positive.
enzyme. d. ATP hydrolysis raises the activation energy for the reaction.
Chapter 6 Energy and Metabolism 129
▲
Chapter concepts are assessed at three different levels at the
end of the chapter. On a first pass through the chapter prior to
▲ class, students might focus on the “Understand” level. As they
At the end of each section, Review of Concept questions gain greater mastery of the material, they should challenge
allow students to check their understanding before themselves with “Apply” and “Synthesize” questions that
moving on to the next concept. require higher cognitive skills.
viii

Think Like a Scientist
Scientific Thinking figures throughout Inquiry & Analysis
▲
the text walk the student through features at the ends of
a scientific experiment, laying out all chapters explore a
the Hypothesis, Predictions, Test scientific investigation in
Procedures, Results, and Conclusion. more detail, presenting
Some also challenge the students to experimental results and
devise further experiments. challenging students to
▲
interpret the data.
Connecting the Concept

Building on the chapter features, the “Connecting the
▲
Concepts” at the end of each Part in the text shows how

seemingly isolated concepts in different chapters are
connected by core concepts of biology.
▲
The end-of-chapter “Connecting the Concepts”
identifies a core concept represented by a gear icon.
Secondary concepts are placed on the cogs, and a list
of observations from the chapter connects the
secondary concept to the core concept. A second gear
or cog is presented that lacks the list of observations.
The student is challenged to complete the open concept
with supporting information from the chapter.
a learning path to understanding biology ix

Strengthen Problem-Solving Skills with Connect®
Detailed Feedback in Connect® framework to scaffold learning, where each step models and
reinforces the learning process.
Learning is a process of iterative development, of making
The feedback for each higher-level Bloom’s question (Apply,
mistakes, reflecting, and adjusting over time. The question and
Analyze, Evaluate) follows a similar process: Clarify Question,
test banks in Connect® for Understanding Biology, 3rd edition,
Gather Content, Choose Answer, Reflect on Process.
are more than direct assessments; they are self-contained
learning experiences that systematically build student learning
over time. Unpacking the Concepts
For many students, choosing the right answer is not We’ve taken problem solving a step further. In each chapter,
necessarily based on applying content correctly; it is more two higher-level Bloom’s questions in the question and test
a matter of increasing their statistical odds of guessing. A banks are broken out by the steps of the detailed feedback.
major fault with this approach is students don’t learn how to Rather than leaving it up to the student to work through
process the information correctly, mostly because they are the Socratic questions, a second version of the question
repeating and reinforcing their mistakes rather than reflecting is presented in a stepwise format. Following the problem-
and learning from them. To help students develop problem- solving steps, students need to answer questions about
solving skills, all higher-level Bloom’s questions in Connect earlier steps, such as “What is the key concept addressed by
are supported with hints, to help students focus on important the question?” before proceeding to answer the question. A
information for answering the questions, as well as detailed professor can choose which version of the question to include
feedback that walks students through the problem-solving in the assignment based on the problem-solving skills of the
process, using Socratic questions in a decision-tree-style students.
x a learning path to understanding biology

Data and Graphing Interactives scientific problem and the opportunity to manipulate variables,
® producing different results on a graph, analyze and evaluate
To help students develop analytical skills, Connect for
data, or view different aspects of the problem. A series of
Understanding Biology, 3rd edition, is enhanced with interactive
questions follows the graphing activity to assess if the student
data and graphing questions. Students are presented with a
understands and is able to interpret the data and results.
Quantitative Question Bank

any chapters also contain a Quantitative Question Bank.
M
These are more challenging algorithmic questions, intended to
help students practice their quantitative reasoning skills. Hints
and guided solution options walk students through a problem,
and then an alternative problem is provided for them to test their
understanding.
a learning path to understanding biology xi

®
FOR INSTRUCTORS
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Want to build your own course? No problem. Prefer to use our turnkey,
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Contents
About the Authors iv
PART II Biology
Changes to This Edition vi
of the Cell 66
A Learning Path to Understanding Biology viii
©Dr. Gopal Murti/Science Source
Acknowledgments xxi 4 Cell Structure 66
Concept 4.1 All Living Organisms Are Composed of Cells 67
Concept 4.2 Prokaryotic Cells Are Relatively Simple 70
PART I The Molecular Concept 4.3 Eukaryotic Cells Are Highly Compartmentalized 72
Concept.4.4 Membranes Organize the Cell Interior into Functional
Basis of Life 1 Compartments 76
©Soames Summerhays/Natural Visions Concept 4.5 Mitochondria and Chloroplasts Are Energy-Processing
Organelles 80
1 The Science of Biology 1
Concept 4.6 An Internal Skeleton Supports the Shape of Cells 82
Concept 1.1 The Diversity of Life Is Overwhelming 2 Concept 4.7 Extracellular Structures Protect Cells 85
Concept 1.2 Biology Is the Science of Life 3 Concept 4.8 Cell-to-Cell Connections Determine How Adjacent
Concept 1.3 Science Is Based on Both Observation and Cells Interact 86
Reasoning 6 Inquiry & Analysis 90
Concept 1.4 The Study of Evolution Is a Good Example of Retracing the Learning Path 91
Scientific Inquiry 9
Assessing the Learning Path 92
Concept 1.5 A Few Important Concepts Form the Core of
Biology 14
Inquiry & Analysis 17 5 Membranes 94
Retracing the Learning Path 18 Concept 5.1 Membranes Are Phospholipid Bilayers with
Assessing the Learning Path 19 Embedded Proteins 95
Concept 5.2 Phospholipids Provide a Membrane’s Structural
Foundation 98
1.5 2 The Nature of Molecules and the
Concept 5.3 Membrane Proteins Enable a Broad Range of
Properties of Water 21 Interactions with the Environment 99
Concept 2.1 All Matter Is Composed of Atoms 22 Concept 5.4 Passive Transport Moves Molecules Across
Membranes by Diffusion 101
Concept 2.2 The Elements in Living Systems Have Low Atomic
Masses 25 Concept 5.5 Active Transport Across Membranes Requires
Energy 105
Concept 2.3 Molecules Are Collections of Atoms Held Together
by Chemical Bonds 27 Concept 5.6 Bulky Materials Cross Membranes Within
Vesicles 108
Concept 2.4 The Properties of Water Result from Its Polar Nature 30
Inquiry & Analysis 111
Concept 2.5 Water Molecules Can Dissociate into Ions 34
Retracing the Learning Path 112
6 Energy and Metabolism 115
3 The Chemical Building Blocks of Life 40 Concept 6.1 Energy Flows Through Living Systems 116
Concept 3.1 Carbon Provides the Framework of Biological Concept 6.2 The Laws of Thermodynamics Govern All Energy
Molecules 41 Changes 117
Concept 3.2 Carbohydrates Form Both Structural and Energy-Storing Concept 6.3 ATP Is the Energy Currency of Cells 119
Molecules 43 Concept 6.4 Enzymes Speed Up Reactions by Lowering Activation
Concept 3.3 Proteins Are the Tools of the Cell 47 Energy 121
Concept 3.4 Nucleic Acids Store and Express Genetic Concept 6.5 Metabolism Is the Sum of a Cell’s Chemical
Information 55 Activities 125
Concept 3.5 Hydrophobic Lipids Form Fats and Membranes 58 Inquiry & Analysis 127
Inquiry & Analysis 61 Retracing the Learning Path 128
Retracing the Learning Path 62 Assessing the Learning Path 129
xiv

7 How Cells Harvest Energy 131 10 How Cells Divide 198
Concept 7.1 Cells Harvest Energy from Organic Compounds by Concept 10.1 Bacterial Cell Division Is Clonal 199
Oxidation 132 Concept 10.2 Eukaryotes Have Large, Linear Chromosomes 200
Concept 7.2 Glycolysis Splits Glucose and Yields a Small Amount Concept 10.3 The Eukaryotic Cell Cycle Is Complex and Highly
of ATP 136 Organized 203
Concept 7.3 The Citric Acid Cycle Is the Oxidative Core of Concept 10.4 During Interphase, Cells Grow and Prepare for
Cellular Respiration 139 Mitosis 204
Concept 7.4 Electrons Removed by Oxidation Pass Along an Concept 10.5 In Mitosis, Chromosomes Segregate 205
Electron Transport Chain 142
Concept 10.6 Events of the Cell Cycle Are Carefully
Concept 7.5 The Total Energy Yield of Aerobic Respiration Far Regulated 210
Exceeds That of Glycolysis 146
Concept 10.7 Cancer Is a Failure of Cell-Cycle Control 214
Concept 7.6 Aerobic Respiration Is Regulated by Feedback
Inhibition 147 Inquiry & Analysis 217
Concept 7.7 Some Forms of Energy Metabolism Do Not Require Retracing the Learning Path 218
O2 148 Assessing the Learning Path 219
Concept 7.8 Carbohydrates Are Not the Only Energy Source Used
by Heterotrophs 149
Retracing the Learning Path 153 PART III Genetics and
Assessing the Learning Path 154 Molecular Biology 222
©McGraw-Hill Education/Steven P. Lynch
8 Photosynthesis 156
11 Sexual Reproduction and Meiosis 222
Concept 8.1 Photosynthesis Uses Sunlight to Power the Synthesis
of Organic Molecules 157 Concept 11.1 Sexual Reproduction Requires Meiosis 223
Concept 8.2 Experiments Revealed That Photosynthesis Is a Concept 11.2 Meiosis Consists of Two Divisions with One Round
Chemical Process 159 of DNA Replication 224
Concept 8.3 Pigments Capture Energy from Sunlight 160 Concept 11.3 The Process of Meiosis Involves Intimate Interactions
Concept 8.4 Photosynthetic Pigments Are Organized into Between Homologues 225
Photosystems 163 Concept 11.4 Meiosis Has Four Distinct Features 230
Concept 8.5 Energy from Sunlight Is Used to Produce a Proton Concept 11.5 Genetic Variation Is the Evolutionary Consequence
Gradient 165 of Sex 232
Concept 8.6 Using ATP and NADPH from the Light Reactions, Inquiry & Analysis 234
CO2 Is Incorporated into Organic Molecules 170 Retracing the Learning Path 235
Concept 8.7 Photorespiration Short-Circuits Photosynthesis 172 Assessing the Learning Path 236
Retracing the Learning Path 176 11 12 Patterns of Inheritance 238
Concept 12.1 Experiments Carried Out by Mendel Explain
Heredity 239
9 Cell Communication 179 Concept 12.2 Mendel’s Principle of Segregation Accounts
Concept 9.1 The Cells of Multicellular Organisms for 3:1 Phenotypic Ratios 241
Communicate 180 Concept 12.3 Mendel’s Principle of Independent
Concept 9.2 Signal Transduction Begins with Cellular Assortment Asserts That Genes Segregate
Receptors 182 Independently 245
Concept 9.3 Intracellular Receptors Respond to Signals by Concept 12.4 Probability Allows Us to Predict the Results of
Regulating Gene Expression 184 Crosses 246
Concept 9.4 Protein Kinase Receptors Respond to Signals by Concept 12.5 Genotype Dictates Phenotype by Specifying Protein
Phosphorylating Proteins 185 Sequences 248
Concept 9.5 G Protein–Coupled Receptors Respond to Signals Concept 12.6 Extending Mendel’s Model Provides a Clearer View of
Through Effector Proteins 189 Genetics in Action 249
Inquiry & Analysis 194 Inquiry & Analysis 255
Retracing the Learning Path 195 Retracing the Learning Path 256
Assessing the Learning Path 196 Assessing the Learning Path 257
contents xv

13 The Chromosomal Basis of 16 Control of Gene Expression 332
Inheritance 259 Concept 16.1 All Organisms Control Expression of Their
Genes 333
Concept 13.1 Chromosomes Are the Vehicles of Mendelian
Inheritance 260 Concept 16.2 Regulatory Proteins Control Genes by Interacting
with Specific DNA Nucleotide Sequences 334
Concept 13.2 Some Genes Do Not Assort Independently:
Linkage 263 Concept 16.3 Prokaryotes Regulate Their Genes in Clusters 336
Concept 13.3 Genetic Crosses Provide Data for Genetic Maps 265 Concept 16.4 Transcription Factors Control Gene Transcription
in Eukaryotes 340
Concept 13.4 Changes in Chromosome Number Can Have Drastic
Effects 267 Concept 16.5 Chromatin Structure Affects Gene Expression 344
Concept 13.5 Inheritance in Humans Can Be Studied by Analyzing Concept 16.6 Eukaryotic Genes Are Also Regulated After
Pedigrees 268 Transcription 345
Concept 13.6 There Are Two Major Exceptions to Chromosomal Concept 16.7 Gene Regulation Determines How Cells Will
Inheritance 273 Develop 351
14 DNA: The Genetic Material 280 17 Biotechnology 358

Concept 14.1 DNA Is the Genetic Material 281 Concept 17.1 Enzymes Allow the Creation of Recombinant
Molecules In Vitro 359
Concept 14.2 The DNA Molecule Is a Double Helix 283
Concept 17.2 The Polymerase Chain Reaction Is Used to Amplify
Concept 14.3 Both Strands Are Copied During DNA Specific DNA Sequences 363
Replication 287
Concept 17.3 Molecular Tools Allow Us to Analyze and Modify
Concept 14.4 Prokaryotes Organize the Enzymes Used to Duplicate Genetic Variation 366
DNA 290
Concept 17.4 Transgenic Organisms Are Used to Analyze Gene
Concept 14.5 Eukaryotic Chromosomes Are Large and Linear 295 Function 368
Concept 14.6 Cells Repair Damaged DNA 297 Concept 17.5 Genetic Tools Are Changing Modern Medicine 371
Inquiry & Analysis 300 Concept 17.6 Genetic Engineering Is Used in Industry and
Retracing the Learning Path 301 Agriculture 374
Assessing the Learning Path 302 Inquiry & Analysis 378
15 Genes and How They Work 304 Assessing the Learning Path 380
Concept 15.1 Experiments Have Revealed the Nature of
Genes 305 18 Genomics 382
Concept 15.2 The Genetic Code Relates Information in DNA Concept 18.1 Mapping Identifies and Locates Functional Elements
and Protein 307 in Genomes 383
Concept 15.3 Prokaryotes Exhibit All the Basic Features of Concept 18.2 The Modernization of DNA Sequencing Has
Transcription 310 Accelerated Discovery 386
Concept 15.4 Eukaryotes Use Three Polymerases and Extensively Concept 18.3 Genome Projects Reveal Insights into Medicine and
Modify Transcripts 313 Agriculture 389
Concept 15.5 Eukaryotic Genes May Contain Noncoding Concept 18.4 Genome Annotation Assigns Functional
Sequences 315 Information to Genomes 391
Concept 15.6 The Ribosome Is the Machine of Protein Concept 18.5 Genome Comparisons Provide Information About
Synthesis 317 Genomic Structure and Function 394
Concept 15.7 The Process of Translation Is Complex and Energy- Concept 18.6 Comparative Genomics Informs Evolutionary
Expensive 320 Biology 399
Concept 15.8 Mutations Are Alterations in the Sequence, Number, Inquiry & Analysis 403
or Position of Genes 325
xvi contents

Concept 21.6 The Pace of Evolution Varies 471
Concept 21.7 Speciation and Extinction Have Molded Biodiversity
PART IV Evolution 408 Through Time 472
©Tetra Images/Getty Images Retracing the Learning Path 475
19 Genes Within Populations 408 Assessing the Learning Path 476
Concept 19.1 Natural Populations Exhibit Genetic Variation 409
Concept 19.2 Frequencies of Alleles Can Change 411
Concept 19.3 Five Agents Are Responsible for Evolutionary
Change 413 PART V The Diversity
Concept 19.4 Selection Can Act on Traits Affected by Many
Genes 418
of Life 479
©Imagemore Co, Ltd./Getty Images
Concept 19.5 Natural Selection Can Be Studied
Experimentally 420 22 Systematics and Phylogeny 479
Concept 19.6 Fitness Is a Measure of Evolutionary Success 422 Concept 22.1 Systematics Reconstructs Evolutionary
Concept 19.7 Evolutionary Processes Sometimes Maintain Relationships 480
Variation 423 Concept 22.2 Cladistics Focuses on Traits Derived from a
Concept 19.8 Sexual Selection Determines Reproductive Common Ancestor 481
Success 426 Concept 22.3 Classification Is a Labeling Process, Not an
Inquiry & Analysis 429 Evolutionary Reconstruction 485
Retracing the Learning Path 430 Concept 22.4 Taxonomy Attempts to Classify Organisms in an
Assessing the Learning Path 431 Evolutionary Context 488
Concept 22.5 The Largest Taxa Are Domains 490
20 The Evidence for Evolution 433 Inquiry & Analysis 495
Concept 20.1 The Beaks of Darwin’s Finches Provide Evidence of Retracing the Learning Path 496
Natural Selection 434 Assessing the Learning Path 497
Concept 20.2 Peppered Moths and Industrial Melanism Illustrate
Natural Selection in Action 437 23 Prokaryotes and Viruses 499
Concept 20.3 Human-Initiated Artificial Selection Is Also a Concept 23.1 Prokaryotes Are the Most Ancient Organisms 500
Powerful Agent of Change 439
Concept 23.2 Prokaryotes Have an Organized but Simple
Concept 20.4 Fossils Provide Direct Evidence of Evolution 441 Structure 503
Concept 20.5 Anatomical Evidence for Evolution Is Extensive Concept 23.3 The Genetics of Prokaryotes Focuses on DNA
and Persuasive 444 Transfer 507
Concept 20.6 Genes Carry a Molecular Record of the Evolutionary Concept 23.4 Prokaryotic Metabolism Is Diverse 511
Past 447
Concept 23.5 Bacteria Cause Important Human Diseases 511
Concept 20.7 Natural Selection Favors Convergent Evolution in
Similar Environments 448 Concept 23.6 Viruses Are Not Organisms 513
Concept 20.8 Addressing Common Criticisms of Evolutionary Concept 23.7 Bacterial Viruses Infect by DNA Injection 515
Theory 450 Concept 23.8 Animal Viruses Infect by Endocytosis 517
21 The Origin of Species 456 24 Protists 524

Concept 21.1 The Biological Species Concept Highlights Concept 24.1 Protists, the First Eukaryotes, Arose by
Reproductive Isolation 457 Endosymbiosis 525
Concept 21.2 Natural Selection May Reinforce Reproductive Concept 24.2 Protists Are a Very Diverse Group 527
Isolation 462 Concept 24.3 The Rough Outlines of Protist Phylogeny Are
Concept 21.3 Natural Selection and Genetic Drift Play Key Roles in Becoming Clearer 529
Speciation 463 Concept 24.4 Excavata Are Flagellated Protists Lacking
Concept 21.4 Speciation Is Influenced by Geography 464 Mitochondria 530
Concept 21.5 Adaptive Radiation Requires Both Speciation and Concept 24.5 SAR: Stramenopiles and Alveolates Exhibit Secondary
Habitat Diversity 466 Endosymbiosis 533
contents xvii

Concept 24.6 SAR: Rhizaria Have Silicon Exoskeletons or Limestone Concept 27.5 Mollusks and Annelids Are the Largest Groups
Shells 539 of Lophotrochozoans 609
Concept 24.7 Archaeplastida Are Descended from a Single Concept 27.6 Lophophorates Are Very Simple Marine Organisms 613
Endosymbiosis Event 540 Concept 27.7 Nematodes and Arthropods Are Both Large Groups
Concept 24.8 Amoebozoa and Opisthokonta Are Closely of Ecdysozoans 614
Related 543 Concept 27.8 Deuterostomes Are Composed of Echinoderms and
Inquiry & Analysis 546 Chordates 619
Retracing the Learning Path 547 Inquiry & Analysis 622
Assessing the Learning Path 548 Retracing the Learning Path 623
25 Fungi 550
Concept 25.1 Fungi Have Unique Reproductive and Nutritional 28 Vertebrates 626
Strategies 551 Concept 28.1 Nonvertebrate Chordates Do Not Form Bone 627
Concept 25.2 Fungi Have an Enormous Ecological Impact 554 Concept 28.2 Almost All Chordates Are Vertebrates 628
Concept 25.3 Fungi Are Important Plant and Animal Concept 28.3 Fishes Are the Earliest and Most Diverse
Pathogens 557 Vertebrates 629
Concept 25.4 Fungi Are Taxonomically Diverse 558 Concept 28.4 Amphibians Are Moist-Skinned Descendants of the
Concept 25.5 Microsporidia Are Unicellular Parasites 559 Early Tetrapods 633
Concept 25.6 Chytridiomycota and Relatives: Fungi with Concept 28.5 Reptiles Are Fully Adapted to Terrestrial Living 635
Zoospores 560 Concept 28.6 Birds Are Essentially Flying Reptiles 639
Concept 25.7 Zygomycota Produce Zygotes 561 Concept 28.7 Mammals Are the Least Diverse of Vertebrates 642
Concept 25.8 Glomeromycota Are Asexual Plant Symbionts 562 Concept 28.8 Primates Include Lemurs, Monkeys, Apes, and
Concept 25.9 Basidiomycota Are the Mushroom Fungi 563 Humans 645
Concept 25.10 Ascomycota Are the Most Diverse Phylum of Inquiry & Analysis 652
Fungi 564 Retracing the Learning Path 653
Inquiry & Analysis 567 Assessing the Learning Path 654
26 Plants 571 PART VI Plant Form

Concept 26.1 Land Plants Are Multicellular Autotrophs Adapted to and Function 657
Terrestrial Life 572
©Susan Singer
Concept 26.2 Bryophytes Have a Dominant Gametophyte
Generation 574 29 Plant Form 657
Concept 26.3 Seedless Vascular Plants Evolved Roots, Stems, Concept 29.1 Meristems Articulate the Body Plan 658
and Leaves 577
Concept 29.2 Plants Contain Three Principal Tissues 661
Concept 26.4 Lycophytes Have a Dominant Sporophyte
Generation 578 Concept 29.3 Roots Have Four Growth Zones 666
Concept 26.5 Pterophytes Are Ferns and Their Relatives 579 Concept 29.4 Stems Provide Support for Aboveground Organs 671
Concept 26.6 Seed Plants Were a Key Step in Plant Evolution 583 Concept 29.5 Leaves Are a Plant’s Photosynthetic Organs 675
Concept 26.7 Gymnosperms Are Plants with “Naked Seeds” 583 Inquiry & Analysis 678
Concept 26.8 Angiosperms Are Flowering Plants 587 Retracing the Learning Path 679
Inquiry & Analysis 591 Assessing the Learning Path 680
30 Flowering Plant Reproduction 682
Concept 30.1 Reproduction Starts with Flowering 683
27 Animal Diversity 595 Concept 30.2 Flowers Attract Pollinators 685
Concept 27.1 The Diversity of Animal Body Plans Arose by a Series Concept 30.3 Fertilization Leads to Embryogenesis 690
of Evolutionary Innovations 596 Concept 30.4 Seeds Protect Angiosperm Embryos 692
Concept 27.2 Molecular Data Are Clarifying the Animal Concept 30.5 Fruits Promote Seed Dispersal 695
Phylogenetic Tree 602 Concept 30.6 Germination Begins Seedling Growth 697
Concept 27.3 True Tissue Evolved in Simple Animals 605 Concept 30.7 Plant Life Spans Vary Widely 699
Concept 27.4 Flatworms and Rotifers Are Very Simple Concept 30.8 Asexual Reproduction Is Common Among Flowering
Bilaterians 607 Plants 700
xviii contents

Inquiry & Analysis 702 Concept 33.6 The Peripheral Nervous System Consists of Both
Retracing the Learning Path 703 Sensory and Motor Neurons 772
Assessing the Learning Path 704 Concept 33.7 Sensory Receptors Provide Information About the
Body’s Environment 774
Concept 33.8 Mechanoreceptors Sense Touch and Pressure 775
31 The Living Plant 706
Concept 33.9 Sounds and Body Position Are Sensed by Vibration
Concept 31.1 Water Moves Through Plants Based on Detectors 776
Potential Differences 707
Concept 33.10 Taste, Smell, and pH Senses Utilize Chemoreceptors 779
Concept 31.2 Roots Absorb Minerals and Water 710
Concept 33.11 Vision Employs Photoreceptors to Perceive Objects at
Concept 31.3 Xylem Transports Water from Root to Shoot 711 a Distance 780
Concept 31.4 Transpiration Rate Reflects Environmental Inquiry & Analysis 785
Conditions 715
Concept 31.5 Plants Are Adapted to Water Stress 716
Concept 31.6 Phloem Transports Organic Molecules 717
Concept 31.7 Plants Require a Variety of Nutrients 719 34 Fueling the Body’s Metabolism 789
Concept 31.8 Plants Use Hormones to Regulate Growth 720
Concept 34.1 Vertebrate Digestive Systems Are Tubular Tracts 790
Concept 31.9 Plant Growth Is Responsive to Light 725
Concept 34.2 Food Is Processed As It Passes Through the Digestive
Concept 31.10 Plant Growth Is Sensitive to Gravity 728 Tract 791
Inquiry & Analysis 731 Concept 34.3 The Digestive Tract Is Regulated by the Nervous
Retracing the Learning Path 732 System and Hormones 796
Assessing the Learning Path 733 Concept 34.4 Respiratory Systems Promote Efficient Exchange of
Gases 797
Concept 34.5 Gills Provide for Efficient Gas Exchange in Water 798
Concept 34.6 Lungs Are the Respiratory Organs of Terrestrial
PART VII Animal Form Vertebrates 800
and Function 736 Concept 34.7 Oxygen and Carbon Dioxide Are Transported by
Fundamentally Different Mechanisms 803
©Stockbyte Concept 34.8 Circulating Blood Carries Metabolites and Gases
32 The Animal Body and How It Moves 736 to the Tissues 806
Concept 34.9 Vertebrate Circulatory Systems Put a Premium on
Concept 32.1 The Vertebrate Body Has a Hierarchical Efficient Circulation 808
Organization 737
Concept 34.10 The Four Chambers of the Heart Contract in a
Concept 32.2 Epithelial Tissue Covers Body Surfaces 739 Cycle 811
Concept 32.3 Nerve Tissue Conducts Signals Rapidly 741 Concept 34.11 The Circulatory Highway Is Composed of Arteries,
Concept 32.4 Connective Tissue Supports the Body 742 Capillaries, and Veins 814
Concept 32.5 Muscle Tissue Powers the Body’s Movements 743 Inquiry & Analysis 817
Concept 32.6 Skeletal Systems Anchor the Body’s Muscles 745 Retracing the Learning Path 818
Concept 32.7 Vertebrate Endoskeletons Are Made of Bone 746 Assessing the Learning Path 819
Concept 32.8 Muscles Contract Because Their Myofilaments
Shorten 750 35 Maintaining Homeostasis 821
Concept 32.9 Animal Locomotion Takes Many Forms 754
Concept 35.1 Homeostasis Maintains a Constant Internal
Inquiry & Analysis 757 Environment 822
Retracing the Learning Path 758 Concept 35.2 Hormones Are Chemical Messages That Direct Body
Assessing the Learning Path 759 Processes 826
Concept 35.3 The Pituitary and the Hypothalamus Are the Body’s
33 The Nervous System 761 Control Centers 830
Concept 35.4 Peripheral Endocrine Glands Play Major Roles in
Concept 33.1 The Nervous System Directs the Body’s Actions 762 Homeostasis 833
Concept 33.2 Neurons Maintain a Resting Potential Across the Concept 35.5 Animals Are Osmoconformers or Osmoregulators 837
Plasma Membrane 763
Concept 35.6 The Kidney Maintains Osmotic Homeostasis in
Concept 33.3 Action Potentials Propagate Nerve Impulses 765 Mammals 838
Concept 33.4 Synapses Are Where Neurons Communicate with Concept 35.7 Hormones Control Osmoregulation 843
Other Cells 767
Concept 35.8 The Immune System Defends the Body 844
Concept 33.5 The Central Nervous System Includes the Brain and
Spinal Cord 770 Concept 35.9 Cell-Mediated Immunity Involves Helper and Killer
T Cells 849
contents xix

Concept 35.10 In Humoral Immunity, B Cells Produce Protective Concept 38.2 Population Growth Depends upon Members’ Age and
Antibodies 851 Sex 921
Inquiry & Analysis 855 Concept 38.3 Evolution Favors Life Histories That Maximize
Retracing the Learning Path 856 Lifetime Reproductive Success 924
Assessing the Learning Path 857 Concept 38.4 Environment Limits Population Growth 926
Concept 38.5 Resource Availability Regulates Population
Growth 928
36 Reproduction and Development 859
Concept 38.6 Earth’s Human Population Is Growing
Concept 36.1 Mammals Are Viviparous 860 Explosively 931
Concept 36.2 The Human Male Reproductive System Is Typical of Inquiry & Analysis 935
Mammals 861
Concept 36.3 The Human Female Reproductive System Undergoes
Cyclic Gamete Development 864
Concept 36.4 The First Step in Development Is Fertilization 869
39 Community Ecology and Ecosystem
Concept 36.5 Cells of the Early Embryo Are Totipotent 872
Concept 36.6 Cleavage Leads to the Blastula Stage 876 Dynamics 939
Concept 36.7 Gastrulation Forms the Basic Body Plan of the Concept 39.1 Competition Shapes How Species Live Together in
Embryo 878 Communities 940
Concept 36.8 The Body’s Organs Form in Organogenesis 880 Concept 39.2 Predator–Prey Relationships Foster Coevolution 944
Concept 36.9 Human Development Takes Nine Months 883 Concept 39.3 Cooperation Among Species Can Lead to
Inquiry & Analysis 887 Coevolution 948
Retracing the Learning Path 888 Concept 39.4 Ecological Succession Is a Consequence of
Habitat Alteration 953
Concept 39.5 Chemical Elements Move Through Ecosystems in
Biogeochemical Cycles 955
Concept 39.6 Energy Flows Through Ecosystems in One
Direction 960
PART VIII Ecology Concept 39.7 Biodiversity May Increase Ecosystem Stability 965
and Behavior 892 Inquiry & Analysis 969
©Mike Powles/Getty Images Retracing the Learning Path 970
37 Behavioral Biology 892
Concept 37.1 An Animal’s Genome Influences Its Behavior 893 40 The Living World 973
Concept 37.2 Learning Also Influences Behavior 895
Concept 40.1 Ecosystems Are Shaped by Sun, Wind, and
Concept 37.3 Thinking Directs the Behavior of Many Animals 896 Water 974
Concept 37.4 Migratory Behavior Is Both Innate and Learned 897 Concept 40.2 Earth Has 14 Major Terrestrial Ecosystems,
Concept 37.5 Animal Communication Plays a Key Role in Called Biomes 978
Ecological and Social Behavior 899 Concept 40.3 Freshwater Habitats Occupy Less Than 2% of Earth’s
Concept 37.6 Behavior Evolves Adaptively 902 Surface 980
Concept 37.7 Behavioral Ecology Examines the Adaptive Concept 40.4 Marine Habitats Dominate the Earth 981
Significance of Behaviors 903 Concept 40.5 Humanity’s Pollution and Resource Depletion Are
Concept 37.8 Behavioral Strategies Have Evolved to Maximize Severely Impacting the Biosphere 983
Reproductive Success 906 Concept 40.6 Human Activity Is Altering Earth’s Climate 986
Concept 37.9 Some Behaviors Decrease Fitness to Benefit Other Inquiry & Analysis 990
Individuals 908
Concept 37.10 Group Living Has Evolved in Both Insects and
Vertebrates 911 Assessing the Learning Path 9926
Retracing the Learning Path 914 Appendix A: Answer Key A-1
Index I-1
38 Ecology of Individuals and
Populations 917
Concept 38.1 Populations Are Groups of a Single Species in One
Place 918
xx contents

Acknowledgments
A revision of this scope relies on the talents and efforts Their feedback shaped this edition. All of these people took
of many people working behind the scenes, and we have time to share their ideas and opinions to help us build a better
benefited greatly from their assistance. edition of Understanding Biology for the next generation of
The copy editor, Linda Davoli labored many hours and introductory biology students, and they have our heartfelt
always improved the clarity and consistency of the text. thanks.
She has made a tremendous contribution to the quality
of the final product. We were fortunate to work with MPS Reviewers for Understanding Biology, 3rd edition
Limited to update the art program and improve the layout of Peggy L. Anderson Navarro College
the pages. Our close collaboration resulted in a text that is Michelle Bell Boissiere Xavier University of Louisiana
pedagogically effective as well as more beautiful than any
Pengli Bu St. John’s University
other biology text on the market.
Don Campanella College of Southern Idaho
We have the continued support of an excellent team at Mary Colon Seminole State College of Florida
McGraw-Hill Education. Andrew Urban the Portfolio Manager
Abdeslem El Idrissi City University of New York / CSI
for Biology, has been a steady leader during a time of change.
Fleur Ferro Community College of Denver
Senior Product Developer, Liz Sievers, provided support in so
many ways it would be impossible to name them all. Kelly Hart, Judyth Gulden Tulsa Community College
Senior Content Project Manager, and Matt Diamond, Designer, Bethany Henderson-Dean University of Findlay
ensured our text was elegantly designed. Kelly Brown, Senior, Linda Johnson University of Maryland Eastern Shore
Marketing Manager, is always a sounding board for more than Gregory Jones Santa Fe College
just marketing, and many more people behind the scenes have Vladimir Jurukovski Suffolk County Community College
all contributed to the success of our text. This includes the Xu Lu University of Findlay
digital team, to whom we owe a great deal for their efforts to
Brad Mehrtens University of Illinois at Urbana-Champaign
continue improving our Connect assessment tools.
Cecile Mioni California State University Monterey Bay
Throughout this edition we have had the support of spouses Mary Phillips Tulsa Community College
who have seen less of us than they might have liked because
Susan Phillips Eastern Florida State College
of the pressures of getting this revision completed. They
Kathleen M. Raley-Susman Vassar College
have adapted to the many hours this book draws us away
from them, and, even more than us, looked forward to its Laurel Roberts University of Pittsburgh
completion. Jeremy D. Rentsch Francis Marion University
In the end, the people we owe the most are the generations Mr. Connie Rye East Mississippi Community College
of students who have passed through our lecture halls. They Robert E. Shinn East Mississippi Community College, Scooba
have taught us at least as much as we have taught them, and Campus
their questions and suggestions continue to improve the text Kaia Skaggs University of Findlay
and supplementary materials. Kim Spaccarotella Kean University
Finally, we need to thank instructors from across the country Tara N. Turley-Stoulig Southeastern Louisiana University
who are continually sharing their knowledge and experience Anthea Warren-Williams Lawson State Community College
with us through market feedback, reviews, and symposia. Michelle Wilson The University of Texas at Dallas
xxi

Part I The Molecular Basis of Life
1The Science of Biology
Lea r ni ng Pa t h
1.1 The Diversity of Life Is Overwhelming
1.2 Biology Is the Science of Life
Science Is Based on Both Observation

1.3
and Reasoning
The Study of Evolution Is a Good Example

1.4
of Scientific Inquiry
A Few Important Concepts Form the

1.5
Core of Biology
s
on
V isi
tu ral
ys /Na
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es Sum
©S oam
mas40908_ch01_001-020.indd 1 10/10/19 2:08 PM

Introduction
Y ou are about to embark on a journey of discovery about the
nature of life. Almost two centuries ago, a young English natu-
ralist named Charles Darwin set sail on a similar journey on board
H.M.S. Beagle; the photo shows a replica of this ship. Darwin’s
observations during the voyage of the Beagle influenced his devel-
opment of the theory of evolution by natural selection, which has
become the core of the science of biology. Before we begin, how-
ever, let’s take a moment to think about what biology is and why it’s
important.
Domain Archaea. This Domain Bacteria. This

1.1 The Diversity of Life prokaryotic domain includes
this methanogen, which
prokaryotic domain includes
this purple sulfur bacteria,
manufactures methane as a which can use light energy to
Is Overwhelming result of its metabolic activity. drive the synthesis of organic
compounds (false color).
Biology is the study of living things—literally the science of life. Figure 1.1 The two prokaryotic domains. Bacteria and
The living world is teeming with a breathtaking variety of creatures— archaea share the feature of lacking a membrane-bounded
whales, butterflies, mushrooms, plants, bacteria—which despite nucleus. Organisms from both of these domains are single-celled.
their obvious differences share features common to all living organ- (Archaea): ©Power and Syred/Science Source; (Bacteria): ©Alfred Pasieka/SPL/
isms. We will begin with a brief history of how we classify this Science Source
diversity.
Biological Diversity Can Be Organized

by Evolutionary History divided into three domains: Eubacteria, Archaea, and Eukarya.
Even this system has been challenged as there is evidence that
LEARNING OBJECTIVE 1.1.1 Describe the three domains of life. eukaryotes arose from within Archaea, leading some to advocate
for a two-domain system. This remains an area of active research,
For as long as scientists have been thinking about living sys- with many new prokaryotes being identified based on the ability
tems, there have been efforts to classify organisms to provide a to analyze genetic material without actually visualizing or cultur-
framework that organizes the incredible diversity of life. Classi- ing organisms.
fication systems first used observable similarities in organisms The kingdoms you are probably most familiar with contain
to define groups. This led to the Linnaean taxonomic system, plants, animals, and fungi (figure 1.2), and these represent single
first proposed in 1735, which provided a hierarchical system to lines of evolutionary descent. The group that has been problem-
classify all organisms. With the subsequent recognition that all atic is the protists, which were grouped based on the shared simi-
organisms have an evolutionary history, biologists looked to larity of being eukaryotic and unicellular. This was formerly a
integrate evolutionary relationships with classification based on kingdom, but clearly does not represent a single line of evolution-
shared similarities. Classification based on shared evolutionary ary descent. This has led to attempts to find clear groups with
history often, but not always, produces results similar to shared shared evolutionary history. We will discuss this in detail in chap-
characteristics (see chapter 22). Although the Linnaean system ters 22 and 24.
does not always reflect evolutionary relationships, it remains The work of biologists affects your everyday life: what you
useful today because it is the only universal system, and its long eat, what happens to you when you go to the hospital, and how
history means that an extraordinary amount of biodiversity has our society will handle environmental issues such as climate
been classified this way. change. Unifying the diverse systems studied by biologists are the
The highest level of organization in the Linnaean system shared characteristics of all living things that have been shaped
is the kingdom. With the advent of microscopes, we also by the process of evolution by natural selection. Keeping this
defined two basic cell types: those that have genetic material in theme in mind will help you to manage the complexity and
a membrane-bounded nucleus (eukaryotes), and those that lack diversity of biology.
this membrane-bounded nucleus (prokaryotes). This was
accommodated within the Linnaean system by creating a king-
REVIEW OF CONCEPT 1.1
dom for prokaryotes, specifically Bacteria (once called
Kingdom Monera). This was later upended by the discovery The living world is incredibly diverse. Various systems of
that there were actually two kinds of prokaryotes: called Bacte- classification have been proposed, which remains an area of
ria (or Eubacteria) and Archaea (figure 1.1; discussed in detail active research. The oldest branching of the tree of life is into
in chapter 23). Bacteria, Archaea, and Eukarya.
This change led to the addition of a new taxonomic rank
■■ What are some shared features of living systems?
above kingdom called a domain. All living organisms can be
2 Part I The Molecular Basis of Life
mas40908_ch01_001-020.indd 2 10/10/19 2:08 PM

Protists. Once considered a Fungi. This kingdom contains Plantae. This kingdom contains Animalia. Organisms in this
kingdom, this term is still used mostly multicellular photosynthetic multicellular kingdom are nonphotosynthetic
informally to refer to unicellular nonphotosynthetic organisms organisms that are terrestrial, multicellular organisms that
eukaryotes. that digest their food such as the flowering plant digest their food internally,
externally, such as mushrooms. pictured here. such as this ram.
Figure 1.2 The eukaryotic domain. Eukaryotes consist of most familiar organisms and many that are not. The eukaryotes can be
divided into three kingdoms: Fungi, Plantae, and Animalia. Protists are a diverse group of unicellular eukaryotic organisms with no
exclusive common ancestor.
(Protista): Source: Dr. Stan Erlandsen and Dr. Dennis Feely/CDC; (Fungi): ©Russell Illig/Getty Images; (Plantae): ©Iconotec/Glow Images; (Animalia): ©Alan and Sandy
Carey/Getty Images
whereas others have complex organization, but all are able

1.2 Biology Is the Science to grow and reproduce. Many organisms possess only a
single cell, like the paramecium in figure 1.3; your body
of Life contains about 10 trillion to 100 trillion cells (depending
on how big you are).
In its broadest sense, biology is the study of living things. So it 2. Energy utilization. All living things use energy. Moving,
would seem that biologists would have no problem defining life. growing, thinking—everything you do requires energy.
In fact, it is quite difficult to provide a simple definition of life. Where does all this energy come from? It is captured from
sunlight by plants and algae through photosynthesis. To get
Life Defies Simple Definition the energy that powers our lives, we extract it from plants
or from plant-eating animals. That’s what the kingfisher is
LEARNING OBJECTIVE 1.2.1 Describe five fundamental doing in figure 1.4, eating a fish that ate algae.
properties of life. 3. Homeostasis. All living things maintain relatively constant
internal conditions so that their complex processes can be
What does it mean to be alive? What properties define a living better coordinated. Although the environment often varies
organism? These questions are not as simple as they appear, considerably, organisms act to keep their interior
because some of the most obvious properties of living organisms
are also properties of many nonliving things—for example, com-
plexity (a computer is complex), movement (clouds move in the
sky), and response to stimulation (a soap bubble pops if you touch
it). To appreciate why these three properties, so common among
living things, do not help us to define life, imagine a mushroom
standing next to a television: The television seems more complex
than the mushroom, the picture on the television screen is mov-
ing but the mushroom just stands there, and the television
responds to a remote-control device but the mushroom continues
to just stand there—yet it is the mushroom that is alive.
All living things also share five more fundamental proper-
ties, passed down over millions of years from the first organisms
to evolve on Earth: cellular organization; energy utilization; homeo-
Figure 1.3 Cellular organization. This paramecium is a
stasis; growth, development, and reproduction; and heredity.
complex single-celled protist that has just ingested several yeast
1. Cellular organization. All living things are composed of one cells. Like this paramecium, many organisms consist of just a
or more cells. Often too tiny to see, cells carry out the single cell, while others are composed of trillions of cells.
basic activities of living. Some cells have simple interiors, ©Melba Photo Agency/Alamy
Chapter 1 The Science of Biology 3
mas40908_ch01_001-020.indd 3 10/10/19 2:09 PM

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Fig. 519
Incision for resection of thorax. (Bergmann.)
Fig. 520
Trap-door thoracotomy. (Lejars.)
The original Estlander operation has been modified by Schede,

and as now practised is made by a long incision passing obliquely
across the lateral aspect of the chest, from the origin of the
pectoralis major, at the level of the axilla, to the tenth rib in the
posterior axillary line, and then ascending to a point between the
spine and the scapula. The large flap thus outlined is made to
envelop all the tissues outside the ribs. The ribs thus exposed are
resected from the tubercles forward to their insertion into the costal
cartilages. The large area of the chest wall thus exposed is then
removed with the underlying pleura, and all hemorrhage checked.
This flap includes the periosteum, the intercostal muscles, the ribs,
and the pleura, and thoroughly uncovers the entire abscess cavity. It
makes a formidable procedure, but is more often life-saving than the
reverse. Over the opening the skin flap may be later drawn down
and tacked in place at points sufficiently near to each other to
properly hold it in place (Figs. 519 and 520).
This procedure may be modified to suit the indications of any
given case, and simply includes what may be done in extreme
cases. The surgeon who thus for the first time uncovers such a
cavity will be surprised at its interior appearance, and at the shreds
of tissue and debris which hang from its walls. The measure thus
described provides for collapse of the chest wall. Fowler and others
have shown, however, that even now the principal obstacle to
expansion of the lung is not removed, and have suggested what
Fowler has aptly described as decortication of the lung—namely, a
removal of its thickened pleura by a process of dissection and
stripping, which may be made partial or complete, as circumstances
permit. In some respects this adds to the gravity of the case and will
perhaps better be done at a second operation. Should it, however,
be justified by the condition of the patient it is best done in
connection with the resection of the chest wall.
When decortication cannot be practised Fowler has advised that a
series of incisions be made, and that by thus gridironing the
thickened membrane it may be weakened or caused to lose its
inelasticity and thus a mild degree of similar effect secured. Fig. 521
illustrates the end result of such an extensive thoracoplasty.
Fig. 521
End result of an extensive thoracoplasty. (Park.)
Pneumotomy.—This is a term applied to an attack upon the lung

itself, it having been exposed by a thoracotomy. It is
necessary in cases of gangrene, abscess, hydatid cyst, and
occasionally in large bronchiectatic cavities. It is not ordinarily a
difficult procedure when the lung has attached itself to the chest wall
in the course of the disease process. Here the lesion having been
located a part of one or more ribs is removed, as may be needed,
thus exposing the lung surface, the cavity is then opened either with
a knife or by dilatation with the blades of a forceps, or preferably with
the thermocautery blade, by which hemorrhage is better controlled
and possibilities of absorption reduced. If such a cavity can be
located it may be opened with a large trocar and cannula, which
should be introduced with great care, lest it be thrust too far, the
method by incision being therefore preferable. If after opening the
chest the lung be found non-adherent, it depends on the character of
the lesion whether adhesion should be provoked or the cavity itself
attacked. In the former case adhesions may be produced by stitching
the exposed lung surface to the margins of the wound, and waiting
for sufficient exudate to be poured out to ensure that the pleural
cavity has been hermetically sealed. The same result may be
obtained more crudely by packing gauze around the opening.
In case of urgency it would probably be best to attach the lung to
the chest wall with sutures and secure it there. This is a
comparatively safe method in dealing with hydatid cysts, and will
give a fair measure of success in many other instances. The
suppurating or gangrenous cavity being opened its contents should
be removed, dead or sloughing tissue excised, and the cavity then
packed for drainage purposes, the external wound being kept open
until it can be safely allowed to close.
Pneumonectomy, that is, removal of a portion of the lung
substance, may be done with comparative safety upon animals, but
rarely upon human patients. It is occasionally required in connection
with the removal of malignant tumors of the chest wall, to which the
lung has affixed itself. In exceedingly rare instances it may be
justified for localized tumors of the lung itself. It would be equally
valuable for circumscribed, primary tuberculosis of the lung, were it
possible to recognize this in time. This an Italian surgeon once
thought that he had done, in the case of his fiancée, and proceeded
to resect the upper lobe of one of her lungs. His lack of success
quickly led to his own suicide a few days later.
The lung is exceedingly vascular and at the same time bears
sutures well. The suturing, however, should be accurate in order to
prevent secondary hemorrhage and favor the process of repair.
Other operations may be practised upon the chest wall for relief of
such conditions as acute osteomyelitis of the ribs or sternum, caries
of the ribs, necrosis, and the like. It should be scarcely necessary to
give explicit directions, save that the pleural cavity should never be
opened unless the pleura itself be involved in the disease. Every
case demanding such operative relief should be measured by its
own needs, and the operative procedure adapted to them. Necrosed
portions of bone may be completely removed. The suppurative and
carious conditions necessitate rather a sufficiently wide exposure
from without and then a judicious use of the bone curette. One need
never hesitate to remove so much bone as is diseased, this being
true even of the sternum.
THE THYMUS.
The possibility of suffocative and other disturbances proceeding
from enlargement of the thymus has been discussed, as well as the
use of long trachea tubes in cases of this character which call for
tracheotomy, as they usually do if they permit of any surgical
intervention. The thymus is seldom the site of primary malignant
disease. Certain acute lesions are due to a peculiar form of
hypertrophy in the young, which takes place instead of that
spontaneous disappearance which should have occurred during the
earliest months of infancy. Its connection with the status lymphaticus,
with thymic asthma, and laryngismus stridulus has already been
mentioned. While it can hardly be considered absolutely exempt
from ordinary infections and the like it nevertheless is rarely involved.
The thymus has been removed by operation, usually with success.
Should it become necessary to resort to such a measure it should be
preceded by the removal of the sternum, for only in this way can
sufficient exposure be obtained, and sufficient opportunity for
checking such hemorrhage as might result from its enucleation.
THE AXILLA.
The axilla as a surgical region belongs as much to the thorax as to
any part of the body, although none of its diseases are peculiar to
this area.
It is frequently the site of furuncles of local origin, which
occasionally assume carbuncular type, and which are expressions of
local infection along the hair follicles or mammary ducts. It is full of
lymph nodes, through which are filtered the lymph streams coming
from the upper extremities. In this way there are entangled therein
septic germs, which frequently give rise to small or large phlegmons
proportionate in size to the magnitude of the lesion beyond them. It
takes but a trifling infection of the finger, for instance, to produce
such involvement of axillary lymph nodes as to make them palpable
under the finger. Such lymph nodes once genuinely inflamed
frequently coalesce, and the resulting abscess cavity may be large,
especially if neglected. The sooner these phlegmons are incised and
cleaned out the better for the patient. In order to do thorough work
an anesthetic is usually required.
In the axilla also are frequently seen tuberculous manifestations,
the result of propagated infection from some part of the arm or hand.
These may be involved in a mixed infection and quickly break down,
or may assume the type of the chronically enlarged nodes, which
undergo caseation and more or less encapsulation, with such
infiltration of the surrounding tissues that when extirpated
considerable difficulty is met in the dissection.
In syphilis, also, the lymph nodes become involved, frequently
enlarging to a degree making them palpable, and sometimes
participating in a mixed infection in such a way as to break down into
abscesses.
Again, in the axilla are occasionally seen conspicuous evidences
of Hodgkin’s disease. Any disease of constitutional character which
precipitates trouble in one axilla will cause nearly duplicate
alterations in the other, whereas disease of local origin is usually
confined to one side.
Any phlegmonous cavity or tuberculous lesion which has been
incised through the axilla should be carefully cleaned out and then
drained, lest the external incision close before the deeper parts are
ready for it. Incisions made in the axilla should be parallel with the
great vessels and nerve trunks, by which they are better exposed
and avoided. A wound made in the axillary vein may be sutured or
the vein be doubly ligated. The former is much the better course,
very fine silk sutures being employed. In some lesions where it has
not been possible to discover the bleeding point the writer has not
hesitated to secure it with the ends of pressure forceps and to leave
these forceps included in the dressings for forty-eight hours. He has
never seen harm result from this procedure.
Finally the axilla is Fig. 522
almost always
involved in cases of
malignant disease of
the breast, of the arm
itself, and sometimes
of the regions
adjoining. Primary
malignant disease in
this region is rare,
while secondary
cancer is not unusual.
According to the
modern plan of
treatment of cancer
there is reason for
scrupulous extirpation
of every particle of
infected tissue and all
involved lymphatics,
and in dealing with
such cases the
surgeon need not
hesitate to divide or
extirpate the pectoral
muscles, in order to
permit of thorough
work. The disease
being present nothing
can be so serious for
the patient as to allow
any particle of it to
remain.
Congenital diaphragmatic hernia, with other
congenital defects. Wood Museum. (Dennis.)
THE
DIAPHRAGM.
The diaphragm may show certain congenital defects, consisting
mainly of fissures or openings which permit displacement of viscera,
usually from the abdomen below into the thorax above. This is often
fatal, constituting a form of diaphragmatic hernia, which is
particularly liable to strangulation. Fig. 522 indicates a case of this
kind, showing the hopelessness of the condition.
Anatomically it is worth while to recall that the diaphragm may rise
to a level with the third cartilage during forced expiration, and
descend to the level of the fifth intercostal space on the right side,
and a little lower on the left, during forced inspiration. When forced
upward by pressure from below it may rise even higher than stated
above. These facts are of surgical interest in considering the
possibility of injury or perforation of the diaphragm in connection with
gunshot and other perforating injuries to the thorax or abdomen.
Diaphragmatic paralysis is the necessary result of injury to the
phrenic nerve. It may occur as the result of injury to the thoracic
viscera, especially those of the posterior mediastinum, or injuries to
the cervical or upper dorsal vertebræ, usually fractures or
dislocations, followed by ascending degeneration and involvement of
the phrenic nerve roots. Double phrenic paralysis is in these cases
obviously fatal. Paralysis of a single side will cause at least serious
embarrassment of respiration. An hysterical form of diaphragmatic
paralysis has also been described.
Primary tumors are exceedingly rare in this muscular partition.
Advancing growths, however, attach themselves to it or perforate it,
as may also aneurysms.
Aside from the ordinary injuries which the diaphragm may suffer
from without, and already mentioned, there are peculiar forms of
rupture, the result of force applied from below, usually at right angles
to the surface of the body, this being permitted on account of the
dome-like shape of the muscle. When thus ruptured abdominal
viscera may be forced into the chest and even out through openings
between the ribs. A gunshot wound of the diaphragm will be serious
mainly in proportion to other injuries involving the viscera above or
below it. These injuries produce no typical symptoms, but are nearly
always accompanied by severe pain radiating toward the shoulders,
with dyspnea and a substitution of abdominal for diaphragmatic
respiration. When the viscera have been forced upward they will
displace the heart, and this may produce cardiac symptoms. It is
said that the so-called “sardonic grin” is still observed on the faces of
corpses who came to sudden death from some injury to the
diaphragm.
Thus diaphragmatic wounds are not of themselves of serious
import. When inferentially present they may, therefore, be
disregarded so long as no serious symptoms are produced. On the
other hand, exploratory celiotomy should be performed at any time,
should conditions seem to justify it.
SUBDIAPHRAGMATIC OR SUBPHRENIC ABSCESS.

While this is a condition pertaining, strictly speaking, to the
abdominal cavity, it nevertheless arises so frequently from
intrathoracic causes as to justify its consideration here, as well as
because of its close relations to the diaphragm. It was Volkmann
who, in 1879, first showed how these abscesses could be
successfully and surgically treated. The term is applied to collections
of pus beneath the diaphragm, usually between it and the liver,
which, however, may extend to and later involve surrounding viscera.
The causes may be divided into those met with above the
diaphragm and those below. The former may include empyema, pus
having escaped beyond the normal pleural limits, advancing
tuberculous disease from any of the structures above the diaphragm,
echinococcus in the lung, or suppurative mediastinitis. From below
the diaphragm the infectious process may travel from the direction of
a gastric or a duodenal ulcer, hydatid disease in the liver, phlegmon
around the liver or kidney. The contained pus may, on culture, show
the presence of colon bacilli or pneumococci, as well as the ordinary
pyogenic cocci and tubercle bacilli. If connected with hydatid disease
hooklets may be seen in pus which is not too old.
Subphrenic abscess may result in large collections of pus, which
may travel a considerable distance, separating the peritoneum from
the diaphragm and from the lateral abdominal walls, appearing even
low down in the pelvis. The same is true of escaping pus from a case
of empyema. The primary trouble gives rise to a localized peritonitis
or perihepatitis, by which are produced certain barriers that serve to
retain pus within bounds, and to keep it from spreading save as
above mentioned. Should it be due to extension of abscess or
disease within the liver it may be confined by adhesions about it. Fig.
523 illustrates the relations which such a collection may sustain to
the liver and the diaphragm, as well as how the opening by which it
may be best evacuated should be made through the thoracic walls.
Even with this condition produced by disease below the diaphragm it
is not infrequent to find some collection of fluid or evidence of
exudate above it.
A study of this condition will nearly always lead one back to a
history of some illness which may furnish the explanation for the
commencement of the trouble. Thus, there may be obtained a
history of pulmonary tuberculosis, of empyema, of gastric ulcer, of
gallstone trouble, or of abscess in the liver or in or about the kidney.
When the result of perforation from above, the chest wall may furnish
signs which will be sufficiently indicative.
The symptoms will include swelling, pain, tenderness, with fixation
of the liver, and apparent enlargement of its boundaries, because it
is pushed away from the diaphragm. The abdominal wall will
frequently be edematous. The ordinary signs of the presence of pus
are rarely absent, including the evidences furnished by a differential
blood count. Diagnosis is proved by the use of the exploring needle.
The disease is nearly always situated upon the right side. The more
distended the abscess cavity the less respiratory murmur will be
heard over the lower part of the chest, while the line of the hepatic
dulness may be considerably above the normal. Sometimes a
succussion sound may be obtained.
Should pus be withdrawn from the lower part of the chest by the
exploring needle there might still be doubt as to its actual location,
whether above or below the diaphragm. The absence of cough and
of indications of pleural involvement would prove much in favor of
the latter.
Subphrenic abscesses tend in time to evacuate themselves. Thus
they sometimes perforate the diaphragm and escape into the pleural
cavity, or through a lung which has attached itself at its base, and
thus afforded an outlet for pus through the bronchi and the mouth.
On the other hand, pus may burrow downward and appear in the
flank or beneath the skin near the liver and in front of it. The nearer it
comes to the surface the more easily it is recognized.
Fig. 523
Transthoracic opening for subphrenic abscess. (Beck.)
Treatment.—The treatment of subdiaphragmatic abscess, like that

of all other abscesses, consists in evacuation of the
contained pus, with provision for drainage. In some instances this
may be done with an ordinary trocar and cannula, but serious cases
are best treated by incision, with resection, if necessary, of a portion
of a rib. When the chest wall is entered the best place is between the
ninth and tenth ribs in the axillary line. Nevertheless pus which is
presenting at any other point may be best reached by taking
advantage of the indication thus afforded. An opening having been
made the question of counteropening may be raised. This should be
decided in each instance upon its merits. While an opening made in
front does not drain so well as one placed posteriorly it may be made
to drain by keeping the patient upon the side or face for a portion of
the ensuing few days. When it seems desirable to go through the
chest wall it should be incised carefully, and if the pleura has been
opened before reaching the abscess, the pleural surfaces may be
either stitched together or packed; after waiting a day or two for
protective adhesions to form the abscess may then be opened. The
less extensive operations may be performed with local anesthesia.
Rib resection and extensive incision will usually require general
anesthesia.
C H A P T E R X L I V.
THE BREAST.
ANOMALIES OF THE BREAST.

Amastia, or complete absence of one or both breasts, is a rare
defect. Polymastia, or the occurrence of supernumerary breasts, is
more frequent.[51] These may be found on any portion of the thorax
or abdomen, and may constitute masses of trifling size or may bear
considerable resemblance to the normal breast. A supernumerary
breast has even been found upon the thigh. The condition is to be
regarded as atavistic, and a return to the polymastia of animals,
which produce a litter at one birth. Similarly absence of the nipple,
amazia, is occasionally seen, or more frequently polymazia, the
occurrence of more than one nipple, either upon the normal breast or
in some abnormal position. Some of these lesions are so small as to
escape observation, or to be considered moles unless carefully
noted and recognized when found.
[51] History records interesting examples of the importance attached to
these conditions. Thus the beautiful Anne Boleyn fell under the displeasure
of King Henry because of a supernumerary breast, and it is said that the
mother of the Roman Emperor Alexander Severus was given the name of
Julia Mammæ because of a similar abnormality.
Ordinarily supernumerary breasts are met near the middle line and
below the normal mammary gland. A more common condition is one
of defect of the nipple, which fails to assume its normal prominence
and remains ill-developed, so as not to be seized by the infant in the
act of attempting to nurse. Nevertheless with the physiological
activity which occurs in the breast at the time of pregnancy these ill-
developed nipples usually expand sufficiently to fulfil their function,
even though imperfectly.
Hemorrhages from the breast sometimes take place idiopathically,
at others as vicarious efforts at menstruation. There is a peculiar
sympathy between the pelvic organs of women and the mammary
glands, and the latter evince this in more than one way, becoming
sometimes extremely tender or swollen at the menstrual period, or at
other times peculiarly sensitive or even neuralgic, while at times
congestion will proceed to the point of hemorrhage. These conditions
do not require particular attention, but are not to be confused with a
bloody discharge that may occur later in life, in connection with
certain forms of malignant disease occurring in the interior of the
breast.
There exist the widest differences in development of the breasts in
different individuals. The term “breast” is used intentionally, since the
difference is not so much in the actual glandular development as in
the surrounding connective tissue and fat. Thus a plump breast may
contain very little more secreting structure than one apparently ill-
developed. Nowhere outside the uterus save in the breast do such
compensatory changes take place under the stimulus of pregnancy.
In fact, a mammary gland in preparation for lactation is a
physiological adenoma. At conclusion of lactation there is absorption
and atrophy from disuse, usually not to the original degree, although
in some instances the fatty tissue disappears irregularly and leaves
the breasts in quite different shape from their originals. In this way
the breasts may become exceedingly pendulous, so much so as to
lead to pain and soreness from traction, and to call for their support.
Idiopathic hypertrophy of one or both breasts is a rare deformity,
occurring usually in the young, sometimes in girls, involving them to
an indefinite degree, but in some producing enormous overgrowth,
with corresponding deformity. For such hypertrophy no known cause
has been assigned. Fig. 524 illustrates an instance of this character
in a young girl, occurring under the observation of my colleague, Dr.
Bebee.
INJURIES TO THE BREAST.

These consist largely of contusions to which, from their positions,
the breasts are peculiarly exposed, and these may be followed by
hemorrhage, by extensive ecchymosis, or by any of the
consequences of infection. They may also be followed by more or
less permanent induration. The fact that in the course of time certain
Fig. 524 contusions of the
breast are followed by
development of
cancer is
incontestable,
although the relation
which may exist
between the accident
and the neoplasm
has not yet been
made clear. The
breasts are also
subject to the same
possibilities of injury
as other parts of the
thorax, which has
been considered in
the previous chapter.
The nipples are more
often injured by
efforts of the nursing
infant, or by the
friction of ill-fitting
stays or rough
clothing, than in any
other way. These
injuries, at first of a
minor character, are
not infrequently
followed by serious
results, erysipelas,
septic infection, or
tuberculosis being
conveyed through
Idiopathic hypertrophy of breasts in a girl of sixteen.
trifling abrasions thus
(Bebee.)
inflicted.
The nipple of a nursing woman once excoriated, or its surface
broken, is kept constantly liable to maceration and surface infection.
In this way a trifling lesion may result in a linear ulcer known as a
fissure (“cracked nipple”), or in a more extensive involvement. These
fissured nipples are very erethistic, and great pain is caused by each
attempt at nursing. On this, account the mother postpones the act as
long as possible, and until her breast has become overdistended, the
result being injury to the breast itself, with a greater possibility of
infection and of subsequent abscess formation.
The slightest excoriation of the nipple, under any circumstances,
should lead to the adoption of every precaution for its cleansing and
protection. Both before and after nursing it should be carefully
washed, while, after removal of the child from the breast, it should be
carefully dried and dusted with dry boric acid or a similar antiseptic.
Any abrasion which fails to heal should be treated with silver nitrate.
More pronounced abrasions and ulcers should be cocainized, then
cauterized, and afterward treated as above. Finally in extreme cases
it may be necessary to discontinue nursing and allow the breast to
dry. If this policy be adopted it should be applied to both breasts, for
such is the sympathy between them that the use of one gland seems
to stimulate the other. The local use of such preparations as
belladonna ointment, etc., is to be avoided. Pressure, rest, and the
care above described afford more relief.
Paget’s disease of the nipple implies an eczematous condition,
first described by Paget as a precursor of many cancers. It is a more
or less chronic affection, involves the nipple and the areola, is quite
intractable to treatment, gives more or less discomfort, and is to be
dreaded when noted. It seems to sustain about the same relation to
later cancerous involvement as does leukoplakia in the mouth and
on the tongue.
There is no reason why any person may not have an attack of
eczema about the nipple, but cases in which the condition is
persistent and obstinate, and especially in which the underlying
tissues gradually become infiltrated or indurated, should be viewed
with suspicion, and should be treated by eradication of the area
involved, even though this may require extirpation of the nipple or of
the entire breast. When the condition is developed no ordinary
treatment will suffice, although a fair trial might be given to the
cathode rays.
MASTITIS.
A true inflammation of the mammary gland may occur at one of
three periods: (1) At birth, when the tiny breasts of the newborn
infant secrete a milk-like fluid, become more or less congested and
tender, and when they are unintelligently treated by massage or
interference of any kind; (2) at puberty, when a perfectly natural
turgescence and congestion occur, which, however, rarely proceed
to suppuration unless infected or unless violence or some indiscreet
treatment have been received; (3) during pregnancy and lactation,
this being the time when mastitis is most common.
Considering that the nipple affords a number of open paths, from
an area which it is difficult to keep clean, extending into the depths of
inflammable tissue, it is strange that infection through the milk ducts
does not occur in most cases. Such a path of infection affords the
explanation for at least a large proportion of mammary abscesses.
Again the presence of excoriations, abrasions of any kind, and
especially of deep fissures which are not easily cleansed, will
account for infection through the lymphatics. In these two ways
nearly all cases of mastitis and of mammary abscess are to be
explained, and both these accidents are likely to occur during
pregnancy and lactation.
The consequence of such infection is mastitis, which begins with
painful induration and local indications of inflammation, but which
may under suitable treatment undergo resolution. This failing, the
infectious process proceeds to suppuration, and the consequence is
a superficial, deep, or retromammary abscess, all but the last named
often in multiple form. The lobular construction of the breast permits
the independent occurrence of distinctive suppuration, occurring
synchronously at several different points, and hence it may be that a
breast is riddled with abscesses, which form successively or almost
simultaneously.
There is a superficial form, which occurs usually near the nipple,
and in which the deeper structure of the breast is scarcely involved.
This comes usually through infection of some surface lesion. Simple
incision is usually sufficient, and the local lesion is thus quickly
terminated. Deep or intramammary abscess, single or multiple, is
always painful, sometimes distressing and occasionally an extremely
serious condition. Occurring in a breast already well developed and
fatty, abscesses may form at such depth as to be recognized with
difficulty. The surgeon infers their existence rather than discovers it.
This is unfortunate, for the longer the delay the greater the local
disturbance, with a tendency to burrowing, and the worse are the
consequences for the patient. It is, therefore, far safer to early note
the minor signs of deep suppuration and to freely incise, than it is to
wait for pus to come toward the surface and give its ordinary surface
indications. The amount of induration, sometimes dense and brawny,
which such conditions will produce within the breast, the size which
the latter may assume, and the consequent suffering to the patient
from neglected conditions of this kind, need to be seen to be fully
appreciated.
Retromammary abscess may be the result of conditions not
primary to the breast itself. Thus the writer has seen spontaneous
perforation of the thoracic wall in a case of empyema, with escape of
pus into the loose cellular tissue behind the breast, and the
consequent protrusion forward of the latter until it presented as an
enormous tumor. Treatment in such cases would mean not alone
evacuation of the retromammary collection, but emptying the pleural
cavity of its accumulated fluid.
An infected breast will produce not only the ordinary local
indications, but will be characterized by extreme tenderness, with
enlargement of the lymph nodes in the axilla and later abscess
formation in this location. In proportion to the amount of pus thus
imprisoned, and the virulence of the infecting organisms,
constitutional symptoms may be mild or extreme.
Nowhere is there greater need for release of an imprisoned
amount of pus than under these circumstances, although the
incisions necessary for the purpose may be sometimes multiple and
deep. Every incision made for evacuation of a mammary abscess
should be placed radially—i. e., in a line radiating from the nipple—in
order that lobules may be incised along their course, and that neither

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