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Biology 11e

Eldra P. Solomon
former affiliations
Hillsborough Community College, Tampa
University of South Florida

Charles E. Martin
professor emeritus, Rutgers University

Diana W. Martin
professor emeritus, Rutgers University

Linda R. Berg
former affiliations
University of Maryland, College Park
St. Petersburg College

Australia • Brazil • Mexico • Singapore • United Kingdom • United States

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Biology, Eleventh Edition © 2019, 2015, 2011 Cengage Learning, Inc.
Eldra P. Solomon, Charles E. Martin, WCN: 02-300
Diana W. Martin, Linda R. Berg Unless otherwise noted, all items are © Cengage.
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To our families, friends, and colleagues who gave
freely of their love, support, knowledge, and time as
we prepared this eleventh edition of Biology, and in
appreciation of all who teach and learn.
Especially to
My grandchildren and their generation
Margaret, Damian, and Ava
Alan, Jennifer, and Pat

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About the Authors

Eldra P. Solomon has writ- Charles E. Martin is professor Diana W. Martin is professor Linda R. Berg is an award-
ten several leading college emeritus of cell biology and emeritus and former direc- winning teacher and textbook
textbooks in biology and in neuroscience at Rutgers Uni- tor of general biology in the author. She received a B.S. in
human anatomy and physi- versity. He received his Ph.D. Division of Life Sciences at science education, an M.S. in
ology. Her books have been in genetics from Florida State Rutgers University. Dr. Martin botany, and a Ph.D. in plant
translated into more than ten University and engaged in received an M.S. from Florida physiology from the Univer-
languages. She earned an M.S. postdoctoral research in ge- State University, where she sity of Maryland. Her research
from the University of Florida netics and membrane biology studied the chromosomes of focused on the evolutionary
and an M.A. and Ph.D. from at the University of Texas at related plant species to under- implications of steroid bio-
the University of South Flor- Austin. He has taught general stand their evolutionary rela- synthetic pathways in various
ida. Dr. Solomon taught biol- biology as well as undergradu- tionships. She earned a Ph.D. organisms.
ogy and nursing students for ate and graduate level courses from the University of Texas Dr. Berg taught at the Uni-
more than 20 years. in genetics and molecular cell at Austin, where she studied versity of Maryland at College
In addition to being a bi- biology throughout his career the genetics of the fruit fly, Park for 17 years and at St. Pe-
ologist and science author, Dr. at Rutgers. An award-winning Drosophila melanogaster, and tersburg College in Florida for
Solomon is a biopsychologist teacher for more than 30 years, then conducted postdoc- 8 years. During her career, she
with a special interest in the in 2011 Dr. Martin was named toral research at Princeton taught introductory courses in
neurophysiology of traumatic Professor of the Year by the University. biology, botany, and environ-
experience. Her research has Molecular Biosciences Gradu- Dr. Martin taught general mental science to thousands
focused on the neurological, ate Student Association. biology and other courses at of students. At the University
endocrine, and psychological His research on gene regu- Rutgers for more than 30 years of Maryland, she received nu-
effects of trauma, including lation of membrane protein and has been involved in writ- merous teaching and service
complex post-traumatic stress enzyme systems in yeast and ing textbooks since 1988. She awards. Dr. Berg is also the
disorder and development of other fungi illustrates the in- is immensely grateful that her recipient of many national and
maladaptive coping strategies. terdisciplinary nature of the decision to study biology in regional awards, including
Dr. Solomon has pre- life sciences. He is most proud college has led to a career that the National Science Teach-
sented her research at numer- of the many generations of allows her many ways to share ers Association Award for In-
ous national and international undergraduate, graduate, and her excitement about all as- novations in College Science
conferences, and her work postdoctoral students who pects of biology. Teaching, the Nation’s Capital
has been published in lead- contributed to this research Area Disabled Student Ser-
ing professional journals. She and have gone on to produc- vices Award, and the Wash-
has been profiled more than tive careers. He continues to be ington Academy of Sciences
30 times in leading publica- committed to teaching and is Award in University Science
tions, including Who’s Who in grateful for the opportunities Teaching.
America, Who’s Who in Science to pursue a teaching and re- During her career as a
and Engineering, Who’s Who search career in what contin- professional science writer,
in Medicine and Healthcare, ues to be the most exciting era Dr. Berg has authored or coau-
Who’s Who in American Edu- of the biological sciences. thored several leading college
cation, Who’s Who of American science textbooks. Her writing
Women, and Who’s Who in the reflects her teaching style and
World. love of science.

iv

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Brief Contents

Preface xxiii 28 Seed Plants 584


To the Student xxx 29 The Fungi 603
part one: The Organization of Life 30 An Introduction to Animal Diversity 628
31 Sponges, Cnidarians, Ctenophores,
1 A View of Life 1 and Protostomes 641
2 Atoms and Molecules: 32 The Deuterostomes 676
The Chemical Basis of Life 26
part six: Structure and Life Processes in Plants
3 The Chemistry of Life: Organic Compounds 46
4 Organization of the Cell 73 33 Plant Structure, Growth, and Development 710

5 Biological Membranes 106 34 Leaf Structure and Function 729


6 Cell Communication 131 35 Stem Structure and Transport 745
36 Roots and Mineral Nutrition 762
part two: Energy Transfer through Living Systems
37 Reproduction in Flowering Plants 782
7 Energy and Metabolism 150
38 Plant Developmental Responses to ­External
8 How Cells Make ATP: and ­Internal Signals 803
Energy-Releasing Pathways 167
9 Photosynthesis: Capturing Light Energy 187 part seven: Structure and Life Processes in Animals

part three: The Continuity of Life: Genetics 39 Animal Structure and Function: An Introduction 821
40 Protection, Support, and Movement 842
10 Chromosomes, Mitosis, and Meiosis 206
41 Neural Signaling 860
11 The Basic Principles of Heredity 228
42 Neural Regulation 882
12 DNA: The Carrier of Genetic Information 253
43 Sensory Systems 911
13 Gene Expression 272
44 Internal Transport 936
14 Gene Regulation 297
45 The Immune System: Internal Defense 962
15 DNA Technology and Genomics 315
46 Gas Exchange 991
16 Human Genetics and the Human
Genome 340 47 Processing Food and Nutrition 1010
17 Developmental Genetics 362 48 Osmoregulation and Disposal of ­Metabolic
Wastes 1032
part four: The Continuity of Life: Evolution
49 Endocrine Regulation 1050
18 Introduction to Darwinian Evolution 385 50 Reproduction 1074
19 Evolutionary Change in Populations 406 51 Animal Development 1104
20 Speciation and Macroevolution 421 52 Animal Behavior 1124
21 The Origin and Evolutionary
History of Life 442 part eight: The Interactions of Life: Ecology
22 The Evolution of Primates 461 53 Introduction to Ecology: Population Ecology 1151
part five: The Diversity of Life 54 Community Ecology 1171
55 Ecosystems and the Biosphere 1194
23 Understanding Diversity: Systematics 478
56 Ecology and the Geography of Life 1216
24 Viruses and Subviral Agents 499
57 Biological Diversity and Conservation Biology 1241
25 Bacteria and Archaea 517
26 Protists 539 Glossary G-1
27 Seedless Plants 563 Index I-1

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Contents

part one: THE ORGANIZATION OF LIFE

1 A View of Life 1 2.2 Chemical Reactions 31


1.1 Major Themes of Biology 2 Atoms form compounds and molecules 31
1.2 Characteristics of Life 2 Simplest, molecular, and structural chemical formulas give
Organisms are composed of cells 3 different information 31
Organisms grow and develop 3 One mole of any substance contains the same number
Organisms regulate their metabolic processes 3 of units 31
Organisms respond to stimuli 4 Chemical equations describe chemical reactions 32
Organisms reproduce 5 2.3 Chemical Bonds 32
Populations evolve and become adapted to the environment 5 In covalent bonds electrons are shared 32
1.3 Levels of Biological Organization 6 The function of a molecule is related to its shape 34
Organisms have several levels of organization 6 Covalent bonds can be nonpolar or polar 34
Several levels of ecological organization can be identified 6 Ionic bonds form between cations and anions 34
1.4 Information Transfer 6 Hydrogen bonds are weak attractions 36
DNA transmits information from one generation to the next 8 van der Waals interactions are weak forces 37
Information is transmitted by chemical and electrical signals 8 2.4 Redox Reactions 37
Organisms also communicate information to one another 8 2.5 Water 38
1.5 The Energy of Life 9 Hydrogen bonds form between water molecules 38
1.6 Evolution: The Basic Unifying Concept of Biology 10 Water molecules interact with hydrophilic substances by
Biologists use a binomial system for naming organisms 11 hydrogen bonding 38
Taxonomic classification is hierarchical 11 Water helps maintain a stable temperature 39
Systematists classify organisms in three domains 11 2.6 Acids, Bases, and Salts 41
Species adapt in response to changes in their environment 14 pH is a convenient measure of acidity 41
Natural selection is an important mechanism by which Buffers minimize pH change 42
­evolution proceeds 14 An acid and a base react to form a salt 43
Populations evolve as a result of selective pressures from
changes in their environment 15
1.7 The Process of Science 15 3 The Chemistry of Life:
Science requires systematic thought processes 16 Organic Compounds 46
Scientists make careful observations and ask critical 3.1 Carbon Atoms and Organic Molecules 47
questions 16 Isomers have the same molecular formula but different
Chance often plays a role in scientific discovery 17 structures 48
A hypothesis is a testable statement 17 Functional groups change the properties of organic
Researchers must avoid bias 18 molecules 49
Scientists interpret the results of experiments and draw Many biological molecules are polymers 50
conclusions 18 3.2 Carbohydrates 51
A scientific theory is supported by tested hypotheses 20 Monosaccharides are simple sugars 51
Many hypotheses cannot be tested by direct experiment 21 Disaccharides consist of two monosaccharide units 52
Paradigm shifts accommodate new discoveries 21 Polysaccharides can store energy or provide structure 53
Systems biology integrates different levels of information 21 Some modified and complex carbohydrates have special
Science has ethical dimensions 21 roles 55
Science, technology, and society interact 22 3.3 Lipids 56
Triacylglycerol is formed from glycerol and three fatty
acids 56
2 Atoms and Molecules: Saturated and unsaturated fatty acids differ in physical
The Chemical Basis of Life 26 properties 57
2.1 Elements and Atoms 27 Phospholipids are components of cell membranes 57
An atom is uniquely identified by its number of protons 28 Carotenoids and many other pigments are derived from
Protons plus neutrons determine atomic mass 29 ­isoprene units 57
Isotopes of an element differ in number of neutrons 29 Steroids contain four rings of carbon atoms 58
Electrons move in orbitals corresponding to energy levels 30 Some chemical mediators are lipids 59

vi

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3.4 Proteins 59 Microfilaments consist of intertwined strings of actin 100
Amino acids are the subunits of proteins 60 Intermediate filaments help stabilize cell shape 102
Peptide bonds join amino acids 61 4.7 Cell Coverings 103
Proteins have four levels of organization 61
The amino acid sequence of a protein determines its
conformation 65 5 Biological Membranes 106
3.5 Nucleic Acids 68 5.1 The Structure of Biological Membranes 107
Some nucleotides are important in energy transfers and other Phospholipids form bilayers in water 107
cell functions 68 The fluid mosaic model explains membrane structure 108
3.6 Identifying Biological Molecules 69 Biological membranes are two-dimensional fluids 109
Biological membranes fuse and form closed vesicles 110
Membrane proteins include integral and peripheral
4 Organization of the Cell 73 proteins 111
4.1 The Cell: Basic Unit of Life 74 Proteins are oriented asymmetrically across the bilayer 111
The cell theory is a unifying concept in biology 74 5.2 Overview of Membrane Protein Functions 113
The organization and basic functions of all cells are 5.3 Cell Membrane Structure and Permeability 114
similar 74 Biological membranes present a barrier to polar
Cell size is limited 74 molecules 114
Cell size and shape are adapted to function 76 Transport proteins transfer molecules across membranes 115
4.2 Methods for Studying Cells 76 5.4 Passive Transport 115
Light microscopes are used to study stained or living cells 76 Diffusion occurs down a concentration gradient 115
Electron microscopes provide a high-resolution image that can Osmosis is diffusion of water across a selectively permeable
be greatly magnified 78 membrane 116
Biologists use biochemical and genetic methods to connect Facilitated diffusion occurs down a concentration gradient 118
cell structures with their functions 79 5.5 Active Transport 120
4.3 Prokaryotic and Eukaryotic Cells 82 Active transport systems “pump” substances against their
Organelles of prokaryotic cells are not surrounded by concentration gradients 120
membranes 82 Carrier proteins can transport one or two solutes 122
Membranes divide the eukaryotic cell into compartments 83 Cotransport systems indirectly provide energy for active
The unique properties of biological membranes allow transport 122
­eukaryotic cells to carry on many diverse functions 83 5.6 Exocytosis and Endocytosis 123
4.4 The Cell Nucleus 84 In exocytosis, vesicles export large molecules 123
Ribosomes manufacture proteins in the cytoplasm 87 In endocytosis, the cell imports materials 123
4.5 Membranous Organelles in the Cytoplasm 88 5.7 Cell Junctions 125
The endoplasmic reticulum is a multifunctional network of Anchoring junctions connect cells of an epithelial sheet 125
membranes 88 Tight junctions seal off intercellular spaces between some
The ER is the primary site of membrane assembly for animal cells 127
­components of the endomembrane system 91 Gap junctions allow the transfer of small molecules and ions 128
The Golgi complex processes, sorts, and routes proteins Plasmodesmata allow certain molecules and ions to move
from the ER to different parts of the endomembrane between plant cells 128
system 91
Lysosomes are compartments for digestion 93
Vacuoles are large, fluid-filled sacs with a variety of 6 Cell Communication 131
functions 94 6.1 Cell Communication: an Overview 132
Peroxisomes metabolize small organic compounds 94 6.2 Sending Signals 133
Mitochondria and chloroplasts are energy-converting 6.3 Reception 134
organelles 95 Cells regulate reception 135
Mitochondria make ATP through aerobic respiration 95 Three types of receptors occur on the cell surface 135
Chloroplasts convert light energy to chemical energy through Some receptors are located inside the cell 137
photosynthesis 97 6.4 Signal Transduction 138
4.6 The Cytoskeleton 98 Signaling molecules can act as molecular switches 138
Microtubules are hollow cylinders 98 Ion channel–linked receptors open or close channels 139
Centrosomes and centrioles function in cell division 99 G protein–linked receptors initiate signal transduction 139
Cilia and flagella are composed of microtubules 99 Second messengers are intracellular signaling agents 139

Contents / vii

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Many activated intracellular receptors are transcription Ras pathways involve tyrosine kinase receptors and
factors 142 G proteins 144
Scaffold proteins increase efficiency 143 The response to a signal is amplified 144
Signals can be transmitted in more than one direction 143 Signals must be terminated 145
6.5 Responses to Signals 143 6.6 Evolution of Cell Communication 146

part two: ENERGY Transfer THROUGH Living Systems

7 Energy and Metabolism 150 Pyruvate is converted to acetyl CoA 171


7.1 Biological Work 151 The citric acid cycle oxidizes acetyl groups derived from
Organisms carry out conversions between potential energy acetyl CoA 171
and kinetic energy 151 The electron transport chain is coupled to ATP
7.2 The Laws of Thermodynamics 151 synthesis 176
The total energy in the universe does not change 151 Aerobic respiration of one glucose yields a maximum of 36 to
The entropy of the universe is increasing 152 38 ATPs 180
7.3 Energy and Metabolism 152 Cells regulate aerobic respiration 181
Enthalpy is the total potential energy of a system 153 8.3 Energy Yield of Nutrients other than
Free energy is available to do cell work 153 Glucose 182
Chemical reactions involve changes in free energy 153 8.4 Anaerobic Respiration and Fermentation 182
Free energy decreases during an exergonic reaction 153 Alcohol fermentation and lactate fermentation are
Free energy increases during an endergonic reaction 154 inefficient 183
Diffusion is an exergonic process 154
Free-energy changes depend on the concentrations of 9 Photosynthesis: Capturing Light
­reactants and products 154 Energy 187
Cells drive endergonic reactions by coupling them to 9.1 Light and Photosynthesis 188
­exergonic reactions 154 9.2 Chloroplasts 189
7.4 Atp, the Energy Currency of the Cell 155 Chlorophyll is found in the thylakoid membrane 190
ATP donates energy through the transfer of a phosphate Chlorophyll is the main photosynthetic pigment 191
group 155 9.3 Overview of Photosynthesis 192
ATP links exergonic and endergonic reactions 156 ATP and NADPH are the products of the light-dependent
The cell maintains a very high ratio of ATP to ADP 156 reactions: An overview 192
7.5 Energy Transfer in Redox Reactions 157 Carbohydrates are produced during the carbon fixation
Most electron carriers transfer hydrogen atoms 157 ­reactions: An overview 193
7.6 Enzymes 158 9.4 The Light-Dependent Reactions 193
All reactions have a required energy of activation 158 Photosystems I and II each consist of a reaction center and
An enzyme lowers a reaction’s activation energy 159 multiple antenna complexes 194
An enzyme works by forming an enzyme–substrate Noncyclic electron transport produces ATP and
complex 159 NADPH 194
Enzymes are specific 160 Cyclic electron transport produces ATP but no
Many enzymes require cofactors 160 NADPH 196
Enzymes are most effective at optimal conditions 161 ATP synthesis occurs by chemiosmosis 196
Enzymes are organized into teams in metabolic pathways 162 9.5 The Carbon Fixation Reactions 198
The cell regulates enzymatic activity 162 Most plants use the Calvin cycle to fix
Enzymes are inhibited by certain chemical agents 163 carbon 198
Some drugs are enzyme inhibitors 164 Photorespiration reduces photosynthetic
efficiency 200
The initial carbon fixation step differs in C4 plants and in
8 How Cells Make ATP: CAM plants 200
Energy-Releasing Pathways 167 CAM plants fix CO2 at night 202
8.1 Redox Reactions 168 9.6 Metabolic Diversity 202
8.2 The Four Stages of Aerobic Respiration 168 9.7 Photosynthesis in Plants and in the
In glycolysis, glucose yields two pyruvates 170 Environment 203

viii / Contents

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part three: The Continuity of Life: Genetics

10 Chromosomes, Mitosis, and The rules of probability can be applied to a variety of


Meiosis 206 calculations 238
10.1 Eukaryotic Chromosomes 207 11.3 Inheritance and Chromosomes 240
DNA is organized into informational units called genes 207 Linked genes do not assort independently 240
DNA is packaged in a highly organized way in Calculating the frequency of crossing-over reveals the linear
chromosomes 207 order of linked genes on a chromosome 240
Chromosome number and informational content differ Sex is generally determined by sex chromosomes 241
among species 208 11.4 Extensions of Mendelian Genetics 246
10.2 The Cell Cycle and Mitosis 210 Dominance is not always complete 246
Chromosomes duplicate during interphase 210 Multiple alleles for a locus may exist in a population 248
During prophase, duplicated chromosomes become visible A single gene may affect multiple aspects of the phenotype 248
with the microscope 211 Alleles of different loci may interact to produce a
Prometaphase begins when the nuclear envelope breaks phenotype 248
down 211 In polygenic inheritance, the offspring exhibit a continuous
Duplicated chromosomes line up on the midplane during variation in phenotypes 249
metaphase 212 Genes interact with the environment to shape phenotype 250
During anaphase, chromosomes move toward the
poles 213
During telophase, two separate nuclei form 215 12 DNA: The Carrier of Genetic
Cytokinesis forms two separate daughter cells 215 Information 253
Mitosis produces two cells genetically identical to the 12.1 Evidence of Dna as the Hereditary Material 254
parent cell 215 DNA is the transforming factor in bacteria 254
Lacking nuclei, prokaryotes divide by binary fission 216 DNA is the genetic material in certain viruses 254
10.3 Regulation of the Cell Cycle 217 12.2 The Structure of Dna 257
10.4 Sexual Reproduction and Meiosis 219 Nucleotides can be covalently linked in any order to form long
Meiosis produces haploid cells with unique gene polymers 257
combinations 220 DNA is made of two polynucleotide chains intertwined to
Prophase I includes synapsis and crossing-over 221 form a double helix 258
During meiosis I, homologous chromosomes separate 221 In double-stranded DNA, hydrogen bonds form between
Chromatids separate in meiosis II 222 A and T and between G and C 261
Mitosis and meiosis lead to contrasting outcomes 223 12.3 DNA Replication 261
10.5 Sexual Life Cycles 224 Meselson and Stahl verified the mechanism of
­semiconservative replication 262
Semiconservative replication explains the perpetuation of
11 The Basic Principles of Heredity 228 mutations 262
11.1 Mendel’s Principles of Inheritance 229 DNA replication requires protein “machinery” 263
Alleles separate before gametes are formed: the principle Enzymes proofread and repair errors in DNA 268
of segregation 231 Telomeres cap eukaryotic chromosome ends 268
Alleles occupy corresponding loci on homologous
chromosomes 232
A monohybrid cross involves individuals with different alleles 13 Gene Expression 272
of a given locus 233 13.1 Discovery of the Gene–Protein Relationship 273
A dihybrid cross involves individuals that have different alleles Beadle and Tatum proposed the one-gene, one-enzyme
at two loci 235 hypothesis 273
Alleles on nonhomologous chromosomes are randomly 13.2 Information Flow from Dna to Protein:
distributed into gametes: the principle of independent An Overview 275
assortment 236 DNA is transcribed to form RNA 276
Recognition of Mendel’s work came during the early RNA is translated to form a polypeptide 276
20th century 236 Biologists cracked the genetic code in the 1960s 277
11.2 Using Probability to Predict Mendelian The genetic code is virtually universal 278
Inheritance 238 The genetic code is redundant 278

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13.3 Transcription 279 15 DNA Technology and Genomics 315
The synthesis of mRNA includes initiation, elongation, 15.1 Dna Cloning 316
and termination 280 Restriction enzymes are “molecular scissors” used to
Messenger RNA contains base sequences that do not directly ­construct recombinant DNA molecules 316
code for protein 281 Recombinant DNA is formed when DNA is spliced into a
Eukaryotic mRNA is modified after transcription and before vector 317
translation 282 Scientists use restriction enzymes and gel electrophoresis to
Biologists debate the evolution of eukaryotic gene examine cloned DNA fragments 318
structure 282 The polymerase chain reaction amplifies DNA
13.4 Translation 284 in vitro 318
An amino acid is attached to tRNA before incorporation into a cDNA clones do not contain introns 319
polypeptide 284 15.2 CRISPR-Based Technologies 321
The components of the translational machinery come CRISPR-based technologies can be used to edit genes in
­together at the ribosomes 285 growing cells 321
Translation begins with the formation of an initiation CRISPR-based tools exploit host DNA repair systems to
complex 285 ­perform many types of recombinant DNA functions 322
During elongation, amino acids are added to the growing Engineered CRISPR systems are used for specialized research
polypeptide chain 286 applications 322
One of three stop codons signals the termination of 15.3 Tools for Studying Dna 323
translation 288 DNA, RNA, and protein blots detect differences in related
Transcription and translation are coupled in bacteria 288 molecules separated by gel electrophoresis 324
13.5 Mutations 290 Automated DNA sequencing methods have been
Base-pair substitution mutations result from the replacement developed 324
of one base pair by another 290 Gene databases are powerful research tools 325
Frameshift mutations result from the insertion or deletion of Reverse transcription of mRNA to cDNA is used to measure
base pairs 290 gene expression in numerous ways 326
Some mutations involve mobile genetic elements 290 15.4 Genomics 328
Mutations have various causes 292 Collaborative genome-wide association studies have radically
13.6 Variations in Gene Expression 292 changed our view of the human genome 328
Many eukaryotic genes produce “non-coding” RNAs with Comparative genomic databases are tools for uncovering
catalytic, regulatory, or other cellular functions 292 gene functions 328
The definition of a gene has evolved 293 RNA interference is used to study gene functions 329
The usual direction of information flow has exceptions 293 15.5 Applications of Dna Technologies 330
DNA technology has revolutionized medicine 330
DNA fingerprinting has numerous applications 331
14 Gene Regulation 297
Transgenic organisms have many research and technological
14.1 Gene Regulation in Bacteria and Eukaryotes:
applications 331
An Overview 298
15.6 CRISPR-Based Gene Drives 334
14.2 Gene Regulation in Bacteria 299
15.7 Dna Technology and Safety Concerns 336
Operons in bacteria facilitate the coordinated control of
­functionally related genes 299
Some posttranscriptional regulation occurs in 16 Human Genetics
bacteria 303 and the Human Genome 340
14.3 Gene Regulation in Eukaryotic Cells 304 16.1 Studying Human Genetics 341
Eukaryotic transcription is controlled at many sites and by Human chromosomes are studied by karyotyping 341
many regulatory molecules 305 Family pedigrees help identify certain inherited
Chromosome organization affects the expression of some conditions 342
genes 307 Human gene databases allow geneticists to map the locations
Long non-coding RNAs (lncRNAs) regulate transcription over of genes on chromosomes 342
long distances within the genome 309 16.2 Abnormalities in Chromosome Number
The mRNAs of eukaryotes are subject to many types of and Structure 344
­posttranscriptional control 309 Down syndrome is usually caused by trisomy 21 345
Posttranslational chemical modifications may alter the activity Most sex chromosome aneuploidies are less severe than
of eukaryotic proteins 312 autosomal aneuploidies 347

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Abnormalities in chromosome structure cause certain 17 Developmental Genetics 362
disorders 348 17.1 Cell Differentiation and Nuclear
Genomic imprinting may determine whether inheritance is Equivalence 363
from the male or female parent 349 Most cell differences are due to differential gene
16.3 Genetic Diseases Caused by Single-Gene expression 363
Mutations 351 A totipotent nucleus contains all the instructions for
Many genetic diseases are inherited as autosomal development 364
recessive traits 351 The first cloned mammal was a sheep 366
Some genetic diseases are inherited as autosomal Stem cells divide and give rise to differentiated
dominant traits 353 cells 367
Some genetic diseases are inherited as X-linked 17.2 The Genetic Control of Development 369
recessive traits 353 A variety of model organisms provide insights into basic
16.4 Gene Therapy 354 ­biological processes 369
Performing clinical trials on humans always has inherent Many genes that control development have been identified in
risks 354 the fruit fly 369
16.5 Genetic Testing and Counseling 355 Caenorhabditis elegans has a relatively rigid developmental
Prenatal diagnosis detects chromosome abnormalities and pattern 374
gene defects 355 The mouse is a model for mammalian development 377
Preimplanation genetic diagnosis is used to screen embryos Arabidopsis is a model for studying plant development,
produced by in vitro fertilization 356 ­including transcription factors 379
Genetic screening searches for genotypes or karyotypes 356 17.3 Cancer and Cell Development 380
Genetic counselors educate people about genetic Oncogenes are usually altered components of cell
diseases 357 ­signaling pathways that control growth and
16.6 Human Genetics, Society, and Ethics 357 differentiation 381
Genetic discrimination provokes heated debate 358 In many familial cancers, tumor suppressor genes must be
Many ethical issues related to human genetics must be inactivated before cells progress to cancer 382
addressed 358 Cancer cells evolve by accumulating new mutations 382

part four: The CONTINUITY of Life: Evolution

18 Introduction to Darwinian 19 Evolutionary Change in


Evolution 385 Populations 406
18.1 What Is Evolution? 386 19.1 Genotype, Phenotype, and Allele
18.2 Pre-Darwinian Ideas about Evolution 386 Frequencies 407
18.3 Darwin and Evolution 387 19.2 The Hardy–Weinberg Principle 407
Darwin proposed that evolution occurs by natural Genetic equilibrium occurs if certain conditions are met 409
selection 389 Human MN blood groups are a valuable illustration of the
The modern synthesis combines Darwin’s scientific theory of Hardy–Weinberg principle 409
evolution with genetics 390 19.3 Microevolution 410
Biologists study the effect of chance on evolution 390 Nonrandom mating changes genotype frequencies 410
18.4 Evidence for Evolution 391 Mutation increases variation within a population 410
The fossil record provides strong evidence for In genetic drift, random events change allele frequencies 411
evolution 391 Gene flow generally increases variation within a population 412
The distribution of plants and animals supports Natural selection changes allele frequencies in a way that
evolution 395 increases adaptation 412
Comparative anatomy of related species demonstrates 19.4 Genetic Variation in Populations 415
­similarities in their structures 396 Genetic polymorphism can be studied in several ways 415
Molecular comparisons among organisms provide evidence Balanced polymorphism exists for long periods 416
for evolution 399 Neutral variation may give no selective advantage or
Developmental biology helps unravel evolutionary disadvantage 418
patterns 400 Populations in different geographic areas often exhibit genetic
Evolutionary hypotheses are tested experimentally 401 adaptations to local environments 418

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20 Speciation and Macroevolution 421 Biological evolution began with the first cells 447
20.1 What is a Species? 422 Photosynthesis was a further step in the evolution of
The biological species concept is based on reproductive cells 448
isolation 422 Aerobes appeared after oxygen increased in the
The phylogenetic species concept defines species based on atmosphere 449
such evidence as molecular sequencing 422 Eukaryotic cells descended from prokaryotic cells 450
20.2 Reproductive Isolation 423 21.3 The History of Life 451
Prezygotic barriers interfere with fertilization 423 Rocks from the Ediacaran period contain fossils of cells and
Postzygotic barriers prevent gene flow when fertilization simple animals 451
occurs 425 A diversity of organisms evolved during the Paleozoic era 451
Biologists are discovering genes responsible for reproductive Dinosaurs and other reptiles dominated the Mesozoic era 454
isolating mechanisms 425 The Cenozoic era is the age of mammals 457
20.3 Speciation 425
Long physical isolation and different selective pressures result
in allopatric speciation 427 22 The Evolution of Primates 461
The evolutionary importance of hybridization is being 22.1 Primate Adaptations 462
re-evaluated 432 22.2 Primate Classification 462
20.4 The Rate of Evolutionary Change 434 Suborder Anthropoidea includes monkeys, apes, and
20.5 Macroevolution 435 humans 463
Evolutionary novelties originate through modifications of Apes are our closest living relatives 465
pre-existing structures 435 22.3 Hominin Evolution 467
Adaptive radiation is the diversification of an ancestral The earliest hominins may have lived 6 mya to 7 mya 468
­species into many species 436 Ardipithecus, Australopithecus, and Paranthropus are
Extinction is an important aspect of evolution 438 ­australopithecines, or “southern man apes” 468
Is microevolution related to speciation and Homo habilis is considered the oldest member of genus
macroevolution? 439 Homo 470
Homo ergaster may have arisen from H. habilis 471
Homo erectus probably evolved from H. ergaster 471
21 The Origin and Evolutionary Archaic humans date from about 1.2 mya to 200,000 years
History of Life 442 ago 471
21.1 Chemical Evolution on Early Earth 443 Neandertals appeared approximately 250,000 years ago 472
Organic molecules formed on primitive Earth 443 Scientists have reached a near consensus on the origin of
21.2 The First Cells 445 modern H. sapiens 473
The origin of a simple metabolism within a membrane 22.4 Cultural Change 474
­boundary may have occurred early in the evolution Development of agriculture resulted in a more dependable
of cells 445 food supply 475
Molecular reproduction was a crucial step in the origin Human culture has had a profound effect on the
of cells 445 biosphere 475

part five: The Diversity of Life

23 Understanding Diversity: Some biologists are moving away from Linnaean


Systematics 478 categories 482
23.1 Classifying Organisms 479 Phylogenetic trees show hypothesized evolutionary
Organisms are named using a binomial system 479 relationships 483
Each taxonomic level is more general than the one Systematists continue to consider other hypotheses 484
below it 480 23.3 Reconstructing Evolutionary History 485
23.2 Determining the Major Branches in the Tree Homologous structures are important in determining
of Life 480 ­evolutionary relationships 485
Systematics is an evolving science 480 Shared derived characters provide clues about phylogeny 486
The three domains form the three main branches of the tree Systematists base taxonomic decisions on recent shared
of life 482 ancestry 487

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Molecular homologies help clarify phylogeny 487 25.4 the phylogeny of the two prokaryote
Taxa are grouped based on their evolutionary domains 525
relationships 488 Key characters distinguish the three domains 526
23.4 Constructing Phylogenetic Trees 490 Taxonomy of archaea and bacteria continuously changes 526
Outgroup analysis is used in constructing and interpreting Most archaea live in marine and soil habitats, and many
cladograms 490 thrive in harsh environments 527
A cladogram is constructed by considering shared derived Bacteria are the most familiar prokaryotes 528
characters 491 25.5 Impact on Ecology, Technology, and
Each branch point represents a major evolutionary step 491 Commerce 528
Systematists use the principles of parsimony and maximum Prokaryotes form intimate relationships with other
likelihood to make decisions 494 organisms 529
23.5 Applying Phylogenetic Information 495 Prokaryotes play key ecological roles 529
Prokaryotes are important in many commercial processes and
in technology 532
24 Viruses and Subviral Agents 499 25.6 bacteria and disease 533
24.1 The Status and Structure of Viruses 500 Many scientists have contributed to our understanding of
Viruses are very small 500 infectious disease 533
A virus consists of nucleic acid surrounded by a Many adaptations contribute to pathogen success 533
protein coat 500 Antibiotic resistance is a major public health problem 535
The capsid is a protective protein coat 501
Some viruses are surrounded by an envelope 502
24.2 Classification of Viruses 502 26 Protists 539
24.3 Viral Replication 503 26.1 diversity in the protists 540
Bacteriophages infect bacteria 503 26.2 how did eukaryotes evolve? 541
Viruses replicate inside host cells 503 Mitochondria and chloroplasts probably originated
24.4 Viral Diseases 504 from endosymbionts 541
Viruses cause serious plant diseases 505 A consensus in eukaryote classification is beginning
Viruses cause serious diseases in animals 505 to emerge 541
24.5 Evolution of Viruses 511 26.3 excavates 544
24.6 Subviral Agents 512 Diplomonads are small, mostly parasitic flagellates 544
Satellites depend on helper viruses 512 Parabasilids are anaerobic endosymbionts that live in
Viroids are short, single strands of naked RNA 513 animals 544
Prions are protein particles 513 Euglenoids and trypanosomes include both free-living species
Defective interfering particles are virus mutants 514 and parasites 545
26.4 chromalveolates 546
Most dinoflagellates are a part of marine plankton 546
25 Bacteria and Archaea 517 Apicomplexans are spore-forming parasites of animals 547
25.1 The Structure of Bacteria and Archaea 518 Ciliates use cilia for locomotion 548
Prokaryotes have several common shapes 518 Water molds produce biflagellate reproductive cells 549
Prokaryotic cells do not have membrane-enclosed Diatoms are stramenopiles with shells composed of
organelles 518 two parts 551
A cell wall protects most prokaryotes 519 Brown algae are multicellular stramenopiles 551
Some bacteria produce capsules or slime layers 520 Most golden algae are unicellular biflagellates 552
Some prokaryotes have fimbriae or pili 520 26.5 rhizarians 553
Some bacteria survive unfavorable conditions by forming Forams extend cytoplasmic projections that form a threadlike,
endospores 520 interconnected net 553
Many types of prokaryotes are motile 521 Actinopods project slender axopods 553
25.2 Prokaryote Reproduction and Evolution 522 26.6 archaeplastids 554
Rapid reproduction contributes to prokaryote success 522 Red algae do not produce motile cells 554
Prokaryotes transfer genetic information 522 Green algae share many similarities with land plants 555
Evolution proceeds rapidly in prokaryote populations 523 26.7 unikonts 555
25.3 Nutritional and Metabolic Adaptations 524 Amoebozoa are unikonts with lobose pseudopodia 556
Most prokaryotes require oxygen 525 Choanoflagellates are opisthokonts closely related to
Some prokaryotes fix and metabolize nitrogen 525 animals 558

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27 Seedless Plants 563 29 The Fungi 603
27.1 adaptations of plants to life on land 564 29.1 Characteristics of Fungi 604
The plant life cycle alternates between haploid and diploid Fungi absorb food from the environment 604
generations 564 Fungi have cell walls that contain chitin 604
Four major groups of plants exist today 565 Most fungi consist of a network of filaments 604
27.2 bryophytes 568 29.2 Fungal Reproduction 605
Moss gametophytes are differentiated into “leaves” Many fungi reproduce asexually 605
and “stems” 568 Most fungi reproduce sexually 605
Liverwort gametophytes are either thalloid or leafy 571 29.3 Fungal Diversity 607
Hornwort gametophytes are inconspicuous thalloid Fungi are assigned to the opisthokont clade 607
plants 572 Diverse groups of fungi have evolved 607
Bryophytes are used for experimental studies 572 Chytrids have flagellate spores 608
Recap: details of bryophyte evolution are based on fossils and Zygomycetes reproduce sexually by forming
on structural and molecular evidence 573 zygospores 609
27.3 seedless vascular plants 574 Microsporidia have been a taxonomic mystery 610
Club mosses are small plants with rhizomes and short, Glomeromycetes have a symbiotic relationship with
erect branches 574 plant roots 611
Ferns are a diverse group of spore-forming vascular Ascomycetes reproduce sexually by forming
plants 575 ascospores 612
Whisk ferns are classified as reduced ferns 576 Basidiomycetes reproduce sexually by forming
Horsetails are an evolutionary line of ferns 576 basidiospores 615
Some ferns and club mosses are heterosporous 577 29.4 Ecological Importance of Fungi 618
Seedless vascular plants are used for experimental studies 578 Fungi form symbiotic relationships with some
Seedless vascular plants arose more than 420 mya 580 animals 618
Mycorrhizae are symbiotic associations between fungi
and plant roots 618
28 Seed Plants 584 A lichen consists of one or more fungi and a
28.1 an introduction to seed plants 585 photoautotroph 619
28.2 gymnosperms 586 29.5 Economic, Biological, and Medical Impact
Conifers are woody plants that produce seeds of Fungi 621
in cones 586 Fungi provide beverages and food 621
Pines represent a typical conifer life cycle 588 Fungi are important to modern biology and medicine 622
Cycads have seed cones and compound leaves 589 Fungi are used in bioremediation and to biologically
Ginkgo biloba is the only living species in its control pests 623
phylum 590 Some fungi cause diseases in humans and other
Gnetophytes include three unusual genera 591 animals 623
28.3 flowering plants 591 Fungi cause many important plant diseases 623
Monocots and eudicots are the two largest classes of
flowering plants 592
Sexual reproduction takes place in flowers 593 30 An Introduction to Animal
The life cycle of flowering plants includes double Diversity 628
fertilization 594 30.1 Animal Characteristics 629
Seeds and fruits develop after fertilization 596 30.2 Adaptations to Ocean, Freshwater, and
Flowering plants have many adaptations that account for ­Terrestrial Habitats 630
their success 596 Marine habitats offer many advantages 630
Floral structure provides insights into the evolutionary Some animals are adapted to freshwater
process 596 habitats 630
28.4 the evolution of seed plants 597 Terrestrial living requires major adaptations 630
Our understanding of the evolution of flowering plants has 30.3 Animal Evolution 631
made great progress in recent years 597 Molecular systematics helps biologists interpret the
The basal angiosperms comprise three clades 599 fossil record 631
The core angiosperms comprise magnoliids, monocots, Biologists develop hypotheses about the evolution
and eudicots 600 of development 631

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30.4 Reconstructing Animal Phylogeny 632 Many sea stars capture prey 678
Animals exhibit two main types of body symmetry 632 Basket stars and brittle stars make up the largest group
Animal body plans are linked to the level of tissue of echinoderms 680
development 633 Sea urchins and sand dollars have movable spines 680
Most bilateral animals have a body cavity lined with Sea cucumbers are elongated, sluggish animals 680
mesoderm 634 32.3 The Chordates: Major Characteristics 681
Bilateral animals form two main clades based on differences 32.4 Invertebrate Chordates 682
in development 635 Tunicates are common marine animals 682
Biologists have identified major animal clades based on Lancelets clearly exhibit chordate characteristics 682
­structure, development, and molecular data 635 Systematists debate chordate phylogeny 683
Segmentation apparently evolved three times 636 32.5 Introducing the Vertebrates 684
The vertebral column is a derived vertebrate character 684
Vertebrate taxonomy is a work in progress 686
31 Sponges, Cnidarians, Ctenophores, 32.6 Jawless Fishes 686
and Protostomes 641 32.7 Evolution of Jaws and Limbs: Jawed Fishes and
31.1 Sponges, Cnidarians, and Ctenophores 642 Tetrapods 688
Sponges have collar cells and other specialized cells 642 Most cartilaginous fishes inhabit marine environments 688
Cnidarians have unique stinging cells 644 Ray-finned fishes gave rise to modern bony fishes 690
Comb jellies have adhesive glue cells that trap prey 648 Tetrapods evolved from sarcopterygian ancestors 691
31.2 The Lophotrochozoa 649 Amphibians were the first successful land
Flatworms are bilateral acoelomates 649 vertebrates 693
Nemerteans are characterized by their proboscis 652 32.8 Amniotes: Terrestrial Vertebrates 694
Mollusks have a muscular foot, visceral mass, and mantle 653 Our understanding of amniote phylogeny is changing 695
Annelids are segmented worms 657 Reptiles have many terrestrial adaptations 695
The lophophorates are distinguished by a ciliated ring Biologists assign reptiles to two major lineages 696
of tentacles 659 Lizards and snakes are common modern reptiles 696
Rotifers have a crown of cilia 661 Tuataras superficially resemble lizards 698
31.3 The Ecdysozoa 662 Turtles have protective shells 698
Roundworms are of great ecological importance 662 Crocodilians have an elongated skull 699
Arthropods are characterized by jointed appendages and How do we know that birds are really dinosaurs? 699
an exoskeleton of chitin 662 Early birds were transitional forms 699
Modern birds are adapted for flight 700
Mammals (class Mammalia) have many unique
32 The Deuterostomes 676 characters 702
32.1 What are Deuterostomes? 677 New fossil discoveries are changing our understanding of
32.2 Echinoderms 677 the early evolution of mammals 702
Feather stars and sea lilies are suspension feeders 678 Modern mammals are assigned to three subclasses 703

part six: Structure and Life Processes in Plants

33 Plant Structure, Growth, and The orientation of cellulose microfibrils affects the direction
Development 710 of cell expansion 724
33.1 The Plant Body 711 Cell differentiation depends in part on a cell’s location 724
The plant body consists of cells and tissues 711 Morphogenesis occurs through pattern formation 725
The ground tissue system is composed of three simple tissues 711
The vascular tissue system consists of two complex tissues 716
The dermal tissue system consists of two complex tissues 718 34 Leaf Structure and Function 729
33.2 Plant Meristems 720 34.1 Leaf Form and Structure 730
Primary growth takes place at apical meristems 721 Leaf structure is adapted for maximum light absorption 730
Secondary growth takes place at lateral meristems 721 34.2 Stomatal Opening and Closing 736
33.3 Development of Form 722 Blue light triggers stomatal opening 736
The plane and symmetry of cell division affect plant form 723 Additional factors affect stomatal opening and closing 737

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34.3 Transpiration and Guttation 737 Many plants have mechanisms that prevent
Some plants exude liquid water 738 self-pollination 786
34.4 Leaf Abscission 739 Flowering plants and their animal pollinators have
In many leaves, abscission occurs at an abscission zone coevolved 786
near the base of the petiole 739 Some flowering plants depend on wind to disperse
34.5 Modified Leaves 740 pollen 788
Modified leaves of carnivorous plants capture insects 742 37.3 Fertilization and Seed and Fruit
Development 789
A unique double fertilization process occurs in flowering
35 Stem Structure and Transport 745 plants 790
35.1 Stem Growth and Structure 746 Embryonic development in seeds is orderly and
Herbaceous eudicot and monocot stems differ in internal predictable 790
structure 746 The mature seed contains an embryonic plant and storage
Woody plants have stems with secondary growth 748 materials 791
35.2 Water Transport 754 Fruits are mature, ripened ovaries 792
Water and minerals are transported in xylem 754 Seed dispersal is highly varied 794
Water movement can be explained by a difference in water 37.4 Germination and Early Growth 796
potential 755 Some seeds do not germinate immediately 797
According to the tension–cohesion model, water is pulled up Eudicots and monocots exhibit characteristic patterns of
a stem 755 early growth 797
Root pressure pushes water from the root up a stem 756 37.5 Asexual Reproduction in Flowering Plants 797
35.3 Translocation of Sugar in Solution 757 Apomixis is the production of seeds without the sexual
The pressure–flow model explains translocation in process 799
phloem 757 37.6 A Comparison of Sexual and Asexual
Reproduction 800
Sexual reproduction has some disadvantages 800
36 Roots and Mineral Nutrition 762
36.1 Root Structure and Function 763
Roots have root caps and root hairs 763 38 Plant Developmental Responses
The arrangement of vascular tissues distinguishes the roots of to ­External and Internal Signals 803
herbaceous eudicots and monocots 764 38.1 Tropisms 804
Woody plants have roots with secondary growth 767 38.2 Plant Hormones and Development 805
Some roots are specialized for unusual functions 768 Plant hormones act by signal transduction 805
36.2 Root Associations and Interactions 769 Auxins promote cell elongation 807
Mycorrhizae facilitate the uptake of essential minerals by Gibberellins promote stem elongation 809
roots 771 Cytokinins promote cell division 810
Rhizobial bacteria fix nitrogen in the roots of leguminous Ethylene promotes abscission and fruit ripening 811
plants 772 Abscisic acid promotes seed dormancy 812
36.3 The Soil Environment 773 Brassinosteroids are plant steroid hormones 812
Soil comprises inorganic minerals, organic matter, air, and Identification of a universal flower-promoting signal remains
water 773 elusive 813
About 50% of soil volume is composed of pore spaces 775 38.3 Light Signals and Plant Development 813
Soil organisms form a complex ecosystem 775 Phytochrome detects day length 814
Soil pH affects soil characteristics and plant growth 775 Competition for sunlight among shade-avoiding plants
Soil provides most of the minerals found in plants 776 involves phytochrome 815
Soil can be damaged by human mismanagement 778 Phytochrome is involved in other responses to light,
including germination 816
Phytochrome acts by signal transduction 816
37 Reproduction in Flowering Plants 782 Light influences circadian rhythms 816
37.1 The Flowering Plant Life Cycle 783 38.4 Responses to Herbivores and Pathogens 817
Flowers develop at apical meristems 783 Jasmonic acid activates several plant defenses 818
Each part of a flower has a specific function 783 Methyl salicylate may induce systemic acquired
37.2 Pollination 786 resistance 818

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part seven: Structure and Life Processes in Animals

39 Animal Structure and Function: 41 Neural Signaling 860


An Introduction 821 41.1 Neural Signaling: an Overview 861
39.1 Tissues, Organs, and Organ Systems 822 41.2 Neurons and Glial Cells 862
Epithelial tissues cover the body and line its Neurons receive stimuli and transmit neural signals 862
cavities 822 Certain regions of the CNS produce new neurons 862
Glands are made of epithelial cells 823 Axons aggregate to form nerves and tracts 863
Epithelial cells form membranes 823 Glial cells play critical roles in neural function 863
Connective tissues support other body 41.3 Transmitting Information along the Neuron 865
structures 823 Ion channels and pumps maintain the resting potential of the
Muscle tissue is specialized to contract 828 neuron 865
Nervous tissue controls muscles and glands 829 Ions cross the plasma membrane by diffusion through ion
Tissues and organs make up the organ systems of the channels 866
body 830 Ion pumping maintains the gradients that determine the
39.2 Regulating the Internal Environment 834 resting potential 867
Negative feedback systems restore homeostasis 834 Graded local signals vary in magnitude 867
A few positive feedback systems operate in the body 835 Axons transmit signals called action potentials 867
39.3 Regulating Body Temperature 836 An action potential is generated when the voltage reaches
Ectotherms absorb heat from their surroundings 836 threshold level 867
Endotherms derive heat from metabolic processes 836 The neuron repolarizes and returns to a resting state 868
Many animals adjust to challenging temperature changes 839 The action potential is an all-or-none response 869
An action potential is self-propagating 870
Several factors determine the velocity of an action
40 Protection, Support, and potential 871
Movement 842 41.4 Transmitting Information across Synapses 872
40.1 Epithelial Coverings 843 Signals across synapses can be electrical or chemical 872
Invertebrate epithelium may secrete a cuticle 843 Neurons use neurotransmitters to signal other cells 873
Vertebrate skin functions in protection and temperature Neurotransmitters bind with receptors on postsynaptic
regulation 843 cells 873
40.2 Skeletal Systems 844 Activated receptors can send excitatory or inhibitory
In hydrostatic skeletons body fluids transmit force 844 signals 874
Mollusks and arthropods have nonliving exoskeletons 845 41.5 Neural Integration 877
Internal skeletons are capable of growth 845 Postsynaptic potentials are summed over time and space 877
The vertebrate skeleton has two main divisions 846 Where does neural integration take place? 877
A typical long bone amplifies the motion generated by 41.6 Neural Circuits: Complex Information
muscles 846 Signaling 877
Bones are remodeled throughout life 847
Joints are junctions between bones 847
40.3 Muscle Contraction 848 42 Neural Regulation 882
Invertebrate muscle varies among groups 848 42.1 Invertebrate Nervous Systems: Trends in
Vertebrate skeletal muscles act antagonistically to one Evolution 883
another 849 42.2 Overview of the Vertebrate Nervous System 884
A vertebrate muscle may consist of thousands of muscle 42.3 Evolution of the Vertebrate Brain 885
fibers 849 The hindbrain develops into the medulla, pons, and
Contraction occurs when actin and myosin filaments move cerebellum 886
past one another 850 The midbrain is prominent in fishes and amphibians 886
ATP powers muscle contraction 853 The forebrain gives rise to the thalamus, hypothalamus, and
The type of muscle fibers determines strength and cerebrum 887
endurance 855 42.4 The Human Central Nervous System 888
Several factors influence the strength of muscle The spinal cord transmits impulses to and from the brain 888
contraction 855 The most prominent part of the human brain is the
Smooth muscle and cardiac muscle are involuntary 856 cerebrum 889
Contents / xvii

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the ax missed his head by the fraction of a second. Then with a whir
the arm rose stiffly again to its original position and remained rigid.
The visitor moistened his lips and sighed.
“That’s a new one, a very new one,” he said under his breath, and
the admiration in his tone was evident. He picked up his overcoat,
flung it over his arm, and mounted half a dozen steps to the next
landing. The inspection of the Chinese cabinet was satisfactory.
The white beam of his lamp flashed into corners and crevices and
showed nothing. He shook the curtain of a window and listened,
holding his breath.
“Not here,” he muttered decisively, “the old man wouldn’t try that
game. Snakes turned loose in a house in London, S.W., take a deal
of collecting in the morning.”
He looked round. From the landing access was gained to three
rooms. That which from its position he surmised faced the street he
did not attempt to enter. The second, covered by a heavy curtain, he
looked at for a time in thought. To the third he walked, and carefully
swathing the door-handle with his silk muffler, he turned it. The door
yielded. He hesitated another moment, and jerking the door wide
open, sprang backward.
The interior of the room was for a second only in pitch darkness,
save for the flicker of light that told of an open fireplace. Then the
visitor heard a click, and the room was flooded with light. In the
darkness on the landing the man waited; then a voice, a cracked old
voice, said grumblingly—
“Come in.”
Still the man on the landing waited.
“Oh, come in, Jimmy—I know ye.”
Cautiously the man outside stepped through the entry into the light
and faced the old man, who, arrayed in a wadded dressing-gown,
sat in a big chair by the fire—an old man, with white face and a
sneering grin, who sat with his lap full of papers.
The visitor nodded a friendly greeting.
“As far as I can gather,” he said deliberately, “we are just above your
dressing-room, and if you dropped me through one of your patent
traps, Reale, I should fetch up amongst your priceless china.”
Save for a momentary look of alarm on the old man’s face at the
mention of the china, he preserved an imperturbable calm, never
moving his eyes from his visitor’s face. Then his grin returned, and
he motioned the other to a chair on the other side of the fireplace.
Jimmy turned the cushion over with the point of his stick and sat
down.
“Suspicious?”—the grin broadened—“suspicious of your old friend,
Jimmy? The old governor, eh?”
Jimmy made no reply for a moment, then—
“You’re a wonder, governor, upon my word you are a wonder. That
man in armor—your idea?”
The old man shook his head regretfully.
“Not mine entirely, Jimmy. Ye see, there’s electricity in it, and I don’t
know much about electricity. I never did, except——”
“Except?” suggested the visitor.
“Oh, that roulette board, that was my own idea; but that was
magnetism, which is different to electricity, by my way of looking.”
Jimmy nodded.
“Ye got past the trap?” The old man had just a glint of admiration in
his eye.
“Yes, jumped it.”
The old man nodded approvingly.
“You always was a one for thinkin’ things out. I’ve known lots of ’em
who would never have thought of jumping it. Connor, and that pig
Massey, they’d have walked right on to it. You didn’t damage
anything?” he demanded suddenly and fiercely. “I heard somethin’
break, an’ I was hoping that it was you.”
Jimmy thought of the marble statue, and remembered that it had
looked valuable.
“Nothing at all,” he lied easily, and the old man’s tense look relaxed.
The pair sat on opposite sides of the fireplace, neither speaking for
fully ten minutes; then Jimmy leant forward.
“Reale,” he said quietly, “how much are you worth?”
In no manner disturbed by this leading question, but rather indicating
a lively satisfaction, the other replied instantly—
“Two millions an’ a bit over, Jimmy. I’ve got the figures in my head.
Reckonin’ furniture and the things in this house at their proper value,
two millions, and forty-seven thousand and forty-three pounds—
floatin’, Jimmy, absolute cash, the same as you might put your hand
in your pocket an’ spend—a million an’ three-quarters exact.”
He leant back in his chair with a triumphant grin and watched his
visitor.
Jimmy had taken a cigarette from his pocket and was lighting it,
looking at the slowly burning match reflectively.
“A million and three-quarters,” he repeated calmly, “is a lot of
money.”
Old Reale chuckled softly.
“All made out of the confiding public, with the aid of me—and Connor
and Massey——”
“Massey is a pig!” the old man interjected spitefully.
Jimmy puffed a cloud of tobacco smoke.
“Wrung with sweat and sorrow from foolish young men who backed
the tiger and played high at Reale’s Unrivaled Temple of Chance,
Cairo, Egypt—with branches at Alexandria, Port Said, and Suez.”
The figure in the wadded gown writhed in a paroxysm of silent
merriment.
“How many men have you ruined, Reale?” asked Jimmy.
“The Lord knows!” the old man answered cheerfully; “only three as I
knows of—two of ’em’s dead, one of ’em’s dying. The two that’s
dead left neither chick nor child; the dying one’s got a daughter.”
Jimmy eyed him through narrowed lids.
“Why this solicitude for the relatives—you’re not going——?”
As he spoke, as if anticipating a question, the old man was nodding
his head with feverish energy, and all the while his grin broadened.
“What a one you are for long words, Jimmy! You always was. That’s
how you managed to persuade your swell pals to come an’ try their
luck. Solicitude! What’s that mean? Frettin’ about ’em, d’ye mean?
Yes, that’s what I’m doin’—frettin’ about ’em. And I’m going to make,
what d’ye call it—you had it on the tip of your tongue a minute or two
ago?”
“Reparation?” suggested Jimmy.
Old Reale nodded delightedly.
“How?”
“Don’t you ask questions!” bullied the old man, his harsh voice rising.
“I ain’t asked you why you broke into my house in the middle of the
night, though I knew it was you who came the other day to check the
electric meter. I saw you, an’ I’ve been waitin’ for you ever since.”
“I knew all about that,” said Jimmy calmly, and flicked the ash of his
cigarette away with his little finger, “and I thought you would——”
Suddenly he stopped speaking and listened.
“Who’s in the house beside us?” he asked quickly, but the look on
the old man’s face reassured him.
“Nobody,” said Reale testily. “I’ve got a special house for the
servants, and they come in every morning after I’ve unfixed my—
burglar-alarms.” He grinned, and then a look of alarm came into his
face.
“The alarms!” he whispered; “you broke them when you came in,
Jimmy. I heard the signal. If there’s some one in the house we
shouldn’t know it now.”
They listened.
Down below in the hall something creaked, then the sound of a soft
thud came up.
“He’s skipped the rug,” whispered Jimmy, and switched out the light.
The two men heard a stealthy footstep on the stair, and waited.
There was the momentary glint of a light, and the sound of some one
breathing heavily. Jimmy leant over and whispered in the old man’s
ear.
Then, as the handle of the door was turned and the door pushed
open, Jimmy switched on the light.
The newcomer was a short, thick-set man with a broad, red face. He
wore a check suit of a particularly glaring pattern, and on the back of
his head was stuck a bowler hat, the narrow brim of which seemed
to emphasize the breadth of his face. A casual observer might have
placed him for a coarse, good-natured man of rude but boisterous
humor. The ethnological student would have known him at once for
what he was—a cruel man-beast without capacity for pity.
He started back as the lights went on, blinking a little, but his hand
held an automatic pistol that covered the occupants of the room.
“Put up your hands,” he growled. “Put ’em up!”
Neither man obeyed him. Jimmy was amused and looked it, stroking
his short beard with his white tapering fingers. The old man was fury
incarnate.
He it was that turned to Jimmy and croaked—
“What did I tell ye, Jimmy? What’ve I always said, Jimmy? Massey is
a pig—he’s got the manners of a pig. Faugh!”
“Put up your hands!” hissed the man with the pistol. “Put ’em up, or
I’ll put you both out!”
“If he’d come first, Jimmy!” Old Reale wrung his hands in his regret.
“S’pose he’d jumped the rug—any sneak-thief could have done that
—d’ye think he’d have spotted the man in armor? If you’d only get
the man in armor ready again.”
“Put your pistol down, Massey,” said Jimmy coolly, “unless you want
something to play with. Old man Reale’s too ill for the gymnastics
you suggest, and I’m not inclined to oblige you.”
The man blustered.
“By God, if you try any of your monkey tricks with me, either of you
——”
“Oh, I’m only a visitor like yourself,” said Jimmy, with a wave of his
hand; “and as to monkey tricks, why, I could have shot you before
you entered the room.”
Massey frowned, and stood twiddling his pistol.
“You will find a safety catch on the left side of the barrel,” continued
Jimmy, pointing to the pistol; “snick it up—you can always push it
down again with your thumb if you really mean business. You are not
my idea of a burglar. You breathe too noisily, and you are built too
clumsily; why, I heard you open the front door!”
The quiet contempt in the tone brought a deeper red into the man’s
face.
“Oh, you are a clever ’un, we know!” he began, and the old man,
who had recovered his self-command, motioned him to a chair.
“Sit down, Mister Massey,” he snapped; “sit down, my fine fellow, an’
tell us all the news. Jimmy an’ me was just speakin’ about you, me
an’ Jimmy was. We was saying what a fine gentleman you was”—his
voice grew shrill—“what a swine, what an overfed, lumbering fool of
a pig you was, Mister Massey!”
He sank back into the depths of his chair exhausted.
“Look here, governor,” began Massey again—he had laid his pistol
on a table by his side, and waved a large red hand to give point to
his remarks—“we don’t want any unpleasantness. I’ve been a good
friend to you, an’ so has Jimmy. We’ve done your dirty work for
years, me an’ Jimmy have, and Jimmy knows it”—turning with an
ingratiating smirk to the subject of his remarks—“and now we want a
bit of our own—that is all it amounts to, our own.”
Old Reale looked under his shaggy eyebrows to where Jimmy sat
with brooding eyes watching the fire.
“So it’s a plant, eh? You’re both in it. Jimmy comes first, he being the
clever one, an’ puts the lay nice an’ snug for the other feller.”
Jimmy shook his head.
“Wrong,” he said. He turned his head and took a long scrutiny of the
newcomer, and the amused contempt of his gaze was too apparent.
“Look at him!” he said at last. “Our dear Massey! Does he look the
sort of person I am likely to share confidence with?”
A cold passion seemed suddenly to possess him.
“It’s a coincidence that brought us both together.”
He rose and walked to where Massey sat, and stared down at him.
There was something in the look that sent Massey’s hand wandering
to his pistol.
“Massey, you dog!” he began, then checked himself with a laugh and
walked to the other end of the room. There was a tantalus with a
soda siphon, and he poured himself a stiff portion and sent the soda
fizzling into the tumbler. He held the glass to the light and looked at
the old man. There was a look on the old man’s face that he
remembered to have seen before. He drank his whisky and gave
utterance to old Reale’s thoughts.
“It’s no good, Reale, you’ve got to settle with Massey, but not the
way you’re thinking. We could put him away, but we should have to
put ourselves away too.” He paused. “And there’s me,” he added.
“And Connor,” said Massey thickly, “and Connor’s worse than me.
I’m reasonable, Reale; I’d take a fair share——”
“You would, would you?”
The old man was grinning again.
“Well, your share’s exactly a million an’ three-quarters in solid cash,
an’ a bit over two millions—all in.”
He paused to notice the effect of his words.
Jimmy’s calm annoyed him; Massey’s indifference was outrageous.
“An’ it’s Jimmy’s share, an’ Connor’s share, an’ it’s Miss Kathleen
Kent’s share.”
This time the effect was better. Into Jimmy’s inexpressive face had
crept a gleam of interest.
“Kent?” he asked quickly. “Wasn’t that the name of the man——?”
Old Reale chuckled.
“The very feller, Jimmy—the man who came in to lose a tenner, an’
lost ten thousand; who came in next night to get it back, and left his
lot. That’s the feller!”
He rubbed his lean hands, as at the memory of some pleasant
happening.
“Open that cupboard, Jimmy.” He pointed to an old-fashioned walnut
cabinet that stood near the door. “D’ye see anything—a thing that
looks like a windmill?”
Jimmy drew out a cardboard structure that was apparently a toy
working-model. He handled it carefully, and deposited it on the table
by the old man’s side. Old Reale touched it caressingly. With his little
finger he set a fly-wheel spinning, and tiny little pasteboard rods ran
to and fro, and little wooden wheels spun easily.
“That’s what I did with his money, invented a noo machine that went
by itself—perpetual motion. You can grin, Massey, but that’s what I
did with it. Five years’ work an’ a quarter of a million, that’s what that
little model means. I never found the secret out. I could always make
a machine that would go for hours with a little push, but it always
wanted the push. I’ve been a chap that went in for inventions and
puzzles. D’ye remember the table at Suez?”
He shot a sly glance at the men.
Massey was growing impatient as the reminiscences proceeded. He
had come that night with an object; he had taken a big risk, and had
not lost sight of the fact. Now he broke in—
“Damn your puzzles, Reale. What about me; never mind about
Jimmy. What’s all this rotten talk about two millions for each of us,
and this girl? When you broke up the place in Egypt you said we
should stand in when the time came. Well, the time’s come!”
“Nearly, nearly,” said Reale, with his death’s-head grin. “It’s nearly
come. You needn’t have troubled to see me. My lawyer’s got your
addresses. I’m nearly through,” he went on cheerfully; “dead I’ll be in
six months, as sure as—as death. Then you fellers will get the
money”—he spoke slowly to give effect to his words—“you Jimmy, or
Massey or Connor or the young lady. You say you don’t like puzzles,
Massey? Well, it’s a bad look out for you. Jimmy’s the clever un, an’
most likely he’ll get it; Connor’s artful, and he might get it from
Jimmy; but the young lady’s got the best chance, because women
are good at puzzles.”
“What in hell!” roared Massey, springing to his feet.
“Sit down!” It was Jimmy that spoke, and Massey obeyed.
“There’s a puzzle about these two millions,” Reale went on, and his
croaky voice, with its harsh cockney accent, grew raucous in his
enjoyment of Massey’s perplexity and Jimmy’s knit brows. “An’ the
one that finds the puzzle out, gets the money.”
Had he been less engrossed in his own amusement he would have
seen a change in Massey’s brute face that would have warned him.
“It’s in my will,” he went on. “I’m goin’ to set the sharps against the
flats; the touts of the gamblin’ hell—that’s you two fellers—against
the pigeons. Two of the biggest pigeons is dead, an’ one’s dying.
Well, he’s got a daughter; let’s see what she can do. When I’m dead
——”
“That’s now!” bellowed Massey, and leant over and struck the old
man.
Jimmy, on his feet, saw the gush of blood and the knife in Massey’s
hand, and reached for his pocket.
Massey’s pistol covered him, and the man’s face was a dreadful
thing to look upon.
“Hands up! It’s God’s truth I’ll kill you if you don’t!”
Jimmy’s hands went up.
“He’s got the money here,” breathed Massey, “somewhere in this
house.”
“You’re mad,” said the other contemptuously. “Why did you hit him?”
“He sat there makin’ a fool of me.” The murderer gave a vicious
glance at the inert figure on the floor. “I want something more than
his puzzle-talk. He asked for it.”
He backed to the table where the decanter stood, and drank a
tumbler half-filled with raw spirit.
“We’re both in this, Jimmy,” he said, still keeping his man covered.
“You can put down your hands; no monkey tricks. Give me your
pistol.”
Jimmy slipped the weapon from his pocket, and handed it butt
foremost to the man. Then Massey bent over the fallen man and
searched his pockets.
“Here are the keys. You stay here,” said Massey, and went out,
closing the door after him.
Jimmy heard the grate of the key, and knew he was a prisoner. He
bent over the old man. He lay motionless. Jimmy tried the pulse, and
felt a faint flutter. Through the clenched teeth he forced a little
whisky, and after a minute the old man’s eyes opened.
“Jimmy!” he whispered; then remembering, “Where’s Massey?” he
asked.
There was no need to inquire the whereabouts of Massey. His
blundering footfalls sounded in the room above.
“Lookin’ for money?” gasped the old man, and something like a smile
crossed his face. “Safe’s up there,” he whispered, and smiled again.
“Got the keys?”
Jimmy nodded.
The old man’s eyes wandered round the room till they rested on
what looked like a switchboard.
“See that handle marked ‘seven’?” he whispered.
Jimmy nodded again.
“Pull it down, Jimmy boy.” His voice was growing fainter. “This is a
new one that I read in a book. Pull it down.”
“Why?”
“Do as I tell you,” the lips motioned, and Jimmy walked across the
room and pulled over the insulated lever.
As he did there was a heavy thud overhead that shook the room,
and then silence.
“What’s that?” he asked sharply.
The dying man smiled.
“That’s Massey!” said the lips.

Half an hour later Jimmy left the house with a soiled slip of paper in
his waistcoat pocket, on which was written the most precious verse
of doggerel that the world has known.
And the discovery of the two dead men in the upper chambers the
next morning afforded the evening press the sensation of the year.
CHAPTER III
ANGEL ESQUIRE

Nobody quite knows how Angel Esquire came to occupy the


position he does at Scotland Yard. On his appointment, “An Officer
of Twenty Years’ Standing” wrote to the Police Review and
characterized the whole thing as “a job.” Probably it was. For Angel
Esquire had been many things in his short but useful career, but
never a policeman. He had been a big game shot, a special
correspondent, a “scratch” magistrate, and his nearest approach to
occupying a responsible position in any police force in the world was
when he was appointed a J.P. of Rhodesia, and, serving on the Tuli
Commission, he hanged M’Linchwe and six of that black
desperado’s companions.
His circle of acquaintances extended to the suburbs of London, and
the suburbanites, who love you to make their flesh creep, would sit in
shivering but pleasurable horror whilst Angel Esquire elaborated the
story of the execution.
In Mayfair Angel Esquire was best known as a successful mediator.
“Who is that old-looking young man with the wicked eye?” asked the
Dowager Duchess of Hoeburn; and her vis-à-vis at the Honorable
Mrs. Carter-Walker’s “sit-down tea”—it was in the days when Mayfair
was aping suburbia—put up his altogether unnecessary eyeglass.
“Oh, that’s Angel Esquire!” he said carelessly.
“What is he?” asked the Duchess.
“A policeman.”
“India?”
“Oh, no, Scotland Yard.”
“Good Heavens!” said Her Grace in a shocked voice. “How very
dreadful! What is he doing? Watching the guests, or keeping a
friendly eye on the Carter woman’s spoons?”
The young man guffawed.
“Don’t despise old Angel, Duchess,” he said. “He’s a man to know.
Great fellow for putting things right. If you have a row with your
governor, or get into the hands of—er—undesirables, or generally, if
you’re in a mess of any kind, Angel’s the chap to pull you out.”
Her Grace surveyed the admirable man with a new interest.
Angel Esquire, with a cup of tea in one hand and a thin grass
sandwich in the other, was the center of a group of men, including
the husband of the hostess. He was talking with some animation.
“I held three aces pat, and opened the pot light to let ’em in. Young
Saville raised the opening to a tenner, and the dealer went ten better.
George Manfred, who had passed, came in for a pony, and took one
card. I took two, and drew another ace. Saville took one, and the
dealer stood pat. I thought it was my money, and bet a pony. Saville
raised it to fifty, the dealer made it a hundred, and George Manfred
doubled the bet. It was up to me. I had four aces; I put Saville with a
‘full,’ and the dealer with a ‘flush.’ I had the beating of that lot; but
what about Manfred? Manfred is a feller with all the sense going. He
knew what the others had. If he bet, he had the goods, so I chucked
my four aces into the discard. George had a straight flush.”
A chorus of approval came from the group.
If “An Officer of Twenty Years’ Standing” had been a listener, he
might well have been further strengthened in his opinion that of all
persons Mr. Angel was least fitted to fill the responsible position he
did.
If the truth be told, nobody quite knew exactly what position Angel
did hold. If you turn into New Scotland Yard and ask the janitor at the
door for Mr. Christopher Angel—Angel Esquire by the way was a
nickname affixed by a pert little girl—the constable, having satisfied
himself as to your bona-fides, would take you up a flight of stairs and
hand you over to yet another officer, who would conduct you through
innumerable swing doors, and along uncounted corridors till he
stopped before a portal inscribed “647.” Within, you would find Angel
Esquire sitting at his desk, doing nothing, with the aid of a Sporting
Life and a small weekly guide to the Turf.
Once Mr. Commissioner himself walked into the room unannounced,
and found Angel so immersed in an elaborate calculation, with big
sheets of paper closely filled with figures, and open books on either
hand, that he did not hear his visitor.
“What is the problem?” asked Mr. Commissioner, and Angel looked
up with his sweetest smile, and recognizing his visitor, rose.
“What’s the problem?” asked Mr. Commissioner again.
“A serious flaw, sir,” said Angel, with all gravity. “Here’s Mimosa
handicapped at seven stone nine in the Friary Nursery, when,
according to my calculations, she can give the field a stone, and beat
any one of ’em.”
The Commissioner gasped.
“My dear fellow,” he expostulated, “I thought you were working on
the Lagos Bank business.”
Angel had a far-away look in his eyes when he answered—
“Oh, that is all finished. Old Carby was poisoned by a man named—
forget his name now, but he was a Monrovian. I wired the Lagos
police, and we caught the chap this morning at Liverpool—took him
off an Elder, Dempster boat.”
The Police Commissioner beamed.
“My congratulations, Angel. By Jove, I thought we shouldn’t have a
chance of helping the people in Africa. Is there a white man in it?”
“We don’t know,” said Angel absently; his eye was wandering up and
down a column of figures on the paper before him.
“I am inclined to fancy there is—man named Connor, who used to be
a croupier or something to old Reale.”
He frowned at the paper, and picking up a pencil from the desk,
made a rapid little calculation. “Seven stone thirteen,” he muttered.
The Commissioner tapped the table impatiently. He had sunk into a
seat opposite Angel.
“My dear man, who is old Reale? You forget that you are our tame
foreign specialist. Lord, Angel, if you heard half the horrid things that
people say about your appointment you would die of shame!”
Angel pushed aside the papers with a little laugh.
“I’m beyond shame,” he said lightheartedly; “and, besides, I’ve
heard. You were asking about Reale. Reale is a character. For
twenty years proprietor of one of the most delightful gambling plants
in Egypt, Rome—goodness knows where. Education—none.
Hobbies—invention. That’s the ‘bee in his bonnet’—invention. If he’s
got another, it is the common or garden puzzle. Pigs in clover,
missing words, all the fake competitions that cheap little papers run
—he goes in for them all. Lives at 43 Terrington Square.”
“Where?” The Commissioner’s eyebrows rose. “Reale? 43 Terrington
Square? Why, of course.” He looked at Angel queerly. “You know all
about Reale?”
Angel shrugged his shoulders.
“As much as anybody knows,” he said.
The Commissioner nodded.
“Well, take a cab and get down at once to 43 Terrington Square.
Your old Reale was murdered last night.”
It was peculiar of Angel Esquire that nothing surprised him. He
received the most tremendous tidings with polite interest, and now
he merely said, “Dear me!” Later, as a swift hansom carried him
along Whitehall he permitted himself to be “blessed.”
Outside No. 43 Terrington Square a small crowd of morbid
sightseers stood in gloomy anticipation of some gruesome
experience or other. A policeman admitted him, and the local
inspector stopped in his interrogation of a white-faced butler to bid
him a curt “Good morning.”
Angel’s preliminary inspection did not take any time. He saw the
bodies, which had not yet been removed. He examined the pockets
of both men, and ran his eye through the scattered papers on the
floor of the room in which the tragedy had occurred. Then he came
back to the big drawing-room and saw the inspector, who was sitting
at a table writing his report.
“The chap on the top floor committed the murder, of course,” said
Angel.
“I know that,” said Inspector Boyden brusquely.
“And was electrocuted by a current passing through the handle of
the safe.”
“I gathered that,” the inspector replied as before, and went on with
his work.
“The murderer’s name is Massey,” continued Angel patiently
—“George Charles Massey.”
The inspector turned in his seat with a sarcastic smile.
“I also,” he said pointedly, “have seen the envelopes addressed in
that name, which were found in his pocket.”
Angel’s face was preternaturally solemn as he continued—
“The third man I am not so sure about.”
The inspector looked up suspiciously.
“Third man—which third man?”
Well-simulated astonishment sent Angel’s eyebrows to the shape of
inverted V’s.
“There was another man in it. Didn’t you know that, Mr. Inspector?”
“I have found no evidence of the presence of a third party,” he said
stiffly; “but I have not yet concluded my investigations.”
“Good!” said Angel cheerfully. “When you have, you will find the ends
of three cigarettes—two in the room where the old man was killed,
and one in the safe room. They are marked ‘Al Kam,’ and are a fairly
expensive variety of Egyptian cigarettes. Massey smoked cigars; old
Reale did not smoke at all. The question is”—he went on speaking
aloud to himself, and ignoring the perplexed police official—“was it
Connor or was it Jimmy?”
The inspector struggled with a desire to satisfy his curiosity at the
expense of his dignity, and resolved to maintain an attitude of
superior incredulity. He turned back to his work.
“It would be jolly difficult to implicate either of them,” Angel went on
reflectively, addressing the back of the inspector. “They would
produce fifty unimpeachable alibis, and bring an action for wrongful
arrest in addition,” he added artfully.
“They can’t do that,” said the inspector gruffly.
“Can’t they?” asked the innocent Angel. “Well, at any rate, it’s not
advisable to arrest them. Jimmy would——”
Inspector Boyden swung round in his chair.
“I don’t know whether you’re ‘pulling my leg,’ Mr. Angel. You are
perhaps unused to the procedure in criminal cases in London, and I
must now inform you that at present I am in charge of the case, and
must request that if you have any information bearing upon this
crime to give it to me at once.”
“With all the pleasure in life,” said Angel heartily. “In the first place,
Jimmy——”
“Full name, please.” The inspector dipped his pen in ink.
“Haven’t the slightest idea,” said the other carelessly. “Everybody
knows Jimmy. He was old Reale’s most successful decoy duck. Had
the presence and the plumage and looked alive, so that all the other
little ducks used to come flying down and settle about him, and long
before they could discover that the beautiful bird that attracted them
was only painted wood and feathers, ‘Bang! bang!’ went old Reale’s
double-barrel, and roast duck was on the menu for days on end.”
Inspector Boyden threw down his pen with a grunt.
“I’m afraid,” he said in despair, “that I cannot include your parable in
my report. When you have any definite information to give, I shall be
pleased to receive it.”
Later, at Scotland Yard, Angel interviewed the Commissioner.
“What sort of a man is Boyden to work with?” asked Mr.
Commissioner.
“A most excellent chap—good-natured, obliging, and as zealous as
the best of ’em,” said Angel, which was his way.
“I shall leave him in charge of the case,” said the Chief.
“You couldn’t do better,” said Angel decisively.
Then he went home to his flat in Jermyn Street to dress for dinner.
It was an immaculate Angel Esquire who pushed through the plate-
glass, turn-table door of the Heinz, and, walking into the magnificent
old rose dining-room, selected a table near a window looking out on
to Piccadilly.
The other occupant of the table looked up and nodded.
“Hullo, Angel!” he said easily.
“Hullo, Jimmy!” greeted the unconventional detective.
He took up the card and chose his dishes with elaborate care. A half-
bottle of Beaujolais completed his order.
“The ridiculous thing is that one has got to pay 7s. 6d. for a small
bottle of wine that any respectable grocer will sell you for tenpence
ha’-penny net.”
“You must pay for the magnificence,” said the other, quietly amused.
Then, after the briefest pause, “What do you want?”
“Not you, Jimmy,” said the amiable Angel, “though my young friend,
Boyden, Inspector of Police, and a Past Chief Templar to boot, will
be looking for you shortly.”
Jimmy carefully chose a toothpick and stripped it of its tissue
covering.
“Of course,” he said quietly, “I wasn’t in it—the killing, I mean. I was
there.”
“I know all about that,” said Angel; “saw your foolish cigarettes. I
didn’t think you had any hand in the killing. You are a property
criminal, not a personal criminal.”
“By which I gather you convey the nice distinction as between crimes
against property and crimes against the person,” said the other.
“Exactly.”
A pause.
“Well?” said Jimmy.
“What I want to see you about is the verse,” said Angel, stirring his
soup.
Jimmy laughed aloud.
“What a clever little devil you are, Angel,” he said admiringly; “and
not so little either, in inches or devilishness.”
He relapsed into silence, and the wrinkled forehead was eloquent.
“Think hard,” taunted Angel.
“I’m thinking,” said Jimmy slowly. “I used a pencil, as there was no
blotting paper. I only made one copy, just as the old man dictated it,
and——”
“You used a block,” said Angel obligingly, “and only tore off the top
sheet. And you pressed rather heavily on that, so that the next sheet
bore a legible impression.”
Jimmy looked annoyed.
“What an ass I am!” he said, and was again silent.
“The verse?” said Angel. “Can you make head or tail of it?”
“No”—Jimmy shook his head—“can you?”
“Not a blessed thing,” Angel frankly confessed.
Through the next three courses neither man spoke. When coffee had
been placed on the table, Jimmy broke the silence—
“You need not worry about the verse. I have only stolen a march of a
few days. Then Connor will have it; and some girl or other will have
it. Massey would have had it too.” He smiled grimly.
“What is it all about?”
Jimmy looked at his questioner with some suspicion.
“Don’t you know?” he demanded.
“Haven’t got the slightest notion. That is why I came to see you.”
“Curious!” mused Jimmy. “I thought of looking you up for the very
same purpose. We shall know in a day or two,” he went on,
beckoning the waiter. “The old man said it was all in the will. He just
told me the verse before he died. The ruling passion, don’t you know.
‘Learn it by heart, Jimmy,’ he croaked; ‘it’s two millions for you if you
guess it’—and that’s how he died. My bill, waiter. Which way do you
go?” he asked as they turned into Piccadilly.
“To the ‘Plait’ for an hour,” said Angel.
“Business?”
“Partly; I’m looking for a man who might be there.”
They crossed Piccadilly, and entered a side turning. The second on
the left and the first on the right brought them opposite a brightly-lit
hotel. From within came the sound of violins. At the little tables with
which the spacious bar-room was set about sat laughing women and
young men in evening dress. A haze of cigarette smoke clouded the
atmosphere, and the music made itself heard above a babel of
laughter and talk. They found a corner, and seated themselves.
“You seem to be fairly well known here,” said Jimmy.
“Yes,” replied Angel ruefully, “a jolly sight too well known. You’re not
quite a stranger, Jimmy,” he added.

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