Third Edition

Third Edition
Physics in Biology and Medicine
Complementary Science Series
ACADEMIC PRESS
2008—To be published
Physics in the Arts (Full Edition)
P.U.P.A. Gilbert Willy Haeberli
2003–2007
Physics in Biology and Medicine, Third Edition
Paul Davidovits
Crystallography Made Crystal Clear, Third Edition
Gale Rhodes
Fusion
Garry M. McCracken Peter Stott
Introduction to Quantum Mechanics
Sy M. Blinder
Fundamentals of Quantum Chemistry, Second Edition
James E. House
2000–2002
Introduction to Relativity
John B. Kogut
Earth Magnetism: A Guided Tour through Magnetic Fields
Wallace H. Campbell
The Physical Basis of Chemistry, Second Edition
Warren S. Warren
www.books.elsevier.com
Third Edition
Physics in Biology
and Medicine
Paul Davidovits
AMSTERDAM • BOSTON • HEIDELBERG • LONDON
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Davidovits, Paul.
Physics in biology and medicine / Paul Davidovits. – 3rd ed.
p. cm. – (Complementary science series)
Includes bibliographical references and index.
ISBN-13: 978-0-12-369411-9 (pbk. : alk. paper) 1. Biophysics. 2. Medical physics. I. Title.
QH505.D36 2008
571.4—dc22
2007021167
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ISBN: 978-0-12-369411-9
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Contents
Preface xiii
Abbreviations xvii
1 Static Forces 1
1.1 Equilibrium and Stability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.2 Equilibrium Considerations for the Human Body. . . . . . . . . . . 3
1.3 Stability of the Human Body under the Action of an
External Force. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.4 Skeletal Muscles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.5 Levers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
1.6 The Elbow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
1.7 The Hip. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
1.7.1 Limping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
1.8 The Back . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
1.9 Standing Tip-Toe on One Foot . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
1.10 Dynamic Aspects of Posture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
2 Friction 23
2.1 Standing at an Incline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
2.2 Friction at the Hip Joint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
v
vi Contents
2.3 Spine Fin of a Catfish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
3 Translational Motion 30
3.1 Vertical Jump. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
3.2 Effect of Gravity on the Vertical Jump . . . . . . . . . . . . . . . . . . . . 35
3.3 Running High Jump. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
3.4 Range of a Projectile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
3.5 Standing Broad Jump . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
3.6 Running Broad Jump (Long Jump) . . . . . . . . . . . . . . . . . . . . . . . 39
3.7 Motion through Air . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
3.8 Energy Consumed in Physical Activity. . . . . . . . . . . . . . . . . . . . 42
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
4 Angular Motion 45
4.1 Forces on a Curved Path . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
4.2 A Runner on a Curved Track. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
4.3 Pendulum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
4.4 Walking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
4.5 Physical Pendulum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
4.6 Speed of Walking and Running. . . . . . . . . . . . . . . . . . . . . . . . . . . 52
4.7 Energy Expended in Running . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
4.8 Alternate Perspectives on Walking And Running . . . . . . . . . . . 56
4.9 Carrying Loads . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
5 Elasticity and Strength of Materials 61
5.1 Longitudinal Stretch and Compression . . . . . . . . . . . . . . . . . . . . 61
5.2 A Spring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
5.3 Bone Fracture: Energy Considerations . . . . . . . . . . . . . . . . . . . . 64
5.4 Impulsive Forces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
5.5 Fracture Due to a Fall: Impulsive Force Considerations . . . . . 67
5.6 Airbags: Inflating Collision Protection Devices . . . . . . . . . . . . 68
5.7 Whiplash Injury . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
5.8 Falling from Great Height . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
5.9 Osteoarthritis and Exercise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Contents vii
6 Insect Flight 73
6.1 Hovering Flight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
6.2 Insect Wing Muscles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
6.3 Power Required for Hovering . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
6.4 Kinetic Energy of Wings in Flight . . . . . . . . . . . . . . . . . . . . . . . . 78
6.5 Elasticity of Wings. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
7 Fluids 82
7.1 Force and Pressure in a Fluid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
7.2 Pascal’s Principle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
7.3 Hydrostatic Skeleton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
7.4 Archimedes’ Principle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
7.5 Power Required to Remain Afloat . . . . . . . . . . . . . . . . . . . . . . . . 87
7.6 Buoyancy of Fish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
7.7 Surface Tension . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
7.8 Soil Water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
7.9 Insect Locomotion on Water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
7.10 Contraction of Muscles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
7.11 Surfactants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
8 The Motion of Fluids 101
8.1 Bernoulli’s Equation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
8.2 Viscosity and Poiseuille’s Law . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
8.3 Turbulent Flow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
8.4 Circulation of the Blood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
8.5 Blood Pressure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
8.6 Control of Blood Flow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
8.7 Energetics of Blood Flow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
8.8 Turbulence in the Blood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
8.9 Arteriosclerosis and Blood Flow. . . . . . . . . . . . . . . . . . . . . . . . . . 111
8.10 Power Produced by the Heart . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
8.11 Measurement of Blood Pressure . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
viii Contents
9 Heat and Kinetic Theory 116
9.1 Heat and Hotness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
9.2 Kinetic Theory of Matter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
9.3 Definitions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
9.3.1 Unit of Heat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
9.3.2 Specific Heat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
9.3.3 Latent Heats. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
9.4 Transfer of Heat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
9.4.1 Conduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
9.4.2 Convection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
9.4.3 Radiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
9.4.4 Diffusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123
9.5 Transport of Molecules by Diffusion . . . . . . . . . . . . . . . . . . . . . . 126
9.6 Diffusion through Membranes. . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
9.7 The Respiratory System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
9.8 Surfactants and Breathing. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
9.9 Diffusion and Contact Lenses . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
10 Thermodynamics 135
10.1 First Law of Thermodynamics . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
10.2 Second Law of Thermodynamics . . . . . . . . . . . . . . . . . . . . . . . . . 137
10.3 Difference between Heat and Other Forms of
Energy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
10.4 Thermodynamics of Living Systems . . . . . . . . . . . . . . . . . . . . . . 140
10.5 Information and the Second Law . . . . . . . . . . . . . . . . . . . . . . . . . 143
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
11 Heat and Life 145
11.1 Energy Requirements of People . . . . . . . . . . . . . . . . . . . . . . . . . . 146
11.2 Energy from Food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
11.3 Regulation of Body Temperature . . . . . . . . . . . . . . . . . . . . . . . . . 149
11.4 Control of Skin Temperature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
11.5 Convection. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
11.6 Radiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
11.7 Radiative Heating by the Sun . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Contents ix
11.8 Evaporation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155
11.9 Resistance to Cold . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156
11.10 Heat and Soil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
12 Waves and Sound 162
12.1 Properties of Sound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
12.2 Some Properties of Waves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165
12.2.1 Reflection and Refraction . . . . . . . . . . . . . . . . . . . . . . . . . . 165
12.2.2 Interference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166
12.2.3 Diffraction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
12.3 Hearing and the Ear . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
12.3.1 Performance of the Ear . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
12.3.2 Frequency and Pitch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172
12.3.3 Intensity and Loudness . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
12.4 Bats and Echoes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
12.5 Sounds Produced by Animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
12.6 Acoustic Traps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
12.7 Clinical Uses of Sound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
12.8 Ultrasonic Waves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
13 Electricity 180
13.1 The Nervous System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
13.1.1 The Neuron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
13.1.2 Electrical Potentials in the Axon . . . . . . . . . . . . . . . . . . . . 183
13.1.3 Action Potential. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
13.1.4 Axon as an Electric Cable. . . . . . . . . . . . . . . . . . . . . . . . . . 186
13.1.5 Propagation of the Action Potential . . . . . . . . . . . . . . . . . . 188
13.1.6 An Analysis of the Axon Circuit . . . . . . . . . . . . . . . . . . . . 190
13.1.7 Synaptic Transmission . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
13.1.8 Action Potentials in Muscles . . . . . . . . . . . . . . . . . . . . . . . 194
13.1.9 Surface Potentials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
13.2 Electricity in Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
13.3 Electricity in the Bone. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
x Contents
13.4 Electric Fish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
14 Electrical Technology 200
14.1 Electrical Technology in Biological Research . . . . . . . . . . . . . . 200
14.2 Diagnostic Equipment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202
14.2.1 The Electrocardiograph. . . . . . . . . . . . . . . . . . . . . . . . . . . . 202
14.2.2 The Electroencephalograph . . . . . . . . . . . . . . . . . . . . . . . . 203
14.3 Physiological Effects of Electricity . . . . . . . . . . . . . . . . . . . . . . . 204
14.4 Control Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206
14.5 Feedback . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
14.6 Sensory Aids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
14.6.1 Hearing Aids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
14.6.2 Cochlear Implant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
15 Optics 214
15.1 Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
15.2 Nature of Light . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
15.3 Structure of the Eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
15.4 Accommodation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
15.5 Eye and the Camera. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
15.5.1 Aperture and Depth of Field . . . . . . . . . . . . . . . . . . . . . . . . 218
15.6 Lens System of the Eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
15.7 Reduced Eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
15.8 Retina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
15.9 Resolving Power of the Eye. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
15.10 Threshold of Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
15.11 Vision and the Nervous System . . . . . . . . . . . . . . . . . . . . . . . . . . 226
15.12 Defects in Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
15.13 Lens for Myopia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
15.14 Lens for Presbyopia and Hyperopia . . . . . . . . . . . . . . . . . . . . . . . 229
15.15 Extension of Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
15.15.1 Telescope . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
15.15.2 Microscope . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
15.15.3 Confocal Microscopy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
Contents xi
15.15.4 Fiber Optics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
16 Atomic Physics 239
16.1 The Atom . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
16.2 Spectroscopy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244
16.3 Quantum Mechanics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246
16.4 Electron Microscope . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
16.5 X-rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
16.6 X-ray Computerized Tomography . . . . . . . . . . . . . . . . . . . . . . . . 250
16.7 Lasers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252
16.7.1 Lasers Surgery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255
17 Nuclear Physics 256
17.1 The Nucleus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256
17.2 Magnetic Resonance Imaging . . . . . . . . . . . . . . . . . . . . . . . . . . . . 257
17.2.1 Nuclear Magnetic Resonance . . . . . . . . . . . . . . . . . . . . . . . 258
17.2.2 Imaging with NMR . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262
17.2.3 Functional Magnetic Resonance Imaging
(fMRI) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265
17.3 Radiation Therapy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266
17.4 Food Preservation by Radiation . . . . . . . . . . . . . . . . . . . . . . . . . . 267
17.5 Isotopic Tracers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268
17.6 Laws of Physics and Life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269
Exercises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
Appendix A: Basic Concepts in Mechanics 272
Appendix B: Review of Electricity 287
Appendix C: Review of Optics 293
Bibliography 302
Answers to Numerical Exercises 310
Index 314
Companion Web Site Information
Instructor support materials for Physics in Biology and Medicine,
Third Edition, can be found at:
www.textbooks.elsevier.com/9780123694119/
Preface
Until the mid 1800s it was not clear to what extent the laws of physics and
chemistry, which were formulated from the observed behavior of inanimate
matter, could be applied to living matter. It was certainly evident that on the
large scale the laws were applicable. Animals are clearly subject to the same
laws of motion as inanimate objects. The question of applicability arose on
a more basic level. Living organisms are very complex. Even a virus, which
is one of the simplest biological organisms, consists of millions of interacting
atoms. A cell, which is the basic building block of tissue, contains on the average
1014 atoms. Living organisms exhibit properties not found in inanimate
objects. They grow, reproduce, and decay. These phenomena are so different
from the predictable properties of inanimate matter that many scientists in
the early 19th century believed that different laws governed the structure and
organization molecules in living matter. Even the physical origin of organic
molecules was in question. These molecules tend to be larger and more complex
than molecules obtained from inorganic sources. It was thought that the
large molecules found in living matter could be produced only by living organisms
through a “vital force” that could not be explained by the existing laws of
physics. This concept was disproved in 1828 when Friedrich W�ohler synthesized
an organic substance, urea, from inorganic chemicals. Soon thereafter
many other organic molecules were synthesized without the intervention of
biological organisms. Today most scientists believe that there is no special
vital force residing in organic substances. Living organisms are governed by
the laws of physics on all levels.
xiii
xiv Preface
Much of the biological research during the past hundred years has been
directed toward understanding living systems in terms of basic physical laws.
This effort has yielded some significant successes. The atomic structure of
many complex biological molecules has now been determined, and the role of
these molecules within living systems has been described. It is now possible to
explain the functioning of cells and many of their interactions with each other.
Yet the work is far from complete. Even when the structure of a complex
molecule is known, it is not possible at present to predict its function from its
atomic structure. The mechanisms of cell nourishment, growth, reproduction,
and communication are still understood only qualitatively. Many of the basic
questions in biology remain unanswered. However, biological research has
so far not revealed any areas where physical laws do not apply. The amazing
properties of life seem to be achieved by the enormously complex organization
in living systems.
The aim of this book is to relate some of the concepts in physics to living
systems. In general, the text follows topics found in basic college physics
texts. The discussion is organized into the following areas: solid mechanics,
fluid mechanics, thermodynamics, sound, electricity, optics, and atomic and
nuclear physics.
Each chapter contains a brief review of the background physics, but most
of the text is devoted to the applications of physics to biology and medicine.
No previous knowledge of biology is assumed. The biological systems to
be discussed are described in as much detail as is necessary for the physical
analysis. Whenever possible, the analysis is quantitative, requiring only basic
algebra and trigonometry.
Many biological systems can be analyzed quantitatively. A few examples
will illustrate the approach. Under the topic of mechanics we calculate the
forces exerted by muscles. We examine the maximum impact a body can
sustain without injury. We calculate the height to which a person can jump,
and we discuss the effect of an animal’s size on the speed at which it can run.
In our study of fluids we examine quantitatively the circulation of blood in
the body. The theory of fluids allows us also to calculate the role of diffusion
in the functioning of cells and the effect of surface tension on the growth of
plants in soil. Using the principles of electricity, we analyze quantitatively
the conduction of impulses along the nervous system. Each section contains
problems that explore and expand some of the concepts.
There are, of course, severe limits on the quantitative application of physics
to biological systems. These limitations are discussed.
Many of the advances in the life sciences have been greatly aided by the
application of the techniques of physics and engineering to the study of living
systems. Some of these techniques are examined in the appropriate sections
of the book.
Preface xv
This new edition has been updated and includes a discussion of information
theory and descriptions of CT scan, endoscopy, MRI and fMRI imaging,
techniques that were not available at the writing of the earlier editions.
A word about units. Most physics and chemistry textbooks now use the
MKS International System of units (SI). In practice, however, a variety of
units continues to be in use. For example, in the SI system, pressure is
expressed in units of pascal (kg/m2). Both in common use and in the scientific
literature one often finds pressure also expressed in units of dynes/cm2,
Torr (mm Hg), psi, and atm. In this book I have used mostly SI units. However,
other units have also been used when common usage so dictated. In
those cases conversion factors have been provided either within the text or in
a compilation at the end of Appendix A .
In the first edition of this book I expressed my thanks to W. Chameides,
M. D. Egger, L. K. Stark, and J. Taplitz for their help and encouragement.
In the second edition I thanked Prof. R. K. Hobbie and David Cinabro for
their careful reading of the manuscript and helpful suggestions. In this third
edition I want to express my appreciation for the encouragement and competent
direction of Tom Singer and Jason Malley editors at Elsevier/Academic
Press and for the help of Sarah Hajduk and Ramesh Gurusubramanian in the
production of this edition.
Paul Davidovits
Chestnut Hill, Massachusetts
May 2007
This page intentionally left blank
Abbreviations
˚A angstrom
av average
atm atmosphere
A ampere
cal calorie (gram calorie)
Cal Calorie (kilo calorie)
C coulomb
CT computerized tomography
cos cosine
cps cycles per second
cm2 square centimeters
cm centimeter
c.g. center of gravity
deg degree
dB decibel
diam diameter
dyn dyne
dyn/cm2 dynes per square centimeter
ft foot
ft/sec feet per second
F farad
F/m farad/meters
g gram
h hour
xvii
xviii Abbreviations
Hz hertz (cps)
J joule
km kilometer
km/h kilometers per hour
kg kilogram
KE kinetic energy
in inch
kph kilometers per hour
lim limit
lb pound
liter/min liters per minute
μ micron
μA microampere
μV microvolt
μV/m microvolt per meter
mV millivolt
ms millisecond
m meter
m/sec meters per second
min minute
mph miles per hour
max maximum
mA milliampere
MRI magnetic resonance imaging
N newton
N-m newton meters
NMR nuclear magnetic resonance
Ω ohm
PE potential energy
psi pounds per sq. in.
rad radian
sin sine
sec second
SMT Soil moisture tension
tan tangent
V volt
W watt
Chapter 1
Static Forces
Mechanics is the branch of physics concerned with the effect of forces on the
motion of bodies. It was the first branch of physics that was applied successfully
to living systems, primarily to understanding the principles governing the
movement of animals. Our present concepts of mechanics were formulated
by Isaac Newton, whose major work on mechanics, Principia Mathematica,
was published in 1687. The study of mechanics, however, began much earlier.
It can be traced to the Greek philosophers of the fourth century B.C. The
early Greeks, who were interested in both science and athletics, were also
the first to apply physical principles to animal movements. Aristotle wrote,
“The animal that moves makes its change of position by pressing against
that which is beneath it. ... Runners run faster if they swing their arms for
in extension of the arms there is a kind of leaning upon the hands and the
wrist.” Although some of the concepts proposed by the Greek philosophers
were wrong, their search for general principles in nature marked the beginning
of scientific thought.
After the decline of ancient Greece, the pursuit of all scientific work
entered a period of lull that lasted until the Renaissance brought about
a resurgence in many activities including science. During this period of
revival, Leonardo da Vinci (1452–1519) made detailed observations of animal
motions and muscle functions. Since da Vinci, hundreds of people have
contributed to our understanding of animal motion in terms of mechanical
principles. Their studies have been aided by improved analytic techniques
and the development of instruments such as the photographic camera and
electronic timers. Today the study of human motion is part of the disciplines
1
2 Chapter 1 Static Forces
of kinesiology, which studies human motion primarily as applied to athletic
activities, and biomechanics, a broader area that is concerned not only with
muscle movement but also with the physical behavior of bones and organs
such as the lungs and the heart. The development of prosthetic devices such
as artificial limbs and mechanical hearts is an active area of biomechanical
research.
Mechanics, like every other subject in science, starts with a certain number
of basic concepts and then supplies the rules by which they are interrelated.
Appendix A summarizes the basic concepts in mechanics, providing a review
rather than a thorough treatment of the subject. We will now begin our discussion
of mechanics by examining static forces that act on the human body.
We will first discuss stability and equilibrium of the human body, and then we
will calculate the forces exerted by the skeletal muscles on various parts of
the body.
1.1 Equilibrium and Stability
The Earth exerts an attractive force on the mass of an object; in fact, every
small element of mass in the object is attracted by the Earth. The sum of
these forces is the total weight of the body. This weight can be considered
a force acting through a single point called the center of mass or center of
gravity. As pointed out in Appendix A, a body is in static equilibrium if the
vectorial sum of both the forces and the torques acting on the body is zero. If a
body is unsupported, the force of gravity accelerates it, and the body is not in
equilibrium. In order that a body be in stable equilibrium, it must be properly
supported.
The position of the center of mass with respect to the base of support determines
whether the body is stable or not. A body is in stable equilibrium under
the action of gravity if its center of mass is directly over its base of support
(Fig. 1.1). Under this condition, the reaction force at the base of support cancels
the force of gravity and the torque produced by it. If the center of mass
is outside the base, the torque produced by the weight tends to topple the
body (Fig. 1.1c).
The wider the base on which the body rests, the more stable it is; that is, the
more difficult it is to topple it. If the wide-based body in Fig. 1.1a is displaced
as shown in Fig. 1.2a, the torque produced by its weight tends to restore it to
its original position (Fr shown is the reaction force exerted by the surface on
the body). The same amount of angular displacement of a narrow-based body
results in a torque that will topple it (Fig. 1.2b). Similar considerations show
that a body is more stable if its center of gravity is closer to its base.
Section 1.2 Equilibrium Considerations for the Human Body 3
FIGURE 1.1 Stability of bodies.
FIGURE 1.2 (a) Torque produced by the weight will restore the body to its original
position. (b) Torque produced by the weight will topple the body.
1.2 Equilibrium Considerations for the Human Body
The center of gravity (c.g.) of an erect person with arms at the side is at
approximately 56% of the person’s height measured from the soles of the feet
(Fig. 1.3). The center of gravity shifts as the person moves and bends. The
act of balancing requires maintenance of the center of gravity above the feet.
A person falls when his center of gravity is displaced beyond the position of
the feet.
When carrying an uneven load, the body tends to compensate by bending
and extending the limbs so as to shift the center of gravity back over the
feet. For example, when a person carries a weight in one arm, the other arm
4 Chapter 1 Static Forces
FIGURE 1.3 Center of gravity for a person.
swings away from the body and the torso bends away from the load (Fig. 1.4).
This tendency of the body to compensate for uneven weight distribution often
causes problems for people who have lost an arm, as the continuous compensatory
bending of the torso can result in a permanent distortion of the spine. It
is often recommended that amputees wear an artificial arm, even if they cannot
use it, to restore balanced weight distribution.
1.3 Stability of the Human Body under the Action of an
External Force
The body may of course be subject to forces other than the downward force
of weight. Let us calculate the magnitude of the force applied to the shoulder
that will topple a person standing at rigid attention. The assumed dimensions
of the person are as shown in Fig. 1.5. In the absence of the force, the person
is in stable equilibrium because his center of mass is above his feet, which are
Section 1.3 Stability of the Human Body under the Action of an External Force 5
FIGURE 1.4 A person carrying a weight.
the base of support. The applied force Fa tends to topple the body. When the
person topples, he will do so by pivoting around point A—assuming that he
does not slide. The counterclockwise torque Ta about this point produced by
the applied force is
Ta Fa � 1.5 m (1.1)
The opposite restoring torque Tw due to the person’s weight is
Tw W � 0.1 m (1.2)
Assuming that the mass m of the person is 70 kg, his weight W is
W mg 70 � 9.8 686 newton (N) (1.3)
(Here g is the gravitational acceleration, which has the magnitude 9.8 m/sec2.)
The restoring torque produced by the weight is therefore 68.6 newton-meter
6 Chapter 1 Static Forces
FIGURE 1.5 A force applied to an erect person.
(N-m). The person is on the verge of toppling when the magnitudes of these
two torques are just equal; that is, Ta Tw or
Fa � 1.5 m 68.6 N-m (1.4)
Therefore, the force required to topple an erect person is
Fa
68.6
1.5
45.7 N (10.3 lb) (1.5)
Actually, a person can withstand a much greater sideways force without
losing balance by bending the torso in the direction opposite to the applied
force (Fig. 1.6). This shifts the center of gravity away from the pivot point A,
increasing the restoring torque produced by the weight of the body.
Stability against a toppling force is also increased by spreading the legs,
as shown in Fig. 1.7 and discussed in Exercise 1-1.
Section 1.4 Skeletal Muscles 7
FIGURE 1.6 Compensating for a side-pushing force.
1.4 Skeletal Muscles
The skeletal muscles producing skeletal movements consist of many thousands
of parallel fibers wrapped in a flexible sheath that narrows at both ends
into tendons (Fig. 1.8). The tendons, which are made of strong tissue, grow
into the bone and attach the muscle to the bone. Most muscles taper to a single
tendon. But some muscles end in two or three tendons; these muscles are
called, respectively, biceps and triceps. Each end of the muscle is attached
to a different bone. In general, the two bones attached by muscles are free to
move with respect to each other at the joints where they contact each other.
This arrangement of muscle and bone was noted by Leonardo da Vinci,
who wrote, “The muscles always begin and end in the bones that touch one
another, and they never begin and end on the same bone. . . .” He also stated,
8 Chapter 1 Static Forces
FIGURE 1.7 Increased stability resulting from spreading the legs.
“It is the function of the muscles to pull and not to push except in the cases of
the genital member and the tongue.”
Da Vinci’s observation about the pulling by muscles is correct. When fibers
in the muscle receive an electrical stimulus from the nerve endings that are
attached to them, they contract. This results in a shortening of the muscle and a
corresponding pulling force on the two bones to which the muscle is attached.
There is a great variability in the pulling force that a given muscle can apply.
The force of contraction at any time is determined by the number of individual
fibers that are contracting within the muscle. When an individual fiber receives
an electrical stimulus, it tends to contract to its full ability. If a stronger pulling
force is required, a larger number of fibers are stimulated to contract.
Experiments have shown that the maximum force a muscle is capable of
exerting is proportional to its cross section. From measurements, it has been
estimated that a muscle can exert a force of about 7 � 106 dyn/cm2 of its area
(7 � 106 dyn/cm2 7 � 105 Pa 102 lb/in2).
Section 1.5 Levers 9
FIGURE 1.8 Drawing of a muscle.
To compute the forces exerted by muscles, the various joints in the body
can be conveniently analyzed in terms of levers. Such a representation implies
some simplifying assumptions. We will assume that the tendons are connected
to the bones at well-defined points and that the joints are frictionless.
Simplifications are often necessary to calculate the behavior of systems in
the real world. Seldom are all the properties of the system known, and even
when they are known, consideration of all the details is usually not necessary.
Calculations are most often based on a model, which is assumed to be a good
representation of the real situation.
1.5 Levers
A lever is a rigid bar free to rotate about a fixed point called the fulcrum. The
position of the fulcrum is fixed so that it is not free to move with respect to
10 Chapter 1 Static Forces
the bar. Levers are used to lift loads in an advantageous way and to transfer
movement from one point to another.
There are three classes of levers, as shown in Fig. 1.9. In a Class 1 lever,
the fulcrum is located between the applied force and the load. A crowbar is an
example of a Class 1 lever. In a Class 2 lever, the fulcrum is at one end of the
bar; the force is applied to the other end; and the load is situated in between.
A wheelbarrow is an example of a Class 2 lever. A Class 3 lever has the
fulcrum at one end and the load at the other. The force is applied between
the two ends. As we will see, many of the limb movements of animals are
performed by Class 3 levers.
It can be shown from the conditions for equilibrium (see Appendix A) that,
for all three types of levers, the force F required to balance a load of weight
W is given by
F
Wd1
d2
, (1.6)
where d1 and d2 are the lengths of the lever arms, as shown in Fig. 1.9 (see
Exercise 1-2). If d1 is less than d2, the force required to balance a load is
smaller than the load. The mechanical advantage M of the lever is defined as
M
W
F
d2
d1
. (1.7)
Depending on the distances from the fulcrum, the mechanical advantage of a
Class 1 lever can be greater or smaller than one. By placing the load close to
the fulcrum, with d1 much smaller than d2, a very large mechanical advantage
can be obtained with a Class 1 lever. In a Class 2 lever, d1 is always smaller
than d2; therefore, the mechanical advantage of a Class 2 lever is greater than
one. The situation is opposite in a Class 3 lever. Here d1 is larger than d2;
therefore, the mechanical advantage is always less than one.
FIGURE 1.9 The three classes of lever.
Section 1.6 The Elbow 11
FIGURE 1.10 Motion of the lever arms in a Class 1 lever.
A force slightly greater than what is required to balance the load will lift
it. As the point at which the force is applied moves through a distance L2, the
load moves a distance L1 (see Fig. 1.10). The relationship between L1 and
L2, (see Exercise 1-2) is given by
L1
L2
d1
d2
. (1.8)
The ratio of velocities of these two points on a moving lever is likewise
given by
v1
v2
d1
d2
. (1.9)
Here v2 is the velocity of the point where the force is applied, and v1 is the
velocity of the load. These relationships apply to all three classes of levers.
Thus, it is evident that the excursion and velocity of the load are inversely
proportional to the mechanical advantage.
1.6 The Elbow
The two most important muscles producing elbow movement are the biceps
and the triceps (Fig. 1.11). The contraction of the triceps causes an extension,
or opening, of the elbow, while contraction of the biceps closes the elbow.
In our analysis of the elbow, we will consider the action of only these two
muscles. This is a simplification, as many other muscles also play a role in
elbow movement. Some of them stabilize the joints at the shoulder as the
elbow moves, and others stabilize the elbow itself.
12 Chapter 1 Static Forces
FIGURE 1.11 The elbow.
FIGURE 1.12 (a) Weight held in hand. (b) A simplified drawing of (a).
Section 1.6 The Elbow 13
FIGURE 1.13 Lever representation of Fig. 1.12.
Figure 1.12a shows a weight W held in the hand with the elbow bent at a
100◦ angle. A simplified diagram of this arm position is shown in Fig. 1.12b.
The dimensions shown in Fig. 1.12 are reasonable for a human arm, but they
will, of course, vary from person to person. The weight pulls the arm downward.
Therefore, the muscle force acting on the lower arm must be in the up
direction. Accordingly, the prime active muscle is the biceps. The position of
the upper arm is fixed at the shoulder by the action of the shoulder muscles.
We will calculate, under the conditions of equilibrium, the pulling force Fm
exerted by the biceps muscle and the direction and magnitude of the reaction
force Fr at the fulcrum (the joint). The calculations will be performed by considering
the arm position as a Class 3 lever, as shown in Fig. 1.13. The x- and
y-axes are as shown in Fig. 1.13. The direction of the reaction force Fr shown
is a guess. The exact answer will be provided by the calculations.
In this problem we have three unknown quantities: the muscle force Fm,
the reaction force at the fulcrum Fr, and the angle, or direction, of this force
φ. The angle θ of the muscle force can be calculated from trigonometric considerations,
without recourse to the conditions of equilibrium. As is shown in
Exercise 1-3, the angle θ is 72.6◦.
For equilibrium, the sum of the x and y components of the forces must
each be zero. From these conditions we obtain
x components of the forces: Fm cos θ Fr cos φ (1.10)
y components of the forces: Fm sin θ W + Fr sin φ (1.11)
These two equations alone are not sufficient to determine the three unknown
quantities. The additional necessary equation is obtained from the torque conditions
for equilibrium. In equilibrium, the torque about any point in Fig. 1.13
must be zero. For convenience, we will choose the fulcrum as the point for
our torque balance.
14 Chapter 1 Static Forces
The torque about the fulcrum must be zero. There are two torques about
this point: a clockwise torque due to the weight and a counterclockwise torque
due to the vertical y component of the muscle force. Since the reaction force
Fr acts at the fulcrum, it does not produce a torque about this point.
Using the dimensions shown in Fig. 1.12, we obtain
4 cm � Fm sin θ 40 cm � W
or
Fm sin θ 10W (1.12)
Therefore, with θ 72.6◦, the muscle force Fm is
Fm
10 W
0.954
10.5W (1.13)
With a 14-kg (31-lb) weight in hand, the force exerted by the muscle is
Fm 10.5 � 14 � 9.8 1440 N (325 lb)
If we assume that the diameter of the biceps is 8 cm and that the muscle
can produce a 7 � 106 dyn force for each square centimeter of area, the arm
is capable of supporting a maximum of 334 N (75 lb) in the position shown in
Fig. 1.13 (see Exercise 1-4).
The solutions of Eqs. 1.10 and 1.11 provide the magnitude and direction
of the reaction force Fr. Assuming as before that the weight supported is
14 kg, these equations become
1440 � cos 72.6 Fr cos φ
1440 � sin 72.6 14 � 9.8 + Fr sin φ (1.14)
or
Fr cos φ 430 N
Fr sin φ 1240 N (1.15)
Squaring both equations, using cos2 φ + sin2 φ 1 and adding them, we
obtain
F2
r 1.74 � 106
N2
or
Fr 1320 N (298 lb) (1.16)
From Eqs. 1.14 and 1.15, the cotangent of the angle is
cot φ
430
1240
0.347 (1.17)
and
φ 70.9◦
Section 1.7 The Hip 15
Exercises 1-5, 1-6, and 1-7 present other similar aspects of biceps mechanics.
In these calculations we have omitted the weight of the arm itself, but this
effect is considered in Exercise 1-8. The forces produced by the triceps muscle
are examined in Exercise 1-9.
Our calculations show that the forces exerted on the joint and by the muscle
are large. In fact, the force exerted by the muscle is much greater than the
weight it holds up. This is the case with all the skeletal muscles in the body.
They all apply forces by means of levers that have a mechanical advantage
less than one. As mentioned earlier, this arrangement provides for greater
speed of the limbs. A small change in the length of the muscle produces a
relatively larger displacement of the limb extremities (see Exercise 1-10). It
seems that nature prefers speed to strength. In fact, the speeds attainable at
limb extremities are remarkable. A skilled pitcher can hurl a baseball at a
speed in excess of 100 mph. Of course, this is also the speed of his hand at the
point where he releases the ball.
1.7 The Hip
Figure 1.14 shows the hip joint and its simplified lever representation, giving
dimensions that are typical for a male body. The hip is stabilized in its socket
by a group of muscles, which is represented in Fig. 1.14b as a single resultant
force Fm. When a person stands erect, the angle of this force is about 71◦
with respect to the horizon. WL represents the combined weight of the leg,
foot, and thigh. Typically, this weight is a fraction (0.185) of the total body
weight W (i.e., WL 0.185 W ). The weight WL is assumed to act vertically
downward at the midpoint of the limb.
We will now calculate the magnitude of the muscle force Fm and the force
FR at the hip joint when the person is standing erect on one foot as in a slow
walk, as shown in Fig. 1.14. The force W acting on the bottom of the lever is
the reaction force of the ground on the foot of the person. This is the force
that supports the weight of the body.
From equilibrium conditions, using the procedure outlined in Section 1.6,
we obtain
Fm cos 71◦
− FR cos θ 0 (x components of the
force 0) (1.18)
Fm sin 71◦
+ W − WL − FR sin θ 0 (y components of the
force 0) (1.19)
(FR sin θ) � 7 cm + WL � 10 cm
−W � 18 cm 0 (torque about
point A 0) (1.20)
16 Chapter 1 Static Forces
FIGURE 1.14 (a) The hip. (b) Its lever representation.
Since WL 0.185 W, from Eq. 1.20 we have
FR sin θ 2.31W
Using the result in Eq. 1.19, we obtain
Fm
1.50W
sin 71◦
1.59W (1.21)
Section 1.8 The Back 17
From Eq. 1.18, we obtain
FR cos θ 1.59W cos 71◦
0.52W
therefore,
θ tan−1
4.44 77.3◦
and
FR 2.37W (1.22)
This calculation shows that the force on the hip joint is nearly two and onehalf
times the weight of the person. Consider, for example, a person whose
mass is 70 kg and weight is 9.8 � 70 686 N (154 lb). The force on the hip
joint is 1625 N (366 lb).
1.7.1 Limping
Persons who have an injured hip limp by leaning toward the injured side as
they step on that foot (Fig. 1.15). As a result, the center of gravity of the
body shifts into a position more directly above the hip joint, decreasing the
force on the injured area. Calculations for the case in Fig. 1.15 show that
the muscle force Fm 0.47W and that the force on the hip joint is 1.28W
(see Exercise 1-11). This is a significant reduction from the forces applied
during a normal one-legged stance.
1.8 The Back
When the trunk is bent forward, the spine pivots mainly on the fifth lumbar
vertebra (Fig. 1.16a). We will analyze the forces involved when the trunk is
bent at 60◦ from the vertical with the arms hanging freely. The lever model
representing the situation is given in Fig. 1.16.
The pivot point A is the fifth lumbar vertebra. The lever arm AB represents
the back. The weight of the trunk W1 is uniformly distributed along the back;
its effect can be represented by a weight suspended in the middle. The weight
of the head and arms is represented by W2 suspended at the end of the lever
arm. The erector spinalis muscle, shown as the connection D-C attached at a
point two-thirds up the spine, maintains the position of the back. The angle
between the spine and this muscle is about 12◦. For a 70-kg man, W1 and W2
are typically 320 N (72 lb) and 160 N (36 lb), respectively.
Solution of the problem is left as an exercise. It shows that just to hold
up the body weight, the muscle must exert a force of 2000 N (450 lb) and
18 Chapter 1 Static Forces
FIGURE 1.15 Walking on an injured hip.
the compressional force of the fifth lumbar vertebra is 2230 N (500 lb). If, in
addition, the person holds a 20-kg weight in his hand, the force on the muscle
is 3220 N (725 lb), and the compression of the vertebra is 3490 N (785 lb)
(see Exercise 1-12).
This example indicates that large forces are exerted on the fifth lumbar
vertebra. It is not surprising that backaches originate most frequently at this
point. It is evident too that the position shown in the figure is not the recommended
way of lifting a weight.
Section 1.10 Dynamic Aspects of Posture 19
FIGURE 1.16 (Left) The bent back. (Right) Lever representation.
1.9 Standing Tip-Toe on One Foot
The position of the foot when standing on tiptoe is shown in Fig. 1.17. The
total weight of the person is supported by the reaction force at point A. This
is a Class 1 lever with the fulcrum at the contact of the tibia. The balancing
force is provided by the muscle connected to the heel by the Achilles tendon.
The dimensions and angles shown in Fig. 1.17b are reasonable values for
this situation. Calculations show that while standing tiptoe on one foot the
compressional force on the tibia is 3.5W and the tension force on the Achilles
tendon is 2.5 � W (see Exercise 1-13). Standing on tiptoe is a fairly strenuous
position.
1.10 Dynamic Aspects of Posture
In our treatment of the human body, we have assumed that the forces exerted
by the skeletal muscles are static. That is, they are constant in time. In fact,
the human body (and bodies of all animals) is a dynamic system continually
responding to stimuli generated internally and by the external environment.
Because the center of gravity while standing erect is about half the height
above the soles of the feet, even a slight displacement tends to topple the body.
20 Chapter 1 Static Forces
FIGURE 1.17 (a) Standing on tip-toe. (b) Lever model.
As has been demonstrated experimentally the simple act of standing upright
requires the body to be in a continual back and forth, left right, swaying motion
to maintain the center of gravity over the base of support. In a typical experiment
designed to study this aspect of posture, the person is instructed to stand,
feet together, as still as possible, on a platform that registers the forces applied
by the soles of the feet (center of pressure). To compensate for the shifting
center of gravity this center of pressure is continually shifting by several centimeters
over the area of the soles of the feet on a time scale of about half a
second. Small back-and-forth perturbations of the center of mass (displacements
less than about 1.5 cm) are compensated by ankle movements. Hip
movements are required to compensate for larger displacements as well as for
left right perturbations.
The maintaining of balance in the process of walking requires a yet more
complex series of compensating movements as the support for the center of
gravity shifts from one foot to the other. Keeping the body upright is a highly
complex task of the nervous system. The performance of this task is most
remarkable when accidentally we slip and the center of gravity is momentarily
displaced from the base of support. As is shown in Chapter 4, Exercise 4-9,
Chapter 1 Exercises 21
without compensating movements an erect human body that looses its balance
will hit the floor in about 1 sec. During this short time interval, the whole muscular
system is called into action by the “righting reflex” to mobilize various
parts of the body into motion so as to shift the center of mass back over the
base of support. The body can perform amazing contortions in the process of
restoring balance.
The nervous system obtains information required to maintain balance principally
from three sources: vision, the vestibular system situated in the inner
ear that monitors movement and position of the head, and somatosensory system
that monitors position and orientation of the various parts of the body. With
age, the efficiency of the functions required to keep a person upright decreases
resulting in an increasing number of injuries due to falls. In the United States,
the number of accidental deaths per capita due to falls for persons above the
age of 80 is about 60 times higher than for people below the age of 70.
Another aspect of the body dynamics is the interconnectedness of the
musculoskeletal system. Through one path or another, all muscles and bones
are connected to one another, and a change in muscle tension or limb position
in one part of the body must be accompanied by a compensating change
elsewhere. The system can be visualized as a complex tentlike structure. The
bones act as the tent poles and the muscles as the ropes bringing into and
balancing the body in the desired posture. The proper functioning of this type
of a structure requires that the forces be appropriately distributed over all the
bones and muscles. In a tent, when the forward-pulling ropes are tightened,
the tension in the back ropes must be correspondingly increased; otherwise,
the tent collapses in the forward direction. The musculoskeletal system operates
in an analogous way. For example, excessive tightness, perhaps through
overexertion, of the large muscles at the front of our legs will tend to pull the
torso forward. To compensate for this forward pull, the muscles in the back
must also tighten, often exerting excess force on the more delicate structures
of the lower back. In this way, excess tension in one set of muscles may be
reflected as pain in an entirely different part of the body.
EXERCISES
1-1. (a) Explain why the stability of a person against a toppling force is
increased by spreading the legs as shown in Fig. 1.7. (b) Calculate the
force required to topple a person of mass 70 kg, standing with his
feet spread 0.9 m apart as shown in Fig. 1.7. Assume the person does
not slide and the weight of the person is equally distributed on both feet.
1-2. Derive the relationships stated in Eqs. 1.6, 1.7, and 1.8.
22 Chapter 1 Static Forces
1-3. Using trigonometry, calculate the angle θ in Fig. 1.13. The dimensions
are specified in Fig. 1.12b.
1-4. Using the data provided in the text, calculate the maximum weight that
the arm can support in the position shown in Fig. 1.12.
1-5. Calculate the force applied by the biceps and the reaction force (Fr)
at the joint as a result of a 14-kg weight held in hand when the elbow
is at (a) 160◦ and (b) 60◦. Dimensions are as in Fig. 1.12.
Assume that the upper part of the arm remains fixed as in Fig. 1.12 and
use calculations from Exercise 1-3. Note that under these conditions
the lower part of the arm is no longer horizontal.
1-6. Consider again Fig. 1.12. Now let the 14-kg weight hang from the middle
of the lower arm (20 cm from the fulcrum). Calculate the biceps
force and the reaction force at the joint.
1-7. Consider the situation when the arm in Fig. 1.13 supports two 14-kg
weights, one held by the hand as in Fig. 1.13 and the other supported in
the middle of the arm as in Exercise 1-6. (a) Calculate the force of the
biceps muscle and the reaction force. (b) Are the forces calculated in
part (a) the same as the sum of the forces produced when the weights
are suspended individually?
1-8. Calculate the additional forces due to the weight of the arm itself in
Fig. 1.13. Assume that the lower part of the arm has a mass of 2 kg and
that its total weight can be considered to act at the middle of the lower
arm, as in Exercise 1-6.
1-9. Estimate the dimensions of your own arm, and draw a lever model for
the extension of the elbow by the triceps. Calculate the force of the triceps
in a one arm push-up in a hold position at an elbow angle of 100◦.
1-10. Suppose that the biceps in Fig. 1.13 contracts 2 cm. What is the upward
displacement of the weight? Suppose that the muscle contraction is uniform
in time and occurs in an interval of 0.5 sec. Compute the velocity
of the point of attachment of the tendon to the bone and the velocity
of the weight. Compare the ratio of the velocities to the mechanical
advantage.
1-11. Calculate the forces in the limping situation shown in Fig. 1.15. At
what angle does the force FR act?
1-12. (a) Calculate the force exerted by the muscle and the compression force
on the fifth lumbar vertebra in Fig. 1.16. Use information provided in
the text. (b) Repeat the calculations in (a) for the case when the person
shown in Fig. 1.16 holds a 20-kg weight in his hand.
1-13. Calculate the force on the tibia and on the Achilles tendon in Fig. 1.17.
Chapter 2
Friction
If we examine the surface of any object, we observe that it is irregular. It has
protrusions and valleys. Even surfaces that appear smooth to the eye show
such irregularities under microscopic examination. When two surfaces are in
contact, their irregularities intermesh, and as a result there is a resistance to
the sliding or moving of one surface on the other. This resistance is called
friction. If one surface is to be moved with respect to another, a force has to
be applied to overcome friction.
Consider a block resting on a surface as shown in Fig. 2.1. If we apply a
force F to the block, it will tend to move. But the intermeshing of surfaces
produces a frictional reaction force Ff that opposes motion. In order to
move the object along the surface, the applied force must overcome the
frictional force. The magnitude of the frictional force depends on the nature
of the surfaces; clearly, the rougher the surfaces, the greater is the frictional
force. The frictional property of the surfaces is represented by the coefficient
of friction μ. The magnitude of the frictional force depends also on the
force Fn perpendicular to the surfaces that presses the surfaces together. The
magnitude of the force that presses the surfaces together determines to what
extent the irregularities are intermeshed.
The frictional force Ff is given by
Ff μFn (2.1)
Distinction has to be made between the frictional force that acts on moving
object (called the kinetic frictional force) and the frictional force that acts on
the object when it is stationary. The kinetic frictional force opposing motion of
23
24 Chapter 2 Friction
FIGURE 2.1 Friction.
the object is obtained from Eq. 2.1 using the kinetic coefficient of friction μk.
In general, it takes a larger force to get the object moving against a frictional
force than to keep it in motion. This is not surprising because in the stationary
case the irregularities of the two surfaces can settle more deeply into each
other. The force that must be applied to an object to get it moving is again
obtained from Eq. 2.1 but this time using the static coefficient of friction μs.
This is the magnitude of the maximum static frictional force.
The magnitude of the frictional force does not depend on the size of the
contact area. If the surface contact area is increased, the force per unit area
(pressure) is decreased, and this reduces the interpenetration of the irregularities.
However, at the same time, the number of irregularities is proportionately
increased. As a result, the total frictional force is unchanged. Coefficients of
static and kinetic friction between some surfaces are shown in Table 2.1. As
is evident, the coefficient of static friction for two given surfaces is somewhat
larger than the coefficient of kinetic friction.
We have illustrated the concept of friction with surfaces sliding along each
other, but frictional forces are encountered also in rolling (rolling friction) and
in fluid flows (viscous friction). Rolling motion is not encountered in living
systems, but viscous friction plays an important role in the flow of blood and
other biological fluids.
Whereas sliding friction is independent of velocity, fluid friction has a
strong velocity dependence. We will discuss this in Chapter 3.
Friction is everywhere around us. It is both a nuisance and an indispensable
factor in the ability of animals to move. Without friction an object that
is pushed into motion would continue to move forever (Newton’s first law,
Appendix A). The slightest force would send us into eternal motion. It is the
frictional force that dissipates kinetic energy into heat and eventually stops the
Section 2.1 Standing at an Incline 25
TABLE 2.1 Coefficients of Friction,
Static (μs) and Kinetic (μk)
Surfaces μs μk
Leather on oak 0.6 0.5
Rubber on dry concrete 0.9 0.7
Steel on ice 0.02 0.01
Dry bone on bone 0.3
Bone on joint, lubricated 0.01 0.003
object (see Exercise 2-1). Without friction we could not walk; nor could we
balance on an inclined plane (see Exercise 2-2). In both cases, friction provides
the necessary reaction force. Friction also produces undesirable wear
and tear and destructive heating of contact surfaces. Both nature and engineers
attempt to maximize friction where it is necessary and minimize it where it is
destructive. Friction is greatly reduced by introducing a fluid such as oil at
the interface of two surfaces. The fluid fills the irregularities and therefore
smooths out the surfaces. A natural example of such lubrication occurs in
the joints of animals, which are lubricated by a fluid called the synovial fluid.
This lubricant reduces the coefficient of friction by about a factor of 100. As
is evident from Table 2.1, nature provides very efficient joint lubrication. The
coefficient of friction here is significantly lower than for steel on ice.
We will illustrate the effects of friction with a few examples.
2.1 Standing at an Incline
Referring to Fig. 2.2, let us calculate the angle of incline θ of an oak board on
which a person of weight W can stand without sliding down. Assume that she
is wearing leather-soled shoes and that she is standing in a vertical position as
shown in the figure.
The force Fn normal to the inclined surface is
Fn W cos θ (2.2)
The static frictional force Ff is
Ff μFn μsW cos θ 0.6W cos θ (2.3)
26 Chapter 2 Friction
FIGURE 2.2 Standing on an incline.
The force parallel to the surface Fp, which tends to cause the sliding, is
Fp W sin θ (2.4)
The person will slide when the force Fp is greater than the frictional force Ff ;
that is,
Fp > Ff (2.5)
At the onset of sliding, these two forces are just equal; therefore,
Ff Fp
0.6W cos θ W sin θ (2.6)
or
sin θ
cos θ
tan θ 0.6
Therefore θ 31◦.
2.2 Friction at the Hip Joint
We have shown in Chapter 1 that the forces acting on the joints are very large.
When the joints are in motion, these large forces produce frictional wear,
Section 2.3 Spine Fin of a Catfish 27
which could be damaging unless the joints are well lubricated. Frictional wear
at the joints is greatly reduced by a smooth cartilage coating at the contact ends
of the bone and by synovial fluid which lubricates the contact areas. We will
now examine the effect of lubrication on the hip joint in a person. When a
person walks, the full weight of the body rests on one leg through most of
each step. Because the center of gravity is not directly above the joint, the
force on the joint is greater than the weight. Depending on the speed of walking,
this force is about 2.4 times the weight (see Chapter 1). In each step, the
joint rotates through about 60◦. Since the radius of the joint is about 3 cm, the
joint slides about 3 cm inside the socket during each step. The frictional force
on the joint is
Ff 2.4Wμ (2.7)
The work expended in sliding the joint against this friction is the product of the
frictional force and the distance over which the force acts (see Appendix A).
Thus, the work expended during each step is
Work Ff � distance 2.4 Wμ(3 cm) 7.2 μW erg (2.8)
If the joint were not lubricated, the coefficient of friction (μ) would be about
0.3. Under these conditions, the work expended would be
Work 2.16 � W erg (2.9)
This is a large amount of work to expend on each step. It is equivalent to
lifting the full weight of the person 2.16 cm. Furthermore, this work would be
dissipated into heat energy, which would destroy the joint.
As it is, the joint is well lubricated, and the coefficient of friction is only
0.003. Therefore, the work expended in counteracting friction and the resultant
heating of the joint are negligible. However, as we age, the joint cartilage
begins to wear, efficiency of lubrication decreases, and the joints may become
seriously damaged. Studies indicate that by the age of 70 about two-thirds of
people have knee joint problems and about one-third have hip problems.
2.3 Spine Fin of a Catfish
Although in most cases good lubrication of bone-contact surfaces is essential,
there are a few cases in nature where bone contacts are purposely unlubricated
to increase friction. The catfish has such a joint connecting its dorsal spine fin
to the rest of its skeleton (Fig. 2.3). Normally the fin is folded flat against the
28 Chapter 2 Friction
body, but when the fish is attacked, the appropriate muscles pull the bone of
the fin into a space provided in the underlying skeleton. Since the coefficient
of friction between the fin bone and the skeleton is high, the frictional force
tends to lock the fin in the up position. In order to remove the fin, a force must
be applied in a predominantly vertical direction with respect to the underlying
skeleton. The erect sharp fin discourages predators from eating the catfish.
Figure 2.3b is a simplified representation of the spine and the protruding
fin. The shaded block represents the movable fin bone, and the horizontal
block is the spine holding the fin. Assume that a force F at an angle θ is
applied at point A to dislodge the bone. The force is shown to act at point A,
FIGURE 2.3 (a) Catfish. (b) Simplified representation of the spine in the catfish.
Chapter 2 Exercises 29
2.5 cm above point B. The dimensions shown in the figure are to be used in
the calculations required for Exercise 2-3. The applied force tips the bone, and
as a result reaction forces are set up at points B and C. The components of
these forces normal to the fin-bone surface produce frictional forces that resist
removal of the bone. Calculation of some of the properties of the locking
mechanism is left as an exercise.
EXERCISES
2-1. (a) Assume that a 50-kg skater, on level ice, has built up her speed to
30 km/h. How far will she coast before the sliding friction dissipates her
energy? (Kinetic energy 1
2 mv2; see Appendix A.) (b) How does the
distance of coasting depend on the mass of the skater?
2-2. Referring to Fig. 1.5, compute the coefficient of friction at which the tendency
of the body to slide and the tendency to topple due to the applied
force are equal.
2-3. (a) Referring to Fig. 2.3, assume that a dislodging force of 0.1 N is
applied at θ 20◦ and the angle between the fin bone and the spine is
45◦. Calculate the minimum value for the coefficient of friction between
the bones to prevent dislodging of the bone. (b) Assuming that the coefficient
of friction is 1.0, what is the value of the angle θ at which a force
of 0.2 N will just dislodge the bone? What would this angle be if the
bones were lubricated (μ 0.01)?
Chapter 3
Translational Motion
In general, the motion of a body can be described in terms of translational and
rotational motion. In pure translational motion all parts of the body have the
same velocity and acceleration (Fig. 3.1). In pure rotational motion, such as
the rotation of a bar around a pivot, the rate of change in the angle θ is the same
for all parts of the body (Fig. 3.2), but the velocity and acceleration along the
body depend on the distance from the center of rotation. Many motions and
movements encountered in nature are combinations of rotation and translation,
as in the case of a body that rotates while falling. It is convenient, however,
to discuss these motions separately. In this chapter, we discuss translation.
Rotation is discussed in the following chapter.
Theequationsoftranslationalmotionforconstantaccelerationarepresented
in Appendix A and may be summarized as follows: In uniform acceleration,
the final velocity (v) of an object that has been accelerated for a time t is
v v0 + at (3.1)
Here v0 is the initial velocity of the object and a is the acceleration. Acceleration
can therefore be expressed as
a
v − v0
t
(3.2)
The average velocity during the time interval t is
vav
v + v0
2
(3.3)
30
Chapter 3 Translational Motion 31
FIGURE 3.1 Translational motion.
FIGURE 3.2 Rotational motion.
The distance s traversed during this time is
s vavt (3.4)
Using Eqs. 3.1 and 3.2, we obtain
s v0t
at2
2
(3.5)
By substituting t (v − v0)/a from Eq. 3.1 into Eq. 3.5, we obtain
v2
v2
0 + 2as (3.6)
Let us now apply these equations to some problems in the life sciences.
Most of our calculations will relate to various aspects of jumping. Although in
32 Chapter 3 Translational Motion
the process of jumping the acceleration of the body is usually not constant, the
assumption of constant acceleration is necessary to solve the problems without
undue difficulties.
3.1 Vertical Jump
Consider a simple vertical jump in which the jumper starts in a crouched
position and then pushes off with her feet (Fig. 3.3).
We will calculate here the height H attained by the jumper. In the crouched
position, at the start of the jump, the center of gravity is lowered by a distance
c. During the act of jumping, the legs generate a force by pressing down
on the surface. Although this force varies through the jump, we will assume
that it has a constant average value F.
Because the feet of the jumper exert a force on the surface, an equal
upward-directed force is exerted by the surface on the jumper (Newton’s third
law). Thus, there are two forces acting on the jumper: her weight (W ), which
is in the downward direction, and the reaction force (F ), which is in the
upward direction. The net upward force on the jumper is F − W (see Fig. 3.4).
This force acts on the jumper until her body is erect and her feet leave the
ground. The upward force, therefore, acts on the jumper through a distance c
FIGURE 3.3 Vertical jump.
Section 3.1 Vertical Jump 33
FIGURE 3.4 Forces on the jumper.
(see Fig. 3.3). The acceleration of the jumper in this stage of the jump (see
Appendix A) is
a
F − W
m
F − W
W/g
(3.7)
where W is the weight of the jumper and g is the gravitational acceleration.
A consideration of the forces acting on the Earth (Fig. 3.5) shows that an equal
force accelerates the Earth in the opposite direction. However, the mass of the
Earth is so large that its acceleration due to the jump is negligible.
The acceleration shown in Eq. 3.7 takes place over a distance c. Therefore,
the velocity v of the jumper at take-off as given by Eq. 3.6 is
v2
v2
0 + 2ac (3.8)
Since the initial velocity at the start of the jump is zero (i.e., v0 0), the
take-off velocity is
v2 2 (F − W ) c
W/g
(3.9)
(Here we have substituted
a
F − W
W/g
into Eq. 3.8.)
34 Chapter 3 Translational Motion
FIGURE 3.5 Forces on the Earth.
After the body leaves the ground, the only force acting on it is the force
of gravity W, which produces a downward acceleration −g on the body. At
the maximum height H, just before the body starts falling back to the ground,
the velocity is zero. The initial velocity for this part of the jump is the take-off
velocity v given by Eq. 3.9. Therefore, from Eq. 3.6, we obtain
0
2 (F − W ) c
W/g
− 2gH (3.10)
From this, the height of the jump is
H
(F − W ) c
W
(3.11)
Now let us estimate the numerical value for the height of the jump. Experiments
have shown that in a good jump a well-built person generates an average
reaction force that is twice his/her weight (i.e., F 2W). In that case, the
height of jump is H c. The distance c, which is the lowering of the center of
gravity in the crouch, is proportional to the length of the legs. For an average
person, this distance is about 60 cm, which is our estimate for the height of a
vertical jump.
The height of a vertical jump can also be computed very simply from
energy considerations. The work done on the body of the jumper by the force
F during the jump is the product of the force F and the distance c over which
this force acts (see Appendix A). This work is converted to kinetic energy as
the jumper is accelerated upward. At the full height of the jump H (before
the jumper starts falling back to ground), the velocity of the jumper is zero.
At this point, the kinetic energy is fully converted to potential energy as the
center of mass of the jumper is raised to a height (c + H). Therefore, from
Section 3.2 Effect of Gravity on the Vertical Jump 35
conservation of energy,
Work done on the body Potential energy at maximum height
or
Fc W(c + H ) (3.12)
From this equation the height of the jump is, as before,
H
(F − W ) c
W
Another aspect of the vertical jump is examined in Exercise 3-1.
3.2 Effect of Gravity on the Vertical Jump
The weight of an object depends on the mass and size of the planet on which
it is located. The gravitational constant of the moon, for example, is one-sixth
that of the Earth; therefore, the weight of a given object on the moon is onesixth
its weight on the Earth. It is a common mistake to assume that the height
to which a person can jump on the moon increases in direct proportion to the
decrease in weight. This is not the case, as the following calculation will show.
From Eq. 3.11, the height of the jump on the Earth is
H
(F − W ) c
W
The force F that accelerates the body upward depends on the strength of the
leg muscles, and for a given person this force is the same on the moon as on
the Earth. Similarly, the lowering of the center of gravity c is unchanged with
location. Therefore, the height of the jump on the moon (H ) is
H
(F − W ) c
W
(3.13)
Here W is the weight of the person on the moon (i.e., W W/6). The ratio
of the jumping heights at the two locations is
H
H
(F − W )W
(F − W)W
(3.14)
If as before we assume that F 2W, we find that H /H 11. That is, if a
person can jump to a height of 60 cm on Earth, that same person can jump up
6.6 m on the moon. Note that the ratio H /H 11 is true only for a particular
choice of F in the calculation (see Exercise 3-2).
36 Chapter 3 Translational Motion
3.3 Running High Jump
In the preceding sections, we calculated the height of a jump from a standing
position and showed that the center of gravity could be raised about 60 cm.
A considerably greater height can be attained by jumping from a running start.
The current high-jump record is about 2.3 m. The additional height is attained
by using part of the kinetic energy of the run to raise the center of gravity
off the ground. Let us calculate the height attainable in a running jump if the
jumper could use all his/her initial kinetic energy (1
2mv2) to raise his/her body
off the ground. If this energy were completely converted to potential energy
by raising the center of gravity to a height H, then
MgH
1
2
mv2
(3.15)
or
H
v2
2g
To complete our estimate, we must consider two additional factors that
increase the height of the jump. First, we should add the 0.6 m, which can
be produced by the legs in the final push-off. Then we must remember that the
center of gravity of a person is already about 1 m above the ground. With little
extra effort, the jumper can alter the position of his body so that it is horizontal
at its maximum height. This maneuver adds 1 m to the height of the bar he
can clear. Thus, our final estimate for the maximum height of the running high
jump is
H
v2
2g
+ 1.6 m (3.16)
The maximum short distance speed of a good runner is about 10 m/sec. At
this speed, our estimate for the maximum height of the jump from Eq. 3.16 is
6.7 m. This estimate is nearly three times the high-jump record. Obviously,
it is not possible for a jumper to convert all the kinetic energy of a full-speed
run into potential energy.
In the unaided running high jump, only the force exerted by the feet is
available to alter the direction of the running start. This limits the amount
of kinetic energy that can be utilized to aid the jump. The situation is quite
different in pole vaulting, where, with the aid of the pole, the jumper can in fact
use most of the kinetic energy to raise his/her center of gravity. The current
men’s pole-vaulting record is 6.15 m (20 ft 2 in), which is remarkably close
Section 3.5 Standing Broad Jump 37
FIGURE 3.6 Projectile.
to our estimate of 6.7 m. These figures would agree even more closely had
we included in our estimate the fact that the jumper must retain some forward
velocity to carry him/her over the bar.
3.4 Range of a Projectile
A problem that is solved in most basic physics texts concerns a projectile
launched at an angle θ and with initial velocity v0. A solution is required for
the range R, the distance at which the projectile hits the Earth (see Fig. 3.6).
It is shown that the range is
R
v2
0 sin 2θ
g
(3.17)
For a given initial velocity the range is maximum when sin 2θ 1 or θ 45◦.
In other words a maximum range is obtained when the projectile is launched
at a 45◦ angle. In that case the range is
Rmax
v2
0
g
(3.18)
Using this result, we will estimate the distance attainable in broad jumping.
3.5 Standing Broad Jump
When the jumper projects himself into the broad jump from a stationary crouching
position (Fig. 3.7), his acceleration is determined by the resultant of two
forces: the downward force of gravity, which is simply equal to his weight W,
38 Chapter 3 Translational Motion
FIGURE 3.7 (a) The standing broad jump. (b) The associated forces.
and the force generated by the feet, which he can apply in any direction.
In order to maximize the distance of the jump, the launching velocity and
therefore also the resultant force should be directed at a 45◦ angle.
We will assume as before that a jumper can generate with his feet a force
equal to twice the body weight. The magnitude of the resultant force (Fr) and
the angle θ at which the legs must apply the force to the body are obtained
from the following considerations.
The horizontal and vertical components of the resultant force (see Fig. 3.7)
are, respectively,
Horizontal component of Fr: Fr cos 45◦
2W cos θ (3.19)
and
Vertical component of Fr: Fr sin 45◦
2W sin θ − W (3.20)
Here we have two equations that can be solved to yield the two unknown
quantities Fr and θ (see Exercise 3-3). The magnitude of the force Fr is
Fr 1.16W
The optimum angle θ at which the legs apply the force 2W is θ 65.8◦. We
will again assume that the force that launches the jumper is applied over a
Section 3.6 Running Broad Jump (Long Jump) 39
distanceof60 cm,whichistheextentofthecrouchingposition. Theacceleration
produced by the resultant force is
a
Fr
m
1.16W
W/g
1.16g
The launching velocity v of the jumper is therefore v2 2as. With s 60 cm,
the velocity is 3.70 m/sec. The distance (R) of the standing broad jump is,
from Eq. 3.18,
R
v2
g
13.7
9.8
1.4 m
The range of the jump can be increased by swinging both the legs and
the arms in the direction of the jump, which results in an increase in the forward
momentum of the body. Other aspects of the standing broad jump are
presented in Exercises 3-4 and 3-5.
3.6 Running Broad Jump (Long Jump)
Let us assume that a jumper launches the jump from a full speed of 10 m/sec.
The push-off force (2W) generated by the legs provides the vertical component
of the launching velocity. From this force we have to subtract the weight (W)
of the jumper. The acceleration produced by the net force is
a
2W − W
m
W
W/g
g
If the push-off force acts on the jumper over a distance of 60 cm (the extent of
the crouch) and if it is directed entirely in the vertical y direction, the vertical
component of the velocity vy during the jump is given by
v2
y 2as 2 � g � 0.6 11.8 m2
/sec2
vy 3.44 m/sec
Since the horizontal component of the launching velocity vx is the running
velocity, the magnitude of the launching velocity is
v v2
x + v2
y 10.6 m/sec
40 Chapter 3 Translational Motion
The launch angle θ is
θ tan−1 vy
vx
tan−1 3.44
10
19◦
From Eq. 3.17, the range R of the jump is
R
v2 sin 2θ
g
112.4 sin 38◦
g
7.06 m
Considering the approximations used in these calculations, this estimate is in
reasonable agreement with the current world record, of about 9 m for men and
7.5 m for women.
3.7 Motion through Air
We have so far neglected the effect of air resistance on the motion of objects,
but we know from experience that this is not a negligible effect. When an
object moves through the air, the air molecules have to be pushed out of its
way. The resulting reaction force pushes back on the body and retards its
motion—this is the source of fluid friction in air. We can deduce some of the
properties of air friction by sticking our hand outside a moving car. Clearly, the
greater the velocity with respect to the air, the larger is the resistive force. By
rotating our hand, we observe that the force is greater when the palms face the
direction of motion. We therefore conclude that the resistive force increases
with the velocity and the surface area in the direction of motion. It has been
found that the force due to air resistance Fa can be expressed approximately as
Fa CAv2
(3.21)
where v is the velocity of the object with respect to the air, A is the area facing
the direction of motion, and C is the coefficient of air friction. The coefficient
C depends somewhat on the shape of the object. In our calculations, we will
use the value C 0.88 kg/m3.
Because of air resistance, there are two forces acting on a falling body: the
downward force of gravity W and the upward force of air resistance. From
Newton’s second law (see Appendix A), we find that the equation of motion
in this case is
W − Fa ma (3.22)
When the body begins to fall, its velocity is zero and the only force acting
on it is the weight; but as the body gains speed, the force of air resistance
Section 3.7 Motion through Air 41
grows, and the net accelerating force on the body decreases. If the body falls
from a sufficiently great height, the velocity reaches a magnitude such that the
force due to air resistance is equal to the weight. Past this point, the body
is no longer accelerated and continues to fall at a constant velocity, called
the terminal velocity vt. Because the force on the body in Eq. 3.22 is not
constant, the solution of this equation cannot be obtained by simple algebraic
techniques. However, the terminal velocity can be obtained without difficulty.
At the terminal velocity, the downward force of gravity is canceled by the
upward force of air resistance, and the net acceleration of the body is zero.
That is,
W − Fa 0 (3.23)
or
Fa W
From Eq. 3.21, the terminal velocity is therefore given by
vt
W
CA
(3.24)
From this equation the terminal velocity of a falling person with mass
70 kg and an effective area of 0.2 m2 is
vt
W
CA
70 � 9.8
0.88 � 0.2
62.4 m/sec (140 mph)
The terminal velocity of different-sized objects that have a similar density
and shape is proportional to the square root of the linear size of the objects.
This can be seen from the following argument. The weight of an object is
proportional to the volume, which is in turn proportional to the cube of the
linear dimension L of the object,
W ∝ L3
The area is proportional to L2. Therefore, from Eq. 3.24, the terminal velocity
is proportional to
√
L as shown here:
vt ∝
W
A
L3
L2
L
This result has interesting implications on the ability of animals to survive
a fall. With proper training, a person can jump from a height of about 10 m
42 Chapter 3 Translational Motion
without sustaining serious injury. From this height, a person hits the ground
at a speed of
v 2gs 14 m/sec (46 ft/sec)
Let us assume that this is the speed with which any animal can hit the
ground without injury. At this speed, the force of air resistance on an animal
the size of man is negligible compared to the weight. But a small animal is
slowed down considerably by air friction at this speed. A speed of 8.6 m/sec is
the terminal velocity of a 1-cm bug (see Exercise 3-6). Such a small creature
can drop from any height without injury. Miners often encounter mice in deep
coal mines but seldom rats. A simple calculation shows that a mouse can
fall down a 100-m mine shaft without severe injury. However, such a fall will
kill a rat.
Air friction has an important effect on the speed of falling raindrops and
hailstones. Without air friction, a 1-cm diameter hailstone, for example, falling
from a height of 1000 m would hit the Earth at a speed of about 140 m/sec.
At such speeds the hailstone would certainly injure anyone on whom it fell.
As it is, air friction slows the hailstone to a safe terminal velocity of about
8.3 m/sec (see Exercise 3-8).
3.8 Energy Consumed in Physical Activity
Animals do work by means of muscular movement. The energy required to
perform the work is obtained from the chemical energy in the food eaten by
the animal. In general, only a small fraction of the energy consumed by the
muscles is converted to work. For example, in bicycling at a rate of one leg
extension per second, the efficiency of the muscles is 20%. In other words
only one fifth of the chemical energy consumed by the muscle is converted
to work. The rest is dissipated as heat. The energy consumed per unit time
during a given activity is called the metabolic rate.
Muscle efficiency depends on the type of work and on the muscles
involved. In most cases, the efficiency of the muscles in converting caloric
food energy to work is less than 20%. However, in our subsequent calculations
we will assume a 20% muscular efficiency.
We will calculate the amount of energy consumed by a 70-kg person jumping
up 60 cm for 10 minutes at a rate of one jump per second. The external
mechanical work performed by the leg muscles in each jump is
Weight � Height of jump 70 kg � 9.8 � 0.6 411 J
Chapter 3 Exercises 43
The total muscle work during the 10 minutes of jumping is
411 � 600 jumps 24.7 � 104
J
If we assume a muscle efficiency of 20%, then in the act of jumping the body
consumes
24.7 � 104
� 5 1.23 � 106
J 294 � 103
cal 294 kcal
This is about the energy content in two doughnuts.
In a similar vein, A. H. Cromer (see Bibliography) calculates the metabolic
rate while running. In the calculation, it is assumed that most of the work done
in running is due to the leg muscles accelerating each leg to the running speed
v, and then decelerating it to 0 velocity as one leg is brought to rest and the
other leg is accelerated. The work in accelerating the leg of mass m is 1
2mv2.
The work done in the deceleration is also 1
2mv2. Therefore, the total amount of
work done during each stride is mv2. As is shown in Exercise 3-9, typically, a
70-kg person (leg mass 10 kg) running at 3 m/sec (9-min. mile) with a muscle
efficiency of 20%, and step length of 1 m, expends 1350 J/sec or 1160 kcal/h.
Thisisingoodagreementwithmeasurements. Theenergyrequiredtoovercome
air resistance in running is calculated in Exercise 3-10.
In connection with the energy consumption during physical activity, we
should note the difference between work and muscular effort. Work is defined
as the product of force and the distance over which the force acts (see
Appendix A). When a person pushes against a fixed wall his/her muscles
are not performing any external work because the wall does not move. Yet
it is evident that considerable energy is used in the act of pushing. All the
energy is expended in the body to keep the muscles balanced in the tension
necessary for the act of pushing.
EXERCISES
3-1. Experiments show that the duration of upward acceleration in the
standing vertical jump is about 0.2 sec. Calculate the power generated in
a 60-cm jump by a 70-kg jumper assuming that c H, as in the text.
3-2. A 70-kg astronaut is loaded so heavily with equipment that on Earth
he can jump only to a height of 10 cm. How high can he jump on the
Moon? (Use the assumptions related to Eq. 3.11 in the text. As in the
text, assume that the force generated by the legs is twice the unloaded
weight of the person and the gravitational constant on the moon is 1/6
that on Earth.)
44 Chapter 3 Translational Motion
3-3. Solve Eqs. 3.19 and 3.20 for the two unknowns Fr and θ.
3-4. What is the time period in the standing broad jump during which the
jumper is in the air? Assume that the conditions of the jump are as
described in the text.
3-5. Consider a person on the moon who launches herself into a standing
broad jump at 45◦. The average force generated during launching is,
as stated in the text, F 2W, and the distance over which this force
acts is 60 cm. The gravitational constant on the moon is 1/6 that on
earth. Compute (a) the range of the jump; (b) the maximum height of
the jump; (c) the duration of the jump.
3-6. Calculate the terminal velocity of a 1-cm bug. Assume that the density
of the bug is 1 g/cm3 and that the bug is spherical in shape with a
diameter of 1 cm. Assume further that the area of the bug subject to air
friction is πr2.
3-7. Calculate the radius of a parachute that will slow a 70-kg parachutist to
a terminal velocity of 14 m/sec.
3-8. Calculate the terminal velocity of a (a) 1-cm diameter hailstone and
(b) a 4-cm diameter hailstone. Density of ice is 0.92 gm/cm3. Assume
that the area subject to air friction is πr2.
3-9. Using the approach discussed in the text, calculate the energy expended
per second by a person running at 3 m/sec (9-min. mile) with a muscle
efficiency of 20%. Assume that the leg mass m 10 kg and the step
length is 1 m.
3-10. Compute the power necessary to overcome air resistance when running
at 4.5 m/sec (6-min. mile) against a 30 km/h wind. (Use data in the text
and assume area of person facing the wind is 0.2 m2.)
Chapter 4
Angular Motion
As was stated in Chapter 3, most natural movements of animals consist of
both linear and angular motion. In this chapter, we will analyze some aspects
of angular motion contained in the movement of animals. The basic equations
and definitions of angular motion used in this chapter are reviewed in
Appendix A.
4.1 Forces on a Curved Path
The simplest angular motion is one in which the body moves along a curved
path at a constant angular velocity, as when a runner travels along a circular
path or an automobile rounds a curve. The usual problem here is to calculate
the centrifugal forces and determine their effect on the motion of the
object.
A common problem solved in many basic physics texts requires determination
of the maximum speed at which an automobile can round a curve
without skidding. We will solve this problem because it leads naturally to an
analysis of running. Consider a car of weight W moving on a curved level
road that has a radius of curvature R. The centrifugal force Fc exerted on the
moving car (see Appendix A) is
Fc
mv2
R
Wv2
gR
(4.1)
45
46 Chapter 4 Angular Motion
For the car to remain on the curved path, a centripetal force must be
provided by the frictional force between the road and the tires. The car begins
to skid on the curve when the centrifugal force is greater than the frictional
force.
When the car is on the verge of skidding, the centrifugal force is just equal
to the frictional force; that is,
Wv2
gR
μW (4.2)
Here μ is the coefficient of friction between the tires and the road surface.
From Eq. 4.2, the maximum velocity vmax without skidding is
vmax μgR (4.3)
Safe speed on a curved path may be increased by banking the road along
the curve. If the road is properly banked, skidding may be prevented without
recourse to frictional forces. Figure 4.1 shows a car rounding a curve banked
at an angle θ. In the absence of friction, the reaction force Fn acting on the
car must be perpendicular to the road surface. The vertical component of this
force supports the weight of the car. That is,
Fn cos θ W (4.4)
To prevent skidding on a frictionless surface, the total centripetal force must
be provided by the horizontal component of Fn; that is,
Fn sin θ
Wv2
gR
(4.5)
where R is the radius of road curvature.
FIGURE 4.1 Banked curve.
Section 4.2 A Runner on a Curved Track 47
The angle θ for the road bank is obtained by taking the ratio of Eqs. 4.4
and 4.5. This yields
tan θ
v2
gR
(4.6)
4.2 A Runner on a Curved Track
A runner on a circular track is subject to the same type of forces described
in discussion of the automobile. As the runner rounds the curve, she leans
toward the center of rotation (Fig. 4.2a). The reason for this position can be
understood from an analysis of the forces acting on the runner. Her foot, as it
makes contact with the ground, is subject to the two forces, shown in Fig. 4.2b:
an upward force W, which supports her weight, and a centripetal reaction force
Fcp, which counteracts the centrifugal force. The resultant force Fr acts on
the runner at an angle θ with respect to the vertical axis.
If the runner were to round the curve remaining perpendicular to the surface,
this resultant force would not pass through her center of gravity and
an unbalancing torque would be applied on the runner (see Exercise 4-1). If
the runner adjusts her position by leaning at an angle θ toward the center of
FIGURE 4.2 (a) Runner on a curved track. (b) Forces acting on the foot of the
runner.
48 Chapter 4 Angular Motion
rotation, the resultant force Fr passes through her center of gravity and the
unbalancing torque is eliminated.
The angle θ is obtained from the relationships (see Fig. 4.2b)
Fr sin θ Fcp
Wv2
gR
(4.7)
and
Fr cos θ W (4.8)
Therefore
tan θ
v2
gR
(4.9)
The proper angle for a speed of 6.7 m/sec (this is a 4-min. mile) on a 15-mradius
track is
tan θ
(6.7)2
9.8 � 15
0.305
θ 17◦
No conscious effort is required to lean into the curve. The body automatically
balances itself at the proper angle. Other aspects of centrifugal force are
examined in Exercises 4-2, 4-3, and 4-4.
4.3 Pendulum
Since the limbs of animals are pivoted at the joints, the swinging motion of
animals is basically angular. Many of the limb movements in walking and
running can be analyzed in terms of the swinging movement of a pendulum.
The simple pendulum shown in Fig. 4.3 consists of a weight attached to
a string, the other end of which is attached to a fixed point. If the pendulum
is displaced a distance A from the center position and then released, it will
swing back and forth under the force of gravity. Such a back-and-forth movement
is called a simple harmonic motion. The number of times the pendulum
swings back and forth per second is called frequency (f ). The time for completing
one cycle of the motion (i.e., from A to A and back to A) is called the
Section 4.3 Pendulum 49
FIGURE 4.3 The simple pendulum.
period T. Frequency and period are inversely related; that is, T 1/f. If the
angle of displacement is small, the period is given by
T
1
f
2π
g
(4.10)
where g is the gravitational acceleration and is the length of the pendulum
arm. Although this expression for T is derived for a small-angle swing, it is a
good approximation even for a relatively wide swing. For example, when the
swing is through 120◦ (60◦ in each direction), the period is only 7% longer
than predicted by Eq. 4.10.
As the pendulum swings, there is continuous interchange between potential
and kinetic energy. At the extreme of the swing, the pendulum is momentarily
stationary. Here its energy is entirely in the form of potential energy. At
this point, the pendulum, subject to acceleration due to the force of gravity,
starts its return toward the center. The acceleration is tangential to the path of
the swing and is at a maximum when the pendulum begins to return toward
the center. The maximum tangential acceleration amax at this point is given by
amax
4π2A
T2
(4.11)
50 Chapter 4 Angular Motion
As the pendulum is accelerated toward the center, its velocity increases,
and the potential energy is converted to kinetic energy. The velocity of the
pendulum is at its maximum when the pendulum passes the center position (0).
At this point the energy is entirely in the form of kinetic energy, and the velocity
(vmax) here is given by
vmax
2πA
T
(4.12)
4.4 Walking
Some aspects of walking can be analyzed in terms of the simple harmonic
motion of a pendulum. The motion of one foot in each step can be considered
as approximately a half-cycle of a simple harmonic motion (Fig. 4.4). Assume
that a person walks at a rate of 120 steps/min (2 steps/sec) and that each step
is 90 cm long. In the process of walking each foot rests on the ground for
0.5 sec and then swings forward 180 cm and comes to rest again 90 cm ahead
of the other foot. Since the forward swing takes 0.5 sec, the full period of the
harmonic motion is 1 sec. The speed of walking v is
v 90 cm � 2 steps/sec 1.8 m/sec (4 mph)
FIGURE 4.4 Walking.
Section 4.5 Physical Pendulum 51
The maximum velocity of the swinging foot vmax is, from Eq. 4.12,
vmax
2πA
T
6.28 � 90
1
5.65 m/sec (12.6 mph)
Thus, at its maximum velocity, the foot moves about three times faster than
the body. The maximum acceleration is
amax
4π2A
T2
35.4 m/sec2
This is 3.6 times acceleration of gravity. These formulas can also be applied
to running (see Exercise 4-5).
4.5 Physical Pendulum
The simple pendulum shown in Fig. 4.3 is not an adequate representation of
the swinging leg because it assumes that the total mass is located at the end
of the pendulum while the pendulum arm itself is weightless. A more realistic
model is the physical pendulum, which takes into account the distribution
of weight along the swinging object (see Fig. 4.5). It can be shown
FIGURE 4.5 The physical pendulum.
52 Chapter 4 Angular Motion
(see [6-23]1) that under the force of gravity the period of oscillation T for a
physical pendulum is
T 2π
I
Wr
(4.13)
Here I is the moment of inertia of the pendulum around the pivot point O (see
Appendix A); W is the total weight of the pendulum, and r is the distance of
the center of gravity from the pivot point. (The expression for the period in
Eq. 4.13 is again strictly correct only for small angular displacement.)
4.6 Speed of Walking and Running
In the analysis of walking and running, the leg may be regarded as a physical
pendulum with a moment of inertia of a thin rod pivoted at one end. The
moment of inertia I for the leg (see Appendix A) is, therefore,
I
m 2
3
W
g
2
3
(4.14)
where W is the weight of the leg and is its length. If we assume that the
center of mass of the leg is at its middle (r 1
2 ), the period of oscillation is
T 2π
I
Wr
2π
(W/g)( 2/3)
W /2
2π
2
3 g
(4.15)
For a 90-cm-long leg, the period is 1.6 sec.
Because each step in the act of walking can be regarded as a half-swing
of a simple harmonic motion, the number of steps per second is simply the
inverse of the half period. In a most effortless walk, the legs swing at their
natural frequency, and the time for one step is T/2. Walking faster or slower
requires additional muscular exertion and is more tiring. In Exercise 4-6 we
calculate that for a person with 90 cm long legs and 90 cm step length the most
effortless walking speed is 1.13 m/sec (2.53 miles/h). Similar considerations
apply to the swinging of the arms (see Exercise 4-7).
We can now deduce the effect of the walker’s size on the speed of walking.
The speed of walking is proportional to the product of the number of steps
taken in a given time and the length of the step. The size of the step is in turn
1
References to the bibliography are given in square brackets.
Section 4.6 Speed of Walking and Running 53
proportional to the length of the leg . Therefore, the speed of walking v is
proportional to
v ∝
1
T
� (4.16)
But because /T is proportional to 1/ (see Eq. 4.15)
v ∝
1
√ (4.17)
Thus, the speed of the natural walk of a person increases as the square root of
the length of his/her legs. The same considerations apply to all animals: The
natural walk of a small animal is slower than that of a large animal.
The situation is different when a person (or any other animal) runs at full
speed. Whereas in a natural walk the swing torque is produced primarily by
gravity, in a fast run the torque is produced mostly by the muscles. Using
some reasonable assumptions, we can show that similarly built animals can
run at the same maximum speed, regardless of differences in leg size.
We assume that the length of the leg muscles is proportional to the length
of the leg ( ) and that the area of the leg muscles is proportional to 2. The
mass of the leg is proportional to 3. In other words, if one animal has a leg
twice as long as that of another animal, the area of its muscle is four times as
large and the mass of its leg is eight times as large.
The maximum force that a muscle can produce Fm is proportional to the
area of the muscle. The maximum torque Lmax produced by the muscle is
proportional to the product of the force and the length of the leg; that is,
Lmax Fm ∝ 3
The expression in the equation for the period of oscillation is applicable
for a pendulum swinging under the force of gravity. In general, the period of
oscillation for a physical pendulum under the action of a torque with maximum
value of Lmax is given by
T 2π
I
Lmax
(4.18)
Because the mass of the leg is proportional to 3, the moment of inertia (from
Eq. 4.14) is proportional to 5. Therefore, the period of oscillation in this case
54 Chapter 4 Angular Motion
is proportional to as shown
T ∝
5
3
The maximum speed of running vmax is again proportional to the product
of the number of steps per second and the length of the step. Because the
length of the step is proportional to the length of the leg, we have
vmax ∝
1
T
∝
1
� 1
This shows that the maximum speed of running is independent of the leg size,
which is in accordance with observation: A fox, for example, can run at about
the same speed as a horse.
Equations 4.10 and 4.15 illustrate another clearly observed aspect of running.
When a person runs at a slow pace, the arms are straight as in walking.
However, as the speed of running (that is the number of steps in a given interval)
increases, the elbows naturally assume a bent position. In this way, the
effective length ( ) of the pendulum decreases. This in turn increases the natural
frequency of the arm, bringing it into closer synchrony with the increased
frequency of steps.
4.7 Energy Expended in Running
In Chapter 3, we obtained the energy expended in running by calculating the
energy needed to accelerate the leg to the speed of the run and then decelerating
it to rest. Here we will use the physical pendulum as a model for the
swinging leg to compute this same quantity. We will assume that in running
the legs swing only at the hips. This model is, of course, not strictly correct
because in running the legs swing not only at the hips but also at the knees.
We will now outline a method for calculating the energy expended in swinging
the legs.
During each step of the run, the leg is accelerated to a maximum angular
velocity ωmax. In our pendulum model, this maximum angular velocity is
reached as the foot swings past the vertical position 0 (see Fig. 4.6).
The rotational kinetic energy at this point is the energy provided by
the leg muscles in each step of the run. This maximum rotational energy Er
(see Appendix A) is
Er
1
2
Iω2
max
Section 4.7 Energy Expended in Running 55
FIGURE 4.6 Running.
Here I is the moment of inertia of the leg. The angular velocity ωmax is
obtained as follows. From the rate of running, we can compute the period
of oscillation T for the leg modeled as a pendulum. Using this value for the
period, we calculate from Eq. 4.12 the maximum linear velocity vmax of the
foot. The angular velocity (see Appendix A) is then
ωmax
vmax
where is the length of the leg. In computing the period T, we must note that
the number of steps per second each leg executes is one half of the total number
of steps per second. In Exercise 4-8, it is shown that, based on the physical
pendulum model for running, the amount of work done during each step
is 1.6 mv2. In Chapter 3, using different considerations, the amount of work
done during each step was obtained as mv2. Considering that both approaches
are approximate, the agreement is certainly acceptable.
In calculating the energy requirements of walking and running, we
assumed that the kinetic energy imparted to the leg is fully (frictionally) dissipated
as the motion of the limb is halted within each step cycle. In fact,
a significant part of the kinetic energy imparted to the limbs during each step
cycle is stored as potential energy and is converted to kinetic energy during the
following part of the gait cycle, as in the motion of an oscillating pendulum
56 Chapter 4 Angular Motion
or a vibrating spring. The assumption of full energy dissipation at each step
results in an overestimate of the energy requirements for walking and running.
This energy overestimate is balanced by the underestimate due to the
neglecting of movement of the center of mass up and down during walking
and running as is discussed in following Sections 4.8 and 4.9.
4.8 Alternate Perspectives on Walking and Running
In Sections 4.4 to 4.7 we presented relatively simple models of walking and
running. More detailed and accurate descriptions can be found in various
technical journals. (For a review of the field see the article by Novacheck,
Bibliography). However, the basic approach in the various methods of analysis
is similar in that the highly complex interactive musculoskeletal system
involved in walking and/or running is represented by a simplified structure
that is amenable to mathematical analysis.
In our treatment of walking and running we considered only the pendulumlike
motion of the legs. A more detailed treatment considers also the motion
of the center of mass. A way to model the center of mass motion in walking
is to consider the motion of the center of mass during the course of a step.
Consider the start of the step when both feet are on the ground with one foot
ahead of the other. At this point the center of mass is between the two feet
and is at its lowest position (see Fig. 4.7). We chose left to be the rear foot.
The step begins with the rear foot leaving the ground and swinging forward.
The center of mass is at its highest point when the swinging foot is in line
with the stationary foot. As the swinging foot passes the stationary foot, it
becomes the forward foot and the step is completed with the two feet once
again on the ground with the right foot now in the rear. The center of mass
trajectory is an arc as depicted in Fig. 4.7.
In the sequence of the step described in the figure, the center of mass is
alternately behind and then in front of the point of the single-foot contact with
the ground as the free leg swings forward. That is, when the rear left foot
starts swinging forward, it is of course off the ground, and the center of mass is
behind the supporting right foot. During this part of the step the center of mass
is swinging toward the stationary right foot and its kinetic energy is converted
to potential energy (as in the upward swing of a pendulum; the supporting
foot being the fulcrum). After the left foot passes the stationary right foot, the
center of mass shifts forward of the right foot and accelerates as the potential
energy is converted to kinetic energy (downward swing of the pendulum).
With a step-length of 90 cm and the center of mass (with feet together) 1 m
above the ground, the center of mass is raised 11 cm during each swinging
Section 4.8 Alternate Perspectives on Walking and Running 57
L R
θ
L
C.M
FIGURE 4.7 Center of mass trajectory in the course of one step.
cycle of the step (see Exercise 4-10). This is an upper limit because in this
simplified treatment it is assumed that the legs remain straight throughout
the step.
For the above case, the work done in raising the center of mass by 0.11 m
is 0.11 m � W (in units of Joule) during each step. Because the body in the
process of walking is not a perfect pendulum, only part of this potential energy
is converted back into kinetic energy. To reduce the energy expenditure, the
body seeks adjustments to minimize the up-and-down movement of the center
of mass (see Section 4.9).
A clear distinction is made between walking and running. During walking,
at one point in the step cycle, both feet are in contact with the ground. In
running there is an interval during the step when both feet are off the ground.
During a walking step the center of mass trajectory is similar to that of an
inverted swinging pendulum with the fulcrum at the point where the two feet
pass one another (Fig. 4.7). Running can be compared to a person on a pogo
stick as if bouncing from one leg to another.
In Fig. 4.8 we show the energy required per meter of distance covered as
a function of speed, for walking and for running. (This figure is based on
the work of Alexander as presented by Novacheck.) The minimum energy
consumed per distance traveled (as shown in Fig. 4.8) occurs at a walking
speed of about 1.3 m/sec (2.9 mile/h). This is only 13% higher than the
most effortless walking speed of 1.13 m/sec we calculated on the basis of
the swinging-leg pendulum model in Section 4.6. This close agreement is
58 Chapter 4 Angular Motion
2
0
100
200
Walking
Running
Energycost
(joulespermeter)
300
400
4
Speed (meters per second)
6
FIGURE 4.8 Energy required per meter of distance covered as a function of speed,
for walking and for running. From T. F. Novacheck/Gait and Posture 7 (1998) 77–95.
somewhat fortuitous, brought about by canceling effect of the approximations
entailed in the calculation as discussed in Section 4.7. As shown in the figure,
the energy consumed per distance traveled increases at both lower and higher
walking speeds. For speeds typically less than 2 m/sec (∼4.5 mile/h) walking
is more efficient than running (that is requiring less energy per distance
traveled). Past this speed most people will spontaneously break into a run
consuming less energy.
As shown in Fig. 4.8 the energy cost of running is about 300 joules per
meter. As discussed in Section 3.8 (Exercise 3-9) the energy required to run
at 3 m/sec (9-min mile) was calculated to be 1350 J/sec, that is 450 joules
per meter. Considering the approximate nature of the calculations and the
difference in the methods, the agreement between the two numbers is again
remarkably good.
4.9 Carrying Loads
Carrying a load requires energy. Measurements have shown that for most
humans, as well animals such as dogs, horses and rats, the energy expended
at a given walking speed increases directly with the weight of the load
Chapter 4 Exercises 59
being carried. Specifically, carrying a load that is 50% of the body weight
increases the energy consumption by 50%. For most people this added energy
expenditure is the same whether they carry the load on their backs or on their
heads.
Recent studies have been focused on the extraordinary load carrying abilities
of women in certain areas of East Africa who can walk with relative ease
carrying large loads balanced on their heads. Quantitative measurements have
shown that women from the Luo and Kikuyu tribes can carry loads up to about
20% of their body weight without any measurable increase in their energy consumption.
Past this weight, the energy consumption increases in proportion
to the weight carried minus the 20%. That is, carrying a load 50% of the
body weight increases their energy consumption by 30% (50%–20%). The
experimenters of Heglund et al. suggest that the high load-carrying efficiency
of these women is due to greater efficiency of exchange from gravitational
potential energy to kinetic energy in the pendulum-like movement of the center
of mass discussed in Section 4.8. In other words their walking more closely
approximates a swinging pendulum. What specific aspect of the movement
or training that brings about these enhanced load carrying abilities is not yet
understood.
EXERCISES
Some of the problems in this chapter require a knowledge of the weight of
human limbs. Use Table 4.1 to compute these weights.
4-1. Explain why a runner is subject to a torque if she rounds a curve maintaining
a vertical position.
TABLE 4.1 Fraction of Body Weight for
Various Parts of the Body
Fraction of body weight
Head and neck 0.07
Trunk 0.43
Upper arms 0.07
Forearms and hands 0.06
Thighs 0.23
Legs and feet 0.14
Total 1.00
From Cooper and Glassow [6-6], p. 174.
60 Chapter 4 Angular Motion
4-2. In the act of walking, the arms swing back and forth through an angle
of 45◦ each second. Using the following data, and the data in Table 4.1,
calculate the average force on the shoulder due to the centrifugal force.
The mass of the person is 70 kg, and the length of the arm is 90 cm.
Assume that the total mass of the arms is located at the midpoint
of the arm.
4-3. Consider the carnival ride in which the riders stand against the wall
inside a large cylinder. As the cylinder rotates, the floor of the cylinder
drops and the passengers are pressed against the wall by the centrifugal
force. Assuming that the coefficient of friction between a rider and the
cylinder wall is 0.5 and that the radius of the cylinder is 5 m, what is
the minimum angular velocity and the corresponding linear velocity of
the cylinder that will hold the rider firmly against the wall?
4-4. If a person stands on a rotating pedestal with his arms loose, the arms
will rise toward a horizontal position. (a) Explain the reason for this
phenomenon. (b) Calculate the angular (rotational) velocity of the
pedestal for the angle of the arm to be at 60◦ with respect to the horizontal.
What is the corresponding number of revolutions per minute?
Assume that the length of the arm is 90 cm and the center of mass is at
mid-length.
4-5. Calculate the maximum velocity and acceleration of the foot of a runner
who does a 100-m dash in 10 sec. Assume that the length of a step is
1 m and that the length of the leg is 90 cm and the center of mass is at
mid-length.
4-6. What is the most effortless walking speed for a person with 90-cm-long
legs if the length of each step is 90 cm?
4-7. While walking, the arms swing under the force of gravity. Compute the
period of the swing. How does this period compare with the period of
the leg swing? Assume arm length of 90 cm.
4-8. Using the physical pendulum model for running described in the text,
derive an expression for the amount of work done during each step.
4-9. Compute the length of time for an erect human body without compensating
movements to hit the floor once it looses its balance. Assume
that the falling body behaves as a physical pendulum pivoted at the
floor with the period given by Eq. 4.13. The full length of the body
is 2 m.
4-10. Calculate the distance the center of mass is raised in the course of
one step with parameters and assumptions as discussed in Section 4.8.
(Refer to Fig. 4.7.)
Chapter 5
Elasticity and Strength
of Materials
So far we have considered the effect of forces only on the motion of a body.
We will now examine the effect of forces on the shape of the body. When a
force is applied to a body, the shape and size of the body change. Depending
on how the force is applied, the body may be stretched, compressed, bent, or
twisted. Elasticity is the property of a body that tends to return the body to its
original shape after the force is removed. If the applied force is sufficiently
large, however, the body is distorted beyond its elastic limit, and the original
shape is not restored after removal of the force. A still larger force will rupture
the body. We will review briefly the theory of deformation and then examine
the damaging effects of forces on bones and tissue.
5.1 Longitudinal Stretch and Compression
Let us consider the effect of a stretching force F applied to a bar (Fig. 5.1). The
applied force is transmitted to every part of the body, and it tends to pull the
material apart. This force, however, is resisted by the cohesive force that holds
the material together. The material breaks when the applied force exceeds the
cohesive force. If the force in Fig. 5.1 is reversed, the bar is compressed, and its
length is reduced. Similar considerations show that initially the compression
is elastic, but a sufficiently large force will produce permanent deformation
and then breakage.
61
62 Chapter 5 Elasticity and Strength of Materials
FIGURE 5.1 Stretching of a bar due to an applied force.
Stress S is the internal force per unit area acting on the material; it is
defined as1
S ≡
F
A
(5.1)
Here F is the applied force and A is the area on which the force is applied.
The force applied to the bar in Fig. 5.1 causes the bar to elongate by an
amount . The fractional change in length / is called the longitudinal
strain St; that is,
St ≡ (5.2)
Here is the length of the bar and is the change in the length due to the
applied force. If reversed, the force in Fig. 5.1 will compress the bar instead
of stretching it. (Stress and strain remain defined as before.) In 1676 Robert
Hooke observed that while the body remains elastic, the ratio of stress to strain
is constant (Hooke’s law); that is,
S
St
Y (5.3)
The constant of proportionality Y is called Young’s modulus. Young’s modulus
has been measured for many materials, some of which are listed in Table 5.1.
The breaking or rupture strength of these materials is also shown.
5.2 A Spring
A useful analogy can be drawn between a spring and the elastic properties of
a material. Consider the spring shown in Fig. 5.2.
1
The ≡ symbol is read “defined as.”
Section 5.2 A Spring 63
TABLE 5.1 Young’s Modulus and Rupture
Strength for Some Materials
Material
Young’s modulus
(dyn/cm2
)
Rupture strength
(dyn/cm2
)
Steel 200 � 1010
450 � 107
Aluminum 69 � 1010
62 � 107
Bone 14 � 1010
100 � 107
compression
83 � 107
stretch
27.5 � 107
twist
Tendon 68.9 � 107
stretch
Muscle 0.55 � 107
stretch
FIGURE 5.2 A stretched spring.
The force F required to stretch (or compress) the spring is directly
proportional to the amount of stretch; that is,
F K (5.4)
The constant of proportionality K is called the spring constant.
A stretched (or compressed) spring contains potential energy; that is, work
can be done by the stretched spring when the stretching force is removed. The
energy E stored in the spring (see [6-23]) is given by
E
1
2
K( )2
(5.5)
64 Chapter 5 Elasticity and Strength of Materials
An elastic body under stress is analogous to a spring with a spring constant
YA/ . This can be seen by expanding Eq. 5.3.
S
St
F/A
/
Y (5.6)
From Eq. 5.6, the force F is
F
YA
(5.7)
This equation is identical to the equation for a spring with a spring constant
K
YA
(5.8)
By analogy with the spring (see Eq. 5.5), the amount of energy stored in a
stretched or compressed body is
E
1
2
YA
( )2
(5.9)
5.3 Bone Fracture: Energy Considerations
Knowledge of the maximum energy that parts of the body can safely absorb
allows us to estimate the possibility of injury under various circumstances. We
shall first calculate the amount of energy required to break a bone of area A
and length . Assume that the bone remains elastic until fracture. Let us designate
the breaking stress of the bone as SB (see Fig. 5.3). The corresponding
force FB that will fracture the bone is, from Eq. 5.7,
FB SBA
YA
(5.10)
The compression at the breaking point is, therefore,
SB
Y
(5.11)
From Eq. 5.9, the energy stored in the compressed bone at the point of fracture is
E
1
2
YA
( )2
(5.12)
Substituting for SB /Y, we obtain
E
1
2
A S2
B
Y
(5.13)
Section 5.3 Bone Fracture: Energy Considerations 65
FIGURE 5.3 Compression of a bone.
As an example, consider the fracture of two leg bones that have a combined
length of about 90 cm and an average area of about 6 cm2. From Table 5.1,
the breaking stress SB is 109 dyn/cm2, and Young’s modulus for the bone is
14 � 1010 dyn/cm2. The total energy absorbed by the bones of one leg at the
point of compressive fracture is, from Eq. 5.13,
E
1
2
6 � 90 � 1018
14 � 1010
19.25 � 108
erg 192.5 J
The combined energy in the two legs is twice this value, or 385 J. This is the
amount of energy in the impact of a 70-kg person jumping from a height of
56 cm (1.8 ft), given by the product mgh. (Here m is the mass of the person,
g is the gravitational acceleration, and h is the height.) If all this energy is
absorbed by the leg bones, they may fracture.
It is certainly possible to jump safely from a height considerably greater
than 56 cm if, on landing, the joints of the body bend and the energy of the fall
is redistributed to reduce the chance of fracture. The calculation does however
point out the possibility of injury in a fall from even a small height. Similar
66 Chapter 5 Elasticity and Strength of Materials
considerations can be used to calculate the possibility of bone fracture in
running (see Exercise 5-1).
5.4 Impulsive Forces
In a sudden collision, a large force is exerted for a short period of time on
the colliding object. The general characteristic of such a collision force as a
function of time is shown in Fig. 5.4. The force starts at zero, increases to
some maximum value, and then decreases to zero again. The time interval
t2 − t1 t during which the force acts on the body is the duration of the
collision. Such a short-duration force is called an impulsive force.
Because the collision takes place in a short period of time, it is usually
difficult to determine the exact magnitude of the force during the collision.
However, it is relatively easy to calculate the average value of the impulsive
force Fav. It can be obtained simply from the relationship between force and
momentum given in Appendix A; that is,
Fav
mvf − mvi
t
(5.14)
Here mvi is the initial momentum of the object and mvf is the final momentum
after the collision. For example, if the duration of a collision is 6 � 10−3 sec
FIGURE 5.4 Impulsive force.
Section 5.5 Fracture Due to a Fall: Impulsive Force Considerations 67
and the change in momentum is 2 kg m/sec, the average force that acted during
the collision is
Fav
2 kg m/sec
6 � 10−3 sec
3.3 � 102
N
Note that, for a given momentum change, the magnitude of the impulsive
force is inversely proportional to the collision time; that is, the collision force
is larger in a fast collision than in a slower collision.
5.5 Fracture Due to a Fall: Impulsive Force
Considerations
In the preceding section, we calculated the injurious effects of collisions from
energy considerations. Similar calculations can be performed using the concept
of impulsive force. The magnitude of the force that causes the damage
is computed from Eq. 5.14. The change in momentum due to the collision
is usually easy to calculate, but the duration of the collision t is difficult
to determine precisely. It depends on the type of collision. If the colliding
objects are hard, the collision time is very short, a few milliseconds. If one of
the objects is soft and yields during the collision, the duration of the collision
is lengthened, and as a result the impulsive force is reduced. Thus, falling into
soft sand is less damaging than falling on a hard concrete surface.
When a person falls from a height h, his/her velocity on impact with the
ground, neglecting air friction (see Eq. 3.6), is
v 2gh (5.15)
The momentum on impact is
mv m 2gh W
2h
g
(5.16)
After the impact the body is at rest, and its momentum is therefore zero
(mvf 0). The change in momentum is
mvi − mvf W
2h
g
(5.17)
The average impact force, from Eq. 5.14, is
F
W
t
2h
g
m
t
2gh (5.18)
68 Chapter 5 Elasticity and Strength of Materials
Now comes the difficult part of the problem: Estimate of the collision
duration. If the impact surface is hard, such as concrete, and if the person
falls with his/her joints rigidly locked, the collision time is estimated to be
about 10−2 sec. The collision time is considerably longer if the person bends
his/her knees or falls on a soft surface.
From Table 5.1, the force per unit area that may cause a bone fracture is
109 dyn/cm2. If the person falls flat on his/her heels, the area of impact may
be about 2 cm2. Therefore, the force FB that will cause fracture is
FB 2 cm2
� 109
dyn/cm2
2 � 109
dyn (4.3 � 103
lb)
From Eq. 5.18, the height h of fall that will produce such an impulsive force
is given by
h
1
2g
F t
m
2
(5.19)
For a man with a mass of 70 kg, the height of the jump that will generate
a fracturing average impact force (assuming t 10−2 sec) is given by
h
1
2g
F t
m
2
1
2 � 980
2 � 109 � 10−2
70 � 103
41.6 cm (1.37 ft)
This is close to the result that we obtained from energy considerations. Note,
however, that the assumption of a 2-cm2 impact area is reasonable but somewhat
arbitrary. The area may be smaller or larger depending on the nature
of the landing; furthermore, we have assumed that the person lands with legs
rigidly straight. Exercises 5-2 and 5-3 provide further examples of calculating
the injurious effect of impulsive forces.
5.6 Airbags: Inflating Collision Protection Devices
The impact force may also be calculated from the distance the center of mass
of the body travels during the collision under the action of the impulsive
force. This is illustrated by examining the inflatable safety device used in
automobiles (see Fig. 5.5). An inflatable bag is located in the dashboard of
the car. In a collision, the bag expands suddenly and cushions the impact of
the passenger. The forward motion of the passenger must be stopped in about
30 cm of motion if contact with the hard surfaces of the car is to be avoided.
The average deceleration (see Eq. 3.6) is given by
a
v2
2s
(5.20)
Section 5.7 Whiplash Injury 69
FIGURE 5.5 Inflating collision protective device.
where v is the initial velocity of the automobile (and the passenger) and s is the
distance over which the deceleration occurs. The average force that produces
the deceleration is
F ma
mv2
2s
(5.21)
where m is the mass of the passenger.
For a 70-kg person with a 30-cm allowed stopping distance, the average
force is
F
70 � 103v2
2 � 30
1.17 � 103
� v2
dyn
At an impact velocity of 70 km/h (43.5 mph), the average stopping force
applied to the person is 4.45 � 106 dyn. If this force is uniformly distributed
over a 1000-cm2 area of the passenger’s body, the applied force per cm2 is
4.45 � 106 dyn. This is just below the estimated strength of body tissue.
The necessary stopping force increases as the square of the velocity. At
a 105-km impact speed, the average stopping force is 1010 dyn and the force
per cm2 is 107 dyn. Such a force would probably injure the passenger.
In the design of this safety system, the possibility has been considered
that the bag may be triggered during normal driving. If the bag were to remain
expanded, it would impede the ability of the driver to control the vehicle;
therefore, the bag is designed to remain expanded for only the short time
necessary to cushion the collision. (For an estimate of this period, see
Exercise 5-4.)
5.7 Whiplash Injury
Neck bones are rather delicate and can be fractured by even a moderate
force. Fortunately the neck muscles are relatively strong and are capable of
70 Chapter 5 Elasticity and Strength of Materials
FIGURE 5.6 Whiplash.
absorbing a considerable amount of energy. If, however, the impact is sudden,
as in a rear-end collision, the body is accelerated in the forward direction by
the back of the seat, and the unsupported neck is then suddenly yanked back
at full speed. Here the muscles do not respond fast enough and all the energy
is absorbed by the neck bones, causing the well-known whiplash injury (see
Fig. 5.6). The whiplash injury is described quantitatively in Exercise 5-5.
5.8 Falling from Great Height
There have been reports of people who jumped out of airplanes with
parachutes that failed to open and yet survived because they landed on soft
snow. It was found in these cases that the body made about a 1-m-deep
depression in the surface of the snow on impact. The credibility of these
reports can be verified by calculating the impact force that acts on the body
during the landing. It is shown in Exercise 5-6 that if the decelerating impact
force acts over a distance of about 1 m, the average value of this force remains
below the magnitude for serious injury even at the terminal falling velocity
of 62.5 m/sec (140 mph).
5.9 Osteoarthritis and Exercise
In the preceding sections of this chapter we discussed possible damaging
effects of large impulsive forces. In the normal course of daily activities our
bodies are subject mostly to smaller repetitive forces such as the impact of feet
with the ground in walking and running. A still not fully resolved question is
to what extent are such smaller repetitive forces particularly those encountered
in exercise and sport, damaging. Osteoarthritis is the commonly suspected
damage resulting from such repetitive impact.
Chapter 5 Exercises 71
Osteoarthritis is a joint disease characterized by a degenerative wearing
out of the components of the joint among them the synovial membrane and
cartilage tissue. As a result of such wear and tear the joint loses flexibility and
strength accompanied by pain and stiffness. Eventually the underlying bone
may also start eroding. Osteoarthritis is a major cause of disability at an older
age. Knees are the most commonly affected joint. After the age of 65, about
60% of men and 75% of women are to some extent affected by this condition.
Over the past several years a number of studies have been conducted to
determine the link between exercise and osteoarthritis. The emerging conclusion
is that joint injury is most strongly correlated with subsequent development
of osteoarthritis. Most likely this is the reason why people engaged in
high impact injury-prone sports are at a significantly greater risk of osteoarthritis.
Further, there appears to be little risk associated with recreational
running 20 to 40 km a week (∼13 to 25 miles).
It is not surprising that an injured joint is more likely to be subsequently
subject to wear and tear. As shown in Chapter 2, Table 2.1, the coefficient of
kinetic friction (μk) of an intact joint is about 0.003. The coefficient of friction
for un-lubricated bones is a hundred times higher. A joint injury usually compromises
to some extent the lubricating ability of the joint leading to increased
frictional wear and osteoarthritis. This simple picture would lead one to expect
that the progress of osteoarthritis would be more rapidly in the joints of people
who are regular runners than in a control group of non-runners. Yet this
does not appear to be the case. Osteoarthritis seems to progress at about the
same rate in both groups, indicating that the joints possess some ability to selfrepair.
These conclusions remain tentative and are subject to further study.
EXERCISES
5-1. Assume that a 50-kg runner trips and falls on his extended hand. If the
bones of one arm absorb all the kinetic energy (neglecting the energy
of the fall), what is the minimum speed of the runner that will cause a
fracture of the arm bone? Assume that the length of arm is 1 m and that
the area of the bone is 4 cm2.
5-2. Repeat the calculations in Exercise 5-1 using impulsive force considerations.
Assume that the duration of impact is 10−2 sec and the area of
impact is 4 cm2. Repeat the calculation with area of impact 1 cm2.
5-3. From what height can a 1-kg falling object cause fracture of the skull?
Assume that the object is hard, that the area of contact with the skull is
1 cm2, and that the duration of impact is 10−3 sec.
72 Chapter 5 Elasticity and Strength of Materials
5-4. Calculate the duration of the collision between the passenger and the
inflated bag of the collision protection device discussed in this chapter.
5-5. In a rear-end collision the automobile that is hit is accelerated to a velocity
v in 10−2/sec. What is the minimum velocity at which there is danger
of neck fracture from whiplash? Use the data provided in the text, and
assume that the area of the cervical vertebra is 1 cm2 and the mass of the
head is 5 kg.
5-6. Calculate the average decelerating impact force if a person falling with
a terminal velocity of 62.5 m/sec is decelerated to zero velocity over
a distance of 1 m. Assume that the person’s mass is 70 kg and that
she lands flat on her back so that the area of impact is 0.3 m2. Is this
force below the level for serious injury? (For body tissue, this is about
5 � 106 dyn/cm2.)
5-7. A boxer punches a 50-kg bag. Just as his fist hits the bag, it travels
at a speed of 7 m/sec. As a result of hitting the bag, his hand comes
to a complete stop. Assuming that the moving part of his hand weighs
5 kg, calculate the rebound velocity and kinetic energy of the bag. Is
kinetic energy conserved in this example? Why? (Use conservation of
momentum.)
Chapter 6
Insect Flight
In this chapter, we will analyze some aspects of insect flight. In particular, we
will consider the hovering flight of insects, using in our calculations many of
the concepts introduced in the previous chapters. The parameters required for
the computations were in most cases obtained from the literature, but some
had to be estimated because they were not readily available. The size, shape,
and mass of insects vary widely. We will perform our calculations for an insect
with a mass of 0.1 g, which is about the size of a bee.
In general, the flight of birds and insects is a complex phenomenon. A
complete discussion of flight would take into account aerodynamics as well
as the changing shape of the wings at the various stages of flight. Differences
in wing movements between large and small insects have only recently been
demonstrated. The following discussion is highly simplified but nevertheless
illustrates some of the basic physics of flight.
6.1 Hovering Flight
Many insects (and also some small birds) can beat their wings so rapidly that
they are able to hover in air over a fixed spot. The wing movements in a hovering
flight are complex. The wings are required to provide sideways stabilization
as well as the lifting force necessary to overcome the force of gravity.
The lifting force results from the downward stroke of the wings. As the wings
push down on the surrounding air, the resulting reaction force of the air on
the wings forces the insect up. The wings of most insects are designed so that
during the upward stroke the force on the wings is small. The lifting force
73
74 Chapter 6 Insect Flight
FIGURE 6.1 Force in flight.
acting on the wings during the wing movement is shown in Fig. 6.1. During
the upward movement of the wings, the gravitational force causes the insect
to drop. The downward wing movement then produces an upward force that
restores the insect to its original position. The vertical position of the insect
thus oscillates up and down at the frequency of the wingbeat.
The distance the insect falls between wingbeats depends on how rapidly its
wings are beating. If the insect flaps its wings at a slow rate, the time interval
during which the lifting force is zero is longer, and therefore the insect falls
farther than if its wings were beating rapidly.
We can easily compute the wingbeat frequency necessary for the insect to
maintain a given stability in its amplitude. To simplify the calculations, let us
assume that the lifting force is at a finite constant value while the wings are
moving down and that it is zero while the wings are moving up. During the
time interval t of the upward wingbeat, the insect drops a distance h under
the action of gravity. From Eq. 3.5, this distance is
h
g( t)2
2
(6.1)
The upward stroke then restores the insect to its original position. Typically,
it may be required that the vertical position of the insect change by no more
Section 6.2 Insect Wing Muscles 75
than 0.1 mm (i.e., h 0.1mm). The maximum allowable time for free fall
is then
t2 2h
g
1/2
2 � 10−2 cm
980 cm/sec2
4.5 � 10−3
sec
Since the up movements and the down movements of the wings are about
equal in duration, the period T for a complete up-and-down wing movement
is twice t; that is,
T 2 t 9 � 10−3
sec (6.2)
The frequency of wingbeats f, that is, the number of wingbeats per second, is
f
1
T
(6.3)
In our example this frequency is 110 wingbeats per second. This is a typical
insect wingbeat frequency, although some insects such as butterflies fly at
much lower frequency, about 10 wingbeats per second (they cannot hover),
and other small insects produce as many as 1000 wingbeats per second. To
restore the vertical position of the insect during the downward wing stroke,
the average upward force, Fav on the body of the insect must be equal to
twice the weight of the insect (see Exercise 6-1). Note that since the upward
force on the insect body is applied only for half the time, the average upward
force on the insect is simply its weight.
6.2 Insect Wing Muscles
A number of different wing-muscle arrangements occur in insects. One
arrangement, found in the dragonfly, is shown, highly simplified, in Fig. 6.2.
The wing movement is controlled by many muscles, which are here represented
by muscles A and B. The upward movement of the wings is produced
by the contraction of muscle A, which depresses the upper part of the thorax
and causes the attached wings to move up. While muscle A contracts, muscle
B is relaxed. Note that the force produced by muscle A is applied to the wing
by means of a Class 1 lever. The fulcrum here is the wing joint marked by
the small circle in Fig. 6.2.
The downward wing movement is produced by the contraction of muscle
B while muscle A is relaxed. Here the force is applied to the wings by means
of a Class 3 lever. In our calculations, we will assume that the length of the
wing is 1 cm.
The physical characteristics of insect flight muscles are not peculiar to
insects. The amount of force per unit area of the muscle and the rate of muscle
76 Chapter 6 Insect Flight
FIGURE 6.2 Wing muscles.
contraction are similar to the values measured for human muscles. Yet insect
wing muscles are required to flap the wings at a very high rate. This is made
possible by the lever arrangement of the wings. Measurements show that during
a wing swing of about 70◦, muscles A and B contract only about 2%.
Assuming that the length of muscle B is 3 mm, the change in length during
the muscle contraction is 0.06 mm (this is 2% of 3 mm). It can be shown that
under these conditions, muscle B must be attached to the wing 0.052 mm from
the fulcrum to achieve the required wing motion (see Exercise 6-2).
If the wingbeat frequency is 110 wingbeats per second, the period for one
up-and-down motion of the wings is 9 � 10−3 sec. The downward wing
movement produced by muscle B takes half this length of time, or 4.5 �
10−3 sec. Thus, the rate of contraction for muscle B is 0.06 mm divided by
4.5 � 10−3 sec, or 13 mm/sec. Such a rate of muscle contraction is commonly
observed in many types of muscle tissue.
6.3 Power Required for Hovering
We will now compute the power required to maintain hovering. Let us consider
again an insect with mass m 0.1 g. As is shown in Exercise 6-1, the
Section 6.3 Power Required for Hovering 77
average force, Fav, applied by the two wings during the downward stroke
is 2W. Because the pressure applied by the wings is uniformly distributed over
the total wing area, we can assume that the force generated by each wing acts
through a single point at the midsection of the wings. During the downward
stroke, the center of the wings traverses a vertical distance d (see Fig. 6.3).
The total work done by the insect during each downward stroke is the product
of force and distance; that is,
Work Fav � d 2Wd (6.4)
If the wings swing through an angle of 70◦, then in our case for the insect with
1-cm-long wings d is 0.57 cm. Therefore, the work done during each stroke
by the two wings is
Work 2 � 0.1 � 980 � 0.57 112 erg
Let us now examine where this energy goes. In our example the mass of the
insect has to be raised 0.1 mm during each downstroke. The energy E required
for this task is
E mgh 0.1 � 980 � 10−2
0.98 erg (6.5)
This is a negligible fraction of the total energy expended. Clearly, most of the
energy is expended in other processes. A more detailed analysis of the problem
FIGURE 6.3 Insect wing motion.
78 Chapter 6 Insect Flight
shows that the work done by the wings is converted primarily into kinetic
energy of the air that is accelerated by the downward stroke of the wings.
Power is the amount of work done in 1 sec. Our insect makes 110 downward
strokes per second; therefore, its power output P is
P 112 erg � 110/sec 1.23 � 104
erg/sec 1.23 � 10−3
W (6.6)
6.4 Kinetic Energy of Wings in Flight
In our calculation of the power used in hovering, we have neglected the kinetic
energy of the moving wings. The wings of insects, light as they are, have a
finite mass; therefore, as they move they possess kinetic energy. Because the
wings are in rotary motion, the maximum kinetic energy during each wing
stroke is
KE
1
2
Iω2
max (6.7)
Here I is the moment of inertia of the wing and ωmax is the maximum angular
velocity during the wing stroke. To obtain the moment of inertia for the wing,
we will assume that the wing can be approximated by a thin rod pivoted at one
end. The moment of inertia for the wing is then
I
m 3
3
(6.8)
where is the length of the wing (1 cm in our case) and m is the mass of two
wings, which may be typically 10−3 g. The maximum angular velocity ωmax
can be calculated from the maximum linear velocity vmax at the center of the
wing
ωmax
vmax
/2
(6.9)
During each stroke the center of the wings moves with an average linear
velocity vav given by the distance d traversed by the center of the wing
divided by the duration t of the wing stroke. From our previous example,
d 0.57 cm and t 4.5 � 10−3 sec. Therefore,
vav
d
t
0.57
4.5 � 10−3
127 cm/sec (6.10)
The velocity of the wings is zero both at the beginning and at the end of
the wing stroke. Therefore, the maximum linear velocity is higher than the
average velocity. If we assume that the velocity varies sinusoidally along the
Section 6.5 Elasticity of Wings 79
wing path, the maximum velocity is twice as high as the average velocity.
Therefore, the maximum angular velocity is
ωmax
254
/2
The kinetic energy is
KE
1
2
Iω2
max
1
2
10−3
2
3
254
/2
2
43 erg
Since there are two wing strokes (up and down) in each cycle of the wing
movement, the kinetic energy is 2 � 43 86 erg. This is about as much
energy as is consumed in hovering itself.
6.5 Elasticity of Wings
As the wings are accelerated, they gain kinetic energy, which is of course
provided by the muscles. When the wings are decelerated toward the end of
the stroke, this energy must be dissipated. During the downstroke, the kinetic
energy is dissipated by the muscles themselves and is converted into heat.
(This heat is used to maintain the required body temperature of the insect.)
Some insects are able to utilize the kinetic energy in the upward movement of
the wings to aid in their flight. The wing joints of these insects contain a pad
of elastic, rubberlike protein called resilin (Fig. 6.4). During the upstroke of
the wing, the resilin is stretched. The kinetic energy of the wing is converted
into potential energy in the stretched resilin, which stores the energy much
like a spring. When the wing moves down, this energy is released and aids in
the downstroke.
Using a few simplifying assumptions, we can calculate the amount of
energy stored in the stretched resilin. Although the resilin is bent into a complex
shape, we will assume in our calculation that it is a straight rod of area A
and length . Furthermore, we will assume that throughout the stretch the
resilin obeys Hooke’s law. This is not strictly true as the resilin is stretched
by a considerable amount and therefore both the area and Young’s modulus
change in the process of stretching.
The energy E stored in the stretched resilin is, from Eq. 5.9,
E
1
2
YA 2
(6.11)
Here Y is the Young’s modulus for resilin, which has been measured to be
1.8 � 107 dyn/cm2.
80 Chapter 6 Insect Flight
FIGURE 6.4 Resilin in the wing.
Typically, in an insect the size of a bee the volume of the resilin may be
equivalent to a cylinder 2 � 10−2 cm long and 4 � 10−4 cm2 in area. We will
assume that the length of the resilin rod is increased by 50% when stretched.
That is, is 10−2 cm. Therefore in our case the energy stored in the resilin
of each wing is
E
1
2
1.8 � 107 � 4 � 10−4 � 10−4
2 � 10−2
18 erg
The stored energy in the two wings is 36 erg, which is comparable to the
kinetic energy in the upstroke of the wings. Experiments show that as much
as 80% of the kinetic energy of the wing may be stored in the resilin. The utilization
of resilin is not restricted to wings. The hind legs of the flea, for example,
also contain resilin, which stores energy for jumping (see Exercise 6-3).
A further application of energy storage in resilin is examined in Exercise 6-4.
EXERCISES
6-1. Compute the force on the body of the insect that must be generated
during the downward wing stroke to keep the insect hovering.
Chapter 6 Exercises 81
6-2. Referring to the discussion in the text, compute the point of attachment
to the wing of muscle B in Fig. 6.2. Assume that the muscle is perpendicular
to the wing throughout the wing motion.
6-3. Assume that the shape of the resilin in each leg of the flea is equivalent
to a cylinder 2 � 10−2 cm long and 10−4 cm2 in area. If the change in
the length of the resilin is 10−2 cm, calculate the energy stored in
the resilin. The flea weighs 0.5 � 10−3 g. How high can the flea jump
utilizing only the stored energy?
6-4. Suppose that a 50 kg person were equipped with resilin pads in her
joints. How large would these pads have to be in order for them to store
enough energy for a 1
2 m jump? Assume that the pad is cubic in shape
and 1
2 .
Chapter 7
Fluids
In the previous chapters, we have examined the behavior of solids under the
action of forces. In the next three chapters, we will discuss the behavior
of liquids and gases, both of which play an important role in the life sciences.
The differences in the physical properties of solids, liquids, and gases
are explained in terms of the forces that bind the molecules. In a solid, the
molecules are rigidly bound; a solid therefore has a definite shape and volume.
The molecules constituting a liquid are not bound together with sufficient
force to maintain a definite shape, but the binding is sufficiently strong
to maintain a definite volume. A liquid adapts its shape to the vessel in which
it is contained. In a gas, the molecules are not bound to each other. Therefore
a gas has neither a definite shape nor a definite volume—it completely fills
the vessel in which it is contained. Both gases and liquids are free to flow and
are called fluids. Fluids and solids are governed by the same laws of mechanics,
but, because of their ability to flow, fluids exhibit some phenomena not
found in solid matter. In this chapter we will illustrate the properties of fluid
pressure, buoyant force in liquids, and surface tension with examples from
biology and zoology.
7.1 Force and Pressure in a Fluid
Solids and fluids transmit forces differently. When a force is applied to one
section of a solid, this force is transmitted to the other parts of the solid with
its direction unchanged. Because of a fluid’s ability to flow, it transmits a force
uniformly in all directions. Therefore, the pressure at any point in a fluid at
82
Section 7.2 Pascal’s Principle 83
rest is the same in all directions. The force exerted by a fluid at rest on any
area is perpendicular to the area. A fluid in a container exerts a force on all
parts of the container in contact with the fluid. A fluid also exerts a force on
any object immersed in it.
The pressure in a fluid increases with depth because of the weight of
the fluid above. In a fluid of constant density ρ, the difference in pressure,
P2 − P1, between two points separated by a vertical distance h is
P2 − P1 ρgh (7.1)
Fluid pressure is often measured in millimeters of mercury, or torr [after
Evangelista Torricelli (1608–1674), the first person to understand the nature
of atmospheric pressure]. One torr is the pressure exerted by a column of mercury
that is 1 mm high. Pascal, abbreviated as Pa is another commonly used
unit of pressure. The relationship between the torr and several of the other
units used to measure pressure follows:
1 torr 1 mm Hg
13.5 mm water
1.33 � 103
dyn/cm2
1.32 � 10−3
atm
1.93 � 10−2
psi
1.33 � 102
Pa (N/m2
)
(7.2)
7.2 Pascal’s Principle
When a force F1 is applied on a surface of a liquid that has an area A1, the
pressure in the liquid increases by an amount P (see Fig. 7.1), given by
P
F1
A1
(7.3)
In an incompressible liquid, the increase in the pressure at any point is transmitted
undiminished to all other points in the liquid. This is known as Pascal’s
principle. Because the pressure throughout the fluid is the same, the force F2
acting on the area A2 in Fig. 7.1 is
F2 PA2
A2
A1
F1 (7.4)
The ratio A2/A1 is analogous to the mechanical advantage of a lever.
84 Chapter 7 Fluids
FIGURE 7.1 An illustration of Pascal’s principle.
7.3 Hydrostatic Skeleton
We showed in Chapter 1 that muscles produce movement by pulling on the
bones of the skeleton. There are, however, soft-bodied animals (such as the
sea anemone and the earthworm) that lack a firm skeleton. Many of these
animals utilize Pascal’s principle to produce body motion. The structure by
means of which this is done is called the hydrostatic skeleton.
For the purpose of understanding the movements of an animal such as a
worm, we can think of the animal as consisting of a closed elastic cylinder
filled with a liquid; the cylinder is its hydrostatic skeleton. The worm produces
its movements with the longitudinal and circular muscles running along
the walls of the cylinder (see Fig. 7.2). Because the volume of the liquid in
the cylinder is constant, contraction of the circular muscles makes the worm
thinner and longer. Contraction of the longitudinal muscles causes the animal
to become shorter and fatter. If the longitudinal muscles contract only on one
side, the animal bends toward the contracting side. By anchoring alternate
ends of its body to a surface and by producing sequential longitudinal and circular
contractions, the animal moves itself forward or backward. Longitudinal
contraction on one side changes the direction of motion.
Let us now calculate the hydrostatic forces inside a moving worm. Consider
a worm that has a radius r. Assume that the circular muscles running
around its circumference are uniformly distributed along the length of the
worm and that the effective area of the muscle per unit length of the worm
Section 7.3 Hydrostatic Skeleton 85
FIGURE 7.2 The hydrostatic skeleton.
is AM. As the circular muscles contract, they generate a force fM, which,
along each centimeter of the worm’s length, is
fM SAM (7.5)
Here S is the force produced per unit area of the muscle. (Note that fM is in
units of force per unit length.) This force produces a pressure inside the worm.
The magnitude of the pressure can be calculated with the aid of Fig. 7.3, which
shows a section of the worm. The length of the section is L. If we were to
cut this section in half lengthwise, as shown in Fig. 7.3, the force due to the
pressure inside the cylinder would tend to push the two halves apart. This
force is calculated as follows. The surface area A along the cut midsection is
A L � 2r (7.6)
Because fluid pressure always acts perpendicular to a given surface area, the
force FP that tends to split the cylinder is
FP P � A P � L � 2r (7.7)
Here P is the fluid pressure produced inside the worm by contraction of the
circular muscles.
In equilibrium, the force FP is balanced by the muscle forces acting along
the two edges of the imaginary cut. Therefore,
FP 2fML
86 Chapter 7 Fluids
FIGURE 7.3 Calculating pressure inside a worm.
or
P � L � 2r 2fML
and
P
fM
r
(7.8)
To make the calculations specific, let us assume that the radius of the worm r
is 0.4 cm, the area of the circular muscles per centimeter length of the worm
is AM 1.5 � 10−3 cm2, and S, the maximum force generated per unit area
of the muscle, is 7 � 106 dyn/cm2. (This is the value we used previously for
human muscles.) Therefore, the pressure inside the worm under maximum
contraction of the circular muscles is
P
fM
r
SAM
r
7 � 106 � 1.5 � 10−3
0.4
2.63 � 104
dyn/cm2
19.8 torr
This is a relatively high pressure. It can raise a column of water to a height
of 26.7 cm. The force Ff in the forward direction generated by this pressure,
which stretches the worm, is
Ff P � πr2
1.32 � 104
dyn
The action of the longitudinal muscles can be similarly analyzed.
Section 7.5 Power Required to Remain Afloat 87
7.4 Archimedes’ Principle
Archimedes’ principle states that a body partially or wholly submerged in a
fluid is buoyed upward by a force that is equal in magnitude to the weight of
the displaced fluid. The derivation of this principle is found in basic physics
texts. We will now use Archimedes’ principle to calculate the power required
to remain afloat in water and to study the buoyancy of fish.
7.5 Power Required to Remain Afloat
Whether an animal sinks or floats in water depends on its density. If its density
is greater than that of water, the animal must perform work in order not to
sink. We will calculate the power P required for an animal of volume V and
density ρ to float with a fraction f of its volume submerged. This problem is
similar to the hovering flight we discussed in Chapter 6, but our approach to
the problem will be different.
Because a fraction f of the animal is submerged, the animal is buoyed up
by a force FB given by
FB gf Vρw (7.9)
where ρw is the density of water. The force FB is simply the weight of the
displaced water.
The net downward force FB on the animal is the difference between its
weight gVρ and the buoyant force; that is,
FD gVρ − gVfρw gV (ρ − fρw) (7.10)
To keep itself floating, the animal must produce an upward force equal to
FD. This force can be produced by pushing the limbs downward against the
water. This motion accelerates the water downward and results in the upward
reaction force that supports the animal.
If the area of the moving limbs is A and the final velocity of the accelerated
water is v, the mass of water accelerated per unit time in the treading motion
is given by (see Exercise 7-1)
m Avρw (7.11)
Because the water is initially stationary, the amount of momentum imparted to
the water each second is mv. (Remember that here m is the mass accelerated
per second.)
Momentum given to the water per second mv
88 Chapter 7 Fluids
This is the rate of change of momentum of the water. The force producing
this change in the momentum is applied to the water by the moving limbs.
The upward reaction force FR, which supports the weight of the swimmer,
is equal in magnitude to FD and is given by
FR FD gV (ρ − fρw) mv (7.12)
Substituting Eq. 7.11 for m, we obtain
ρwAv2
gV (ρ − fρw)
or
v
gV (ρ − fρw)
Aρw
(7.13)
The work done by the treading limbs goes into the kinetic energy of the accelerated
water. The kinetic energy given to the water each second is half the
product of the mass accelerated each second and the squared final velocity of
the water. This kinetic energy imparted to the water each second is the power
generated by the limbs; that is,
KE/sec Power generated by the limbs, P
1
2
mv2
Substituting equations for m and v, we obtain (see Exercise 7-1)
P
1
2
W 1 −
fρw
ρ
3
Aρw
(7.14)
Here W is the weight of the animal (W gVρ).
It is shown in Exercise 7-2 that a 50-kg woman expends about 7.8 W to
keep her nose above water. Note that, in our calculation, we have neglected
the kinetic energy of the moving limbs. In Eq. 7.14 it is assumed that the
density of the animal is greater than the density of water. The reverse case is
examined in Exercise 7-3.
7.6 Buoyancy of Fish
The bodies of some fish contain porous bones or air-filled swim bladders that
decrease their average density and allow them to float in water without an
expenditure of energy. The body of the cuttlefish, for example, contains a
Section 7.7 Surface Tension 89
porous bone that has a density of 0.62 g/cm3. The rest of its body has a
density of 1.067 g/cm3. We can find the percentage of the body volume X
occupied by the porous bone that makes the average density of the fish be
the same as the density of sea water (1.026 g/cm3) by using the following
equation (see Exercise 7-4):
1.026
0.62X + (100 − X) 1.067
100
(7.15)
In this case X 9.2%.
The cuttlefish lives in the sea at a depth of about 150 m. At this depth, the
pressure is 15 atm (see Exercise 7-5). The spaces in the porous bone are filled
with gas at a pressure of about 1 atm. Therefore, the porous bone must be able
to withstand a pressure of 14 atm. Experiments have shown that the bone can
in fact survive pressures up to 24 atm.
In fish that possess swim bladders, the decrease in density is provided by
the gas in the bladder. Because the density of the gas is negligible compared
to the density of tissue, the volume of the swim bladder required to reduce
the density of the fish is smaller than that of the porous bone. For example,
to achieve the density reduction calculated in the preceding example, the
volume of the bladder is only about 4% of the total volume of the fish (see
Exercise 7-6).
Fish possessing porous bones or swim bladders can alter their density.
The cuttlefish alters its density by injecting or withdrawing fluid from its
porous bone. Fish with swim bladders alter their density by changing the
amount of gas in the bladder. Another application of buoyancy is examined
in Exercise 7-7.
7.7 Surface Tension
The molecules constituting a liquid exert attractive forces on each other.
A molecule in the interior of the liquid is surrounded by an equal number
of neighboring molecules in all directions. Therefore, the net resultant intermolecular
force on an interior molecule is zero. The situation is different,
however, near the surface of the liquid. Because there are no molecules
above the surface, a molecule here is pulled predominantly in one direction,
toward the interior of the surface. This causes the surface of a liquid to contract
and behave somewhat like a stretched membrane. This contracting tendency
results in a surface tension that resists an increase in the free surface of
the liquid. It can be shown (see reference [7-7]) that surface tension is a
force acting tangential to the surface, normal to a line of unit length on the
surface (Fig. 7.4). The surface tension T of water at 25◦C is 72.8 dyn/cm. The
90 Chapter 7 Fluids
FIGURE 7.4 Surface tension.
total force FT produced by surface tension tangential to a liquid surface of
boundary length L is
FT TL (7.16)
When a liquid is contained in a vessel, the surface molecules near the wall
are attracted to the wall. This attractive force is called adhesion. At the same
time, however, these molecules are also subject to the attractive cohesive force
exerted by the liquid, which pulls the molecules in the opposite direction.
If the adhesive force is greater than the cohesive force, the liquid wets the
container wall, and the liquid surface near the wall is curved upward. If the
opposite is the case, the liquid surface is curved downward (see Fig. 7.5). The
angle θ in Fig. 7.5 is the angle between the wall and the tangent to the liquid
surface at the point of contact with the wall. For a given liquid and surface
material, θ is a well-defined constant. For example, the contact angle between
glass and water is 25◦.
If the adhesion is greater than the cohesion, a liquid in a narrow tube will
rise to a specific height h (see Fig. 7.6a), which can be calculated from the following
considerations. The weight W of the column of the supported liquid is
W π R2
hρg (7.17)
where R is the radius of the column and ρ is the density of the liquid. The maximum
force Fm due to the surface tension along the periphery of the liquid is
Fm 2π RT (7.18)
The upward component of this force supports the weight of the column of
liquid (see Fig. 7.6a); that is,
2π RT cos θ π R2
hρg (7.19)
Section 7.7 Surface Tension 91
FIGURE 7.5 Angle of contact when (a) liquid wets the wall and (b) liquid does not
wet the wall.
FIGURE 7.6 (a) Capillary rise. (b) Capillary depression.
Therefore, the height of the column is
h
2T cos θ
Rρg
(7.20)
If the adhesion is smaller than the cohesion, the angle θ is greater than 90◦.
In this case, the height of the fluid in the tube is depressed (Fig. 7.6b). Equation
7.20 still applies, yielding a negative number for h. These effects are
called capillary action.
Another consequence of surface tension is the tendency of liquid to assume
a spherical shape. This tendency is most clearly observed in a liquid outside
a container. Such an uncontained liquid forms into a sphere that can be noted
in the shape of raindrops. The pressure inside the spherical liquid drop is
92 Chapter 7 Fluids
higher than the pressure outside. The excess pressure P in a liquid sphere of
radius R is
P
2T
R
(7.21)
This is also the expression for the excess pressure inside an air bubble in a
liquid. In other words, to create gas bubble of radius R in a liquid with surface
tension T, the pressure of the gas injected into the liquid must be greater than
the pressure of the surrounding liquid by P as given in Eq. 7.21.
As will be shown in the following sections, the effects of surface tension
are evident in many areas relevant to the life sciences.
7.8 Soil Water
Most soil is porous with narrow spaces between the small particles. These
spaces act as capillaries and in part govern the motion of water through the soil.
When water enters soil, it penetrates the spaces between the small particles
and adheres to them. If the water did not adhere to the particles, it would run
rapidly through the soil until it reached solid rock. Plant life would then be
severely restricted. Because of adhesion and the resulting capillary action, a
significant fraction of the water that enters the soil is retained by it. For a plant
to withdraw this water, the roots must apply a negative pressure, or suction, to
the moist soil. The required negative pressure may be quite high. For example,
if the effective capillary radius of the soil is 10−3 cm, the pressure required to
withdraw the water is 1.46 � 105 dyn/cm2, or 0.144 atm (see Exercise 7-8).
The pressure required to withdraw water from the soil is called the soil
moisture tension (SMT). The SMT depends on the grain size of the soil, its
moisture content, and the material composition of the soil. The SMT is an
important parameter in determining the quality of the soil. The higher the
SMT, the more difficult it is for the roots to withdraw the water necessary for
plant growth.
The dependence of the SMT on the grain size can be understood from the
following considerations. The spaces between the particles of soil increase
with the size of the grains. Because capillary action is inversely proportional
to the diameter of the capillary, finely grained soil will hold water more tightly
than soil of similar material with larger grains (see Fig. 7.7).
When all the pores of the soil are filled with water, the surface moisture
tension is at its lowest value. In other words, under these conditions the
required suction pressure produced by the plant roots to withdraw the water
from the soil is the lowest. Saturated soil, however, is not the best medium for
plant growth. The roots need some air, which is absent when the soil is fully
Section 7.9 Insect Locomotion on Water 93
FIGURE 7.7 Fine-grained soil (a) holds water more tightly than coarse-grained
soil (b).
saturated with water. As the amount of water in the soil decreases, the SMT
increases. In loam, for example, with a moisture content of 20% the SMT is
about 0.19 atm. When the moisture content drops to 12%, the SMT increases
to 0.76 atm.
The rise in SMT with decreasing moisture content can be explained in
part by two effects. As the soil loses moisture, the remaining water tends to
be bound into the narrower capillaries. Therefore the withdrawal of water
becomes more difficult. In addition, as the moisture content decreases, sections
of water become isolated and tend to form droplets. The size of these
droplets may be very small. If, for example, the radius of a droplet decreases
to 10−5 cm, the pressure required to draw the water out of the droplet is about
14.5 atm.
Capillary action also depends on the strength of adhesion, which in turn
depends on the material composition of the capillary surface. For example,
under similar conditions of grain size and moisture content, the SMT in clay
may be ten times higher than in loam. There is a limit to the pressure that roots
can produce in order to withdraw water from the soil. If the SMT increases
above 15 atm, wheat, for example, cannot obtain enough water to grow. In hot
dry climates where vegetation requires more water, plants may wilt even at an
SMT of 2 atm. The ability of a plant to survive depends not so much on the
water content as on the SMT of the soil. A plant may thrive in loam and yet
wilt in a clayey soil with twice the moisture content. Other aspects of SMT
are treated in Exercises 7-9 and 7-10.
7.9 Insect Locomotion on Water
About 3% of all insects are to some extent aquatic. In one way or another their
lives are associated with water. Many of these insects are adapted to utilize the
surface tension of water for locomotion. The surface tension of water makes
it possible for some insects to stand on water and remain dry. Let us now
94 Chapter 7 Fluids
FIGURE 7.8 Insect standing on water.
estimate the maximum weight of an insect that can be supported by surface
tension.
When the insect lands on water, the surface is depressed as shown in
Fig. 7.8. The legs of such an insect, however, must not be wetted by water.
A waxlike coating can provide the necessary water-repulsive property. The
weight W of the insect is supported by the upward component of the surface
tension; that is,
W LT sin θ (7.22)
where L is the combined circumference of all the insect legs in contact with
the water.
To perform quantitative calculations, we must introduce some assumptions.
We assume that the insect is in the shape of a cube with side dimensions .
The weight of the insect of density ρ is then
W 3
ρg (7.23)
Let us further assume that the circumference of the legs in contact with water
is approximately equal to the dimension of the cube; that is, from Eq. 7.23,
L
W
ρg
1/3
(7.24)
The greatest supporting force provided by surface tension occurs at the angle
θ 90◦ (see Fig. 7.8). (At this point the insect is on the verge of sinking.)
The maximum weight Wm that can be supported by surface tension is obtained
from Eq. 7.22; that is,
Wm LT
Wm
ρg
1/3
T
or
W
2/3
m
T
(ρg)1/3
(7.25)
Section 7.10 Contraction of Muscles 95
If the density of the insect is 1 g/cm3, then with T 72.8 dyn/cm, the
maximum weight is
W
2/3
m
72.8
(980)1/3
or
Wm 19.7 dyn
The mass of the insect is therefore about 2 � 10−2 g. The corresponding
linear size of such an insect is about 3 mm.
As is shown in Exercise 7-11, a 70 kg person would have to stand on a
platform about 10 km in perimeter to be supported solely by surface tension.
(This is a disk about 3.2 km in diameter.)
7.10 Contraction of Muscles
An examination of skeletal muscles shows that they consist of smaller muscle
fibers, which in turn are composed of yet smaller units called myofibrils.
Further, examination with an electron microscope reveals that the myofibril
is composed of two types of threads, one made of myosin, which is about
160 ˚A (1 ˚A 10−8 cm) in diameter, and the other made of actin, which has
a diameter of about 50 ˚A. Each myosin-actin unit is about 1 mm long. The
threads are aligned in a regular pattern with spaces between threads so that the
threads can slide past one another, as shown in Fig. 7.9.
FIGURE 7.9 Contraction of muscles.
96 Chapter 7 Fluids
Muscle contraction begins with an electrical nerve impulse that results in
a release of Ca2+ ions into the myosin-actin structure. The calcium ions in
turn produce conformational changes that result in the sliding of the threads
through each other, shortening the myosin-actin structure. The collective
effect of this process is the contraction of the muscle.
Clearly, a force must act along the myosin-actin threads to produce such a
contracting motion. The physical nature of this force is not fully understood.
It has been suggested by Gamow and Ycas [7-5] that this force may be due
to surface tension, which is present not only in liquids but also in jellylike
materials such as tissue cells. The motion of the threads is then similar to
capillary movement of a liquid. Here the movement is due to the attraction
between the surfaces of the two types of thread. The surface attraction may
be triggered by a release of the Ca2+ ions. Let us now estimate the force
per square centimeter of muscle tissue that could be generated by the surface
tension proposed in this model.
If the average diameter of the threads is D, the number of threads N per
square centimeter of muscle is approximately
N
1
π
4 � D2
(7.26)
The maximum pulling force Ff produced by the surface tension on each
fiber is, from Eq. 7.16,
Ff πDT (7.27)
The total maximum force Fm due to all the fibers in a 1-cm2 area of muscle is
Fm NFf
4T
D
(7.28)
The average diameter D of the muscle fibers is about 100 ˚A (10−6 cm). Therefore,
the maximum contracting force that can be produced by surface tension
per square centimeter of muscle area is
Fm T � 4 � 106
dyn/cm2
A surface tension of 1.75 dyn/cm can account for the 7 � 106 dyn/cm2 measured
force capability of muscles. Because this is well below surface tensions
commonly encountered, we can conclude that surface tension could be the
source of muscle contraction. This proposed mechanism, however, should
not be taken too seriously. The actual processes in muscle contraction are
much more complex and cannot be reduced to a simple surface tension model
(see [7-7 and 7-9]).
Section 7.11 Surfactants 97
7.11 Surfactants
Surfactants are molecules that lower surface tension of liquids. (The word is an
abbreviation of surface active agent.) The most common surfactant molecules
have one end that is water-soluble (hydrophilic) and the other end water insoluble
(hydrophobic) (see Fig. 7.10). As the word implies, the hydrophilic end
is strongly attracted to water while the hydrophobic has very little attraction
to water but is attracted and is readily soluble in oily liquids. Many different
types of surfactant molecules are found in nature or as products of laboratory
synthesis.
When surfactant molecules are placed in water, they align on the surface
with the hydrophobic end pushed out of the water as shown in Fig. 7.11. Such
an alignment disrupts the surface structure of water, reducing the surface tension.
A small concentration of surfactant molecules can typically reduce surface
tension of water from 73 dyn/cm to 30 dyn/cm. In oily liquids, surfactants
are aligned with the hydrophilic end squeezed out of the liquid. In this case
the surface tension of the oil is reduced.
Hydrophobe Hydrophile
FIGURE 7.10 Schematic of a surfactant molecule.
FIGURE 7.11 Surface layer of surfactant molecules.
98 Chapter 7 Fluids
(a)
(b)
(c)
FIGURE 7.12 Action of detergents. (a) Oil drop on a wet spot. (b) The hydrophobic
end of surfactant molecules enter the oil spot. (c) The oil spot break up into smaller
section surrounded by hydrophilic ends.
The most familiar use of surfactants is as soaps and detergents to wash
away oily substances. Here the hydrophobic end of the surfactants dissolves
into the oil surface while the hydrophilic end remains exposed to the surrounding
water as shown in Fig. 7.12. The aligned surfactant molecules reduce the
surface tension of the oil. As a result, the oil breaks up into small droplets
surrounded by the hydrophilic end of the surfactants. The small oil droplets
are solubilized (that is suspended or dissolved) in the water and can now be
washed away.
Surfactants are widely used in experimental biochemistry. In certain types
of experiments, for example, proteins that are hydrophobic such as membrane
proteins and lipoproteins must be dissolved in water. Here surfactants are used
to solubilize the proteins in a process similar to that illustrated in Fig. 7.12.
The hydrophobic ends of the surfactant molecules dissolve into the surface of
the protein. The aligned hydrophilic ends surround the protein, solubilizing it
in the ambient water.
Chapter 7 Exercises 99
Some insects such as the Microvelia not only stand on water but also
utilize surface tension for propulsion. They secrete a substance from their
abdomen that reduces the surface tension behind them. As a result they are
propelled in a forward direction. Here the effect is similar to cutting a taut
rubber membrane which then draws apart, each section moving away from the
cut. This effect known as Marangoni propulsion can be demonstrated simply
by coating one end of a toothpick with soap, and placing it in water. The
soap acting as the surfactant reduces the surface tension behind the coated
end resulting in the acceleration of the toothpick away from the dissolved
soap.
Experiments have shown that the surfactant excreted by insects reduces the
surface tension of water from 73 dyn/cm to about 50 dyn/cm. Measurements
show that during Marangoni propulsion, Microvelia can attain peak speeds of
17 cm/sec. This value is in agreement with a simple calculation presented in
Exercise 7-12.
The importance of surfactants in the process of breathing is described in
Chapter 9.
EXERCISES
7-1. Verify Eqs. 7.11 and 7.14.
7-2. With the nose above the water, about 95% of the body is submerged.
Calculate the power expended by a 50-kg woman treading water in this
position. Assume that the average density of the human body is about
the same as water (ρ ρw 1 g/cm3) and that the area A of the limbs
acting on the water is about 600 cm2.
7-3. In Eq. 7.14, it is assumed that the density of the animal is greater than the
density of the fluid in which it is submerged. If the situation is reversed,
the immersed animal tends to rise to the surface, and it must expend
energy to keep itself below the surface. How is Eq. 7.14 modified for
this case?
7-4. Derive the relationship shown in Eq. 7.15.
7-5. Calculate the pressure 150 m below the surface of the sea. The density
of sea water is 1.026 g/cm3.
7-6. Calculate volume of the swim bladder as a percent of the total volume
of the fish in order to reduce the average density of the fish from
1.067 g/cm3 to 1.026 g/cm3.
100 Chapter 7 Fluids
7-7. The density of an animal is conveniently obtained by weighing it first
in air and then immersed in a fluid. Let the weight in air and in the fluid
be respectively W1 and W2. If the density of the fluid is ρ1, the average
density ρ2 of the animal is
ρ2 ρ1
W1
W1 − W2
Derive this relationship.
7-8. Starting with Eq. 7.20, show that the pressure P required to withdraw
the water from a capillary of radius R and contact angle θ is
P
2T cos θ
R
With the contact angle θ 0◦, determine the pressure required to withdraw
water from a capillary with a 10−3 cm radius. Assume that the
surface tension T 72.8 dyn/cm.
7-9. If a section of coarse-grained soil is adjacent to a finer grained soil of
the same material, water will seep from the coarse-grained to the finer
grained soil. Explain the reason for this.
7-10. Design an instrument to measure the SMT. (You can find a description
of one such device in [7-4].)
7-11. Calculate the perimeter of a platform required to support a 70 kg person
solely by surface tension.
7-12. (a) Estimate the maximum acceleration of the insect that can be produced
by reducing the surface tension as described in the text. Assume
that the linear dimension of the insect is 3 � 10−1 cm and its mass is
3 � 10−2 g. Further, assume that the surface tension difference between
the clean water and surfactant altered water provides the force to accelerate
the insect. Use surface tension values provided in the text.
(b) Calculate the speed of the insect assuming that the surfactant
release lasts 0.5 sec.
Chapter 8
The Motion of Fluids
The study of fluids in motion is closely related to biology and medicine. In
fact, one of the foremost workers in this field, L. M. Poiseuille (1799–1869),
was a French physician whose study of moving fluids was motivated by his
interest in the flow of blood through the body. In this chapter, we will review
briefly the principles governing the flow of fluids and then examine the flow
of blood in the circulatory system.
8.1 Bernoulli’s Equation
If frictional losses are neglected, the flow of an incompressible fluid is governed
by Bernoulli’s equation, which gives the relationship between velocity,
pressure, and elevation in a line of flow. Bernoulli’s equation states that at any
point in the channel of a flowing fluid the following relationship holds:
P + ρgh +
1
2
ρv2
Constant (8.1)
Here P is the pressure in the fluid, h is the height, ρ is the density, and v is
the velocity at any point in the flow channel. The first term in the equation
is the potential energy per unit volume of the fluid due to the pressure in the
fluid. (Note that the unit for pressure, which is dyn/cm2, is identical to erg/cm3,
which is energy per unit volume.) The second term is the gravitational potential
energy per unit volume, and the third is the kinetic energy per unit volume.
Bernoulli’s equation follows from the law of energy conservation. Because
the three terms in the equation represent the total energy in the fluid, in the
101
102 Chapter 8 The Motion of Fluids
FIGURE 8.1 Flow of fluid through a pipe with two segments of different areas.
absence of friction their sum must remain constant no matter how the flow is
altered.
We will illustrate the use of Bernoulli’s equation with a simple example.
Consider a fluid flowing through a pipe consisting of two segments with crosssectional
areas A1 and A2, respectively (see Fig. 8.1). The volume of fluid
flowing per second past any point in the pipe is given by the product of the
fluid velocity and the area of the pipe, A � v. If the fluid is incompressible, in
a unit time as much fluid must flow out of the pipe as flows into it. Therefore,
the rates of flow in segments 1 and 2 are equal; that is,
A1v1 A2v2 or v2
A1
A2
v1 (8.2)
In our case A1 is larger than A2 so we conclude that the velocity of the fluid
in segment 2 is greater than in segment 1.
Bernoulli’s equation states that the sum of the terms in Eq. 8.1 at any point
in the flow is equal to the same constant. Therefore the relationship between
the parameters P, ρ, h, and v at points 1 and 2 is
P1 + ρgh1 +
1
2
ρv2
1 P2 + ρgh2 +
1
2
ρv2
2 (8.3)
where the subscripts designate the parameters at the two points in the flow.
Because in our case the two segments are at the same height (h1 h2), Eq. 8.2
can be written as
P1 +
1
2
ρv2
1 P2 +
1
2
ρv2
2 (8.4)
Because v2 (A1/A2)v1, the pressure in segment 2 is
P2 P1 −
1
2
ρv2
1
A1
A2
2
− 1 (8.5)
This relationship shows that while the flow velocity in segment 2 increases,
the pressure in that segment decreases.
Section 8.2 Viscosity and Poiseuille’s Law 103
FIGURE 8.2 Laminar flow. The length of the arrows indicates the magnitude of the
velocity of the fluid.
8.2 Viscosity and Poiseuille’s Law
Frictionless flow is an idealization. In a real fluid, the molecules attract each
other; consequently, relative motion between the fluid molecules is opposed
by a frictional force, which is called viscous friction. Viscous friction is proportional
to the velocity of flow and to the coefficient of viscosity for the given
fluid. As a result of viscous friction, the velocity of a fluid flowing through a
pipe varies across the pipe. The velocity is highest at the center and decreases
toward the walls; at the walls of the pipe, the fluid is stationary. Such fluid
flow is called laminar. Figure 8.2 shows the velocity profile for laminar flow
in a pipe. The lengths of the arrows are proportional to the velocity across the
pipe diameter.
If viscosity is taken into account, it can be shown (see reference [8-5]) that
the rate of laminar flow Q through a cylindrical tube of radius R and length L
is given by Poiseuille’s law, which is
Q
πR4 (P1 − P2)
8ηL
cm3
/sec (8.6)
where P1 − P2 is the difference between the fluid pressures at the two ends
of the cylinder and η is the coefficient of viscosity measured in units of dyn
(sec/cm2), which is called a poise. The viscosities of some fluids are listed in
Table 8.1. In general, viscosity is a function of temperature and increases as
the fluid becomes colder.
There is a basic difference between frictionless and viscous fluid flow.
A frictionless fluid will flow steadily without an external force applied to it.
This fact is evident from Bernoulli’s equation, which shows that if the height
and velocity of the fluid remain constant, there is no pressure drop along the
flow path. But Poiseuille’s equation for viscous flow states that a pressure
104 Chapter 8 The Motion of Fluids
TABLE 8.1 Viscosities of
Selected Fluids
Temperature Viscosity
Fluid (◦C) (poise)
Water 20 0.01
Glycerin 20 8.3
Mercury 20 0.0155
Air 20 0.00018
Blood 37 0.04
drop always accompanies viscous fluid flow. By rearranging Eq. 8.6, we can
express the pressure drop as
P1 − P2
Q8ηL
πR4
(8.7)
The expression P1 − P2 is the pressure drop that accompanies the flow rate Q
along a length L of the pipe. The product of the pressure drop and the area
of the pipe is the force required to overcome the frictional forces that tend to
retard the flow in the pipe segment. Note that for a given flow rate the pressure
drop required to overcome frictional losses decreases as the fourth power of
the pipe radius. Thus, even though all fluids are subject to friction, if the area
of the flow is large, frictional losses and the accompanying pressure drop are
small and can be neglected. In these cases, Bernoulli’s equation may be used
with little error.
8.3 Turbulent Flow
If the velocity of a fluid is increased past a critical point, the smooth laminar
flow shown in Fig. 8.2 is disrupted. The flow becomes turbulent with eddies
and whirls disrupting the laminar flow (see Fig. 8.3). In a cylindrical pipe the
critical flow velocity vc above which the flow is turbulent, is given by
vc
η
ρD
(8.8)
Here D is the diameter of the cylinder, ρ is the density of the fluid, and η
is the viscosity. The symbol is the Reynold’s number, which for most fluids
has a value between 2000 and 3000. The frictional forces in turbulent flow are
greater than in laminar flow. Therefore, as the flow turns turbulent, it becomes
more difficult to force a fluid through a pipe.
Section 8.4 Circulation of the Blood 105
FIGURE 8.3 Turbulent fluid flow.
8.4 Circulation of the Blood
The circulation of blood through the body is often compared to a plumbing
system with the heart as the pump and the veins, arteries, and capillaries as
the pipes through which the blood flows. This analogy is not entirely correct.
Blood is not a simple fluid; it contains cells that complicate the flow, especially
when the passages become narrow. Furthermore, the veins and arteries are
not rigid pipes but are elastic and alter their shape in response to the forces
applied by the fluid. Still, it is possible to analyze the circulatory system with
reasonable accuracy using the concepts developed for simple fluids flowing in
rigid pipes.
Figure 8.4 is a drawing of the human circulatory system. The blood in the
circulatory system brings oxygen, nutrients, and various other vital substances
to the cells and removes the metabolic waste products from the cells. The
blood is pumped through the circulatory system by the heart, and it leaves the
heart through vessels called arteries and returns to it through veins.
The mammalian heart consists of two independent pumps, each made of
two chambers called the atrium and the ventricle. The entrances to and exits
from these chambers are controlled by valves that are arranged to maintain the
flow of blood in the proper direction. Blood from all parts of the body except
the lungs enters the right atrium, which contracts and forces the blood into the
right ventricle. The ventricle then contracts and drives the blood through the
pulmonary artery into the lungs. In its passage through the lungs, the blood
releases carbon dioxide and absorbs oxygen. The blood then flows into the
left atrium via the pulmonary vein. The contraction of the left atrium forces
the blood into the left ventricle, which on contraction drives the oxygen-rich
blood through the aorta into the arteries that lead to all parts of the body except
the lungs. Thus, the right side of the heart pumps the blood through the lungs,
and the left side pumps it through the rest of the body.
106 Chapter 8 The Motion of Fluids
FIGURE 8.4 Schematic diagram showing various routes of the circulation.
The large artery, called the aorta, which carries the oxygenated blood away
from the left chamber of the heart, branches into smaller arteries, which lead
to the various parts of the body. These in turn branch into still smaller arteries,
the smallest of which are called arterioles. As we will explain later, the arterioles
play an important role in regulating the blood flow to specific regions in
Section 8.5 Blood Pressure 107
the body. The arterioles branch further into narrow capillaries that are often
barely wide enough to allow the passage of single blood cells.
The capillaries are so profusely spread through the tissue that nearly all
the cells in the body are close to a capillary. The exchange of gases, nutrients,
and waste products between the blood and the surrounding tissue occurs by
diffusion through the thin capillary walls (see Chapter 9). The capillaries join
into tiny veins called venules, which in turn merge into larger and larger veins
that lead the oxygen-depleted blood back to the right atrium of the heart.
8.5 Blood Pressure
The contraction of the heart chambers is triggered by electrical pulses that
are applied simultaneously both to the left and to the right halves of the heart.
First the atria contract, forcing the blood into the ventricles; then the ventricles
contract, forcing the blood out of the heart. Because of the pumping action of
the heart, blood enters the arteries in spurts or pulses. The maximum pressure
driving the blood at the peak of the pulse is called the systolic pressure. The
lowest blood pressure between the pulses is called the diastolic pressure. In a
young healthy individual the systolic pressure is about 120 torr (mm Hg) and
the diastolic pressure is about 80 torr. Therefore the average pressure of the
pulsating blood at heart level is 100 torr.
As the blood flows through the circulatory system, its initial energy, provided
by the pumping action of the heart, is dissipated by two loss mechanisms:
losses associated with the expansion and contraction of the arterial
walls and viscous friction associated with the blood flow. Due to these energy
losses, the initial pressure fluctuations are smoothed out as the blood flows
away from the heart, and the average pressure drops. By the time the blood
reaches the capillaries, the flow is smooth and the blood pressure is only about
30 torr. The pressure drops still lower in the veins and is close to zero just
before returning to the heart. In this final stage of the flow, the movement of
blood through the veins is aided by the contraction of muscles that squeeze
the blood toward the heart. One-way flow is assured by unidirectional valves
in the veins.
The main arteries in the body have a relatively large radius. The radius
of the aorta, for example, is about 1 cm; therefore, the pressure drop along
the arteries is small. We can estimate this pressure drop using Poiseuille’s
law (Eq. 8.7). However, to solve the equation, we must know the rate of
blood flow. The rate of blood flow Q through the body depends on the level
of physical activity. At rest, the total flow rate is about 5 liter/min. During
intense activity the flow rate may rise to about 25 liter/min. Exercise 8-1
shows that at peak flow the pressure drop per centimeter length of the aorta
108 Chapter 8 The Motion of Fluids
FIGURE 8.5 Blood pressure in a reclining and in an erect person.
is only 42.5 dyn/cm2 (3.19 � 10−2 torr), which is negligible compared to the
total blood pressure.
Of course, as the aorta branches, the size of the arteries decreases, resulting
in an increased resistance to flow. Although the blood flow in the narrower
arteries is also reduced, the pressure drop is no longer negligible (see
Exercise 8-2). The average pressure at the entrance to the arterioles is about
90 torr. Still, this is only a 10% drop from the average pressure at the heart.
The flow through the arterioles is accompanied by a much larger pressure drop,
about 60 torr. As a result, the pressure at the capillaries is only about 30 torr.
Since the pressure drop in the main arteries is small, when the body is
horizontal, the average arterial pressure is approximately constant throughout
the body. The arterial blood pressure, which is on the average 100 torr, can
support a column of blood 129 cm high (see Eq. 7.1 and Exercise 8-3). This
means that if a small tube were introduced into the artery, the blood in it would
rise to a height of 129 cm (see Fig. 8.5).
If a person is standing erect, the blood pressure in the arteries is not uniform
in the various parts of the body. The weight of the blood must be taken
into account in calculating the pressure at various locations. For example, the
average pressure in the artery located in the head, 50 cm above the heart (see
Exercise 8-4a) is Phead Pheart − ρgh 61 torr. In the feet, 130 cm below
the heart, the arterial pressure is 200 torr (see Exercise 8-4b).
Section 8.6 Control of Blood Flow 109
The cardiovascular system has various flow-control mechanisms that can
compensate for the large arterial pressure changes that accompany shifts in
the position of the body. Still, it may take a few seconds for the system to
compensate. Thus, a person may feel momentarily dizzy as he/she jumps up
from a prone position. This is due to the sudden decrease in the blood pressure
of the brain arteries, which results in a temporary decrease of blood flow to
the brain.
The same hydrostatic factors operate also in the veins, and here their effect
may be more severe than in the arteries. The blood pressure in the veins
is lower than in the arteries. When a person stands motionless, the blood
pressure is barely adequate to force the blood from the feet back to the heart.
Thus when a person sits or stands without muscular movement, blood gathers
in the veins of the legs. This increases the pressure in the capillaries and may
cause temporary swelling of the legs.
8.6 Control of Blood Flow
The pumping action of the heart (that is, blood pressure, flow volume and rate
of heart beat) is regulated by a variety of hormones. Hormones are molecules,
often proteins, that are produced by organs and tissues in different parts of
the body. They are secreted into the blood stream and carry messages from
one part of the body to another. Hormones affecting the heart are produced in
response to stimuli such as need for more oxygen, changes in body temperature,
and various types of emotional stress.
The flow of blood to specific parts of the body is controlled by the arterioles.
These small vessels that receive blood from the arteries have an average
diameter of about 0.1 mm. The walls of the arterioles contain smooth muscle
fibers that contract when stimulated by nerve impulses and hormones. The contraction
of the arterioles in one part of the body reduces the blood flow to that
region and diverts it to another. Since the radius of the arterioles is small, constriction
is an effective method for controlling blood flow. Poiseuille’s equation
shows that if the pressure drop remains constant, a 20% decrease in the radius
reduces the blood flow by more than a factor of 2 (see Exercise 8-5).
A stress-induced heart condition called stress cardiomyopathy (broken
heart syndrome) has only recently been clearly identified by Western medicine.
The syndrome occurs most frequently after a sudden intense emotional trauma
such as death in the family, an experience of violence, or extreme anger. The
symptoms are similar to an acute heart attack, but the coronary arteries are
found to be normal and the heart tissue is not damaged. It has suggested that
the condition is triggered by an excessive release of stress-related hormones
called chatecholamines.
110 Chapter 8 The Motion of Fluids
8.7 Energetics of Blood Flow
For an individual at rest, the rate of blood flow is about 5 liter/min. This
implies that the average velocity of the blood through the aorta is 26.5 cm/sec
(see Exercise 8-6). However, the blood in the aorta does not flow continuously.
It moves in spurts. During the period of flow, the velocity of the blood is about
three times as high as the overall average value calculated in Exercise 8-6.
Therefore, the kinetic energy per cubic centimeter of flowing blood is
KE
1
2
ρv2 1
2
(1.05) � (79.5)2
3330 erg/cm3
We mentioned earlier that energy density (energy per unit volume) and
pressure are measured by the same unit (i.e., 1 erg/cm3 1 dyn/cm2); therefore,
they can be compared to each other. The kinetic energy of 3330 erg/cm3
is equivalent to 2.50 torr pressure; this is small compared to the blood pressure
in the aorta (which is on the average 100 torr). The kinetic energy in the
smaller arteries is even less because, as the arteries branch, the overall area
increases and, therefore, the flow velocity decreases. For example, when the
total flow rate is 5 liter/min, the blood velocity in the capillaries is only about
0.33 mm/sec.
The kinetic energy of the blood becomes more significant as the rate of
blood flow increases. For example, if during physical activity the flow rate
increases to 25 liter/min, the kinetic energy of the blood is 83,300 erg/cm3,
which is equivalent to a pressure of 62.5 torr. This energy is no longer negligible
compared to the blood pressure measured at rest. In healthy arteries,
the increased velocity of blood flow during physical activity does not present
a problem. During intense activity, the blood pressure rises to compensate for
the pressure drop.
8.8 Turbulence in the Blood
Equation 8.8 shows that if the velocity of a fluid exceeds a specific critical
value, the flow becomes turbulent. Through most of the circulatory system the
blood flow is laminar. Only in the aorta does the flow occasionally become
turbulent. Assuming a Reynold’s number of 2000, the critical velocity for the
onset of turbulence in the 2-cm-diameter aorta is, from Eq. 8.8,
Vc
η
ρD
2000 � 0.04
1.05 � 2
38 cm/sec
Section 8.9 Arteriosclerosis and Blood Flow 111
For the body at rest, the flow velocity in the aorta is below this value. But as
the level of physical activity increases, the flow in the aorta may exceed the
critical rate and become turbulent. In the other parts of the body, however, the
flow remains laminar unless the passages are abnormally constricted.
Laminar flow is quiet, but turbulent flow produces noises due to vibrations
of the various surrounding tissues, which indicate abnormalities in the circulatory
system. These noises, called bruit, can be detected by a stethoscope and
can help in the diagnosis of circulatory disorders.
8.9 Arteriosclerosis and Blood Flow
Arteriosclerosis is the most common of cardiovascular diseases. In the United
States, an estimated 200,000 people die annually as a consequence of this
disease. In arteriosclerosis, the arterial wall becomes thickened, and the artery
is narrowed by deposits called plaque. This condition may seriously impair
the functioning of the circulatory system. A 50% narrowing (stenosis) of the
arterial area is considered moderate. Sixty to seventy percent is considered
severe, and a narrowing above 80% is deemed critical. One problem caused
by stenosis is made clear by Bernoulli’s equation. The blood flow through the
region of constriction is speeded up. If, for example, the radius of the artery
is narrowed by a factor of 3, the cross-sectional area decreases by a factor
of 9, which results in a nine-fold increase in velocity. In the constriction, the
kinetic energy increases by 92, or 81. The increased kinetic energy is at the
expense of the blood pressure; that is, in order to maintain the flow rate at
the higher velocity, the potential energy due to pressure is converted to kinetic
energy. As a result, the blood pressure in the constricted region drops. For
example, if in the unobstructed artery the flow velocity is 50 cm/sec, then in
the constricted region, where the area is reduced by a factor of 9, the velocity
is 450 cm/sec. Correspondingly, the pressure is decreased by about 80 torr
(see Exercise 8-8). Because of the low pressure inside the artery, the external
pressure may actually close off the artery and block the flow of blood. When
such a blockage occurs in the coronary artery, which supplies blood to the
heart muscle, the heart stops functioning.
Stenosis above 80% is considered critical because at this point the blood
flow usually becomes turbulent with inherently larger energy dissipation than
is associated with laminar flow. As a result, the pressure drop in the situation
presented earlier is even larger than calculated using Bernoulli’s equation.
Further, turbulent flow can damage the circulatory system because parts of the
flow are directed toward the artery wall rather than parallel to it, as in laminar
112 Chapter 8 The Motion of Fluids
flow. The blood impinging on the arterial wall may dislodge some of the
plaque deposit which downstream may clog a narrower part of the artery. If
such clogging occurs in a cervical artery, blood flow to some part of the brain
is interrupted causing an ischemic stroke.
There is another problem associated with arterial plaque deposit. The
artery has a specific elasticity; therefore, it exhibits certain springlike properties.
Specifically, in analogy with a spring, the artery has a natural frequency
at which it can be readily set into vibrational motion. (See Chapter 5,
Eq. 5.6.) The natural frequency of a healthy artery is in the range 1 to 2 kilohertz.
Deposits of plaque cause an increase in the mass of the arterial wall and
a decrease in its elasticity. As a result, the natural frequency of the artery is
significantly decreased, often down to a few hundred hertz. Pulsating blood
flow contains frequency components in the range of 450 hertz. The plaquecoated
artery with its lowered natural frequency may now be set into resonant
vibrational motion, which may dislodge plaque deposits or cause further damage
to the arterial wall.
8.10 Power Produced by the Heart
The energy in the flowing blood is provided by the pumping action of the
heart. We will now compute the power generated by the heart to keep the
blood flowing in the circulatory system.
The power PH produced by the heart is the product of the flow rate Q and
the energy E per unit volume of the blood; that is,
PH Q
cm3
sec
� E
erg
cm2
Q � E erg/sec (8.9)
At rest, when the blood flow rate is 5 liter/min, or 83.4 cm3/sec, the kinetic
energy of the blood flowing through the aorta is 3.33 � 103 erg/cm3. (See previous
section.) The energy corresponding to the systolic pressure of 120 torr
is 160 � 103 erg/cm3. The total energy is 1.63 � 105 erg/cm3—the sum of the
kinetic energy and the energy due to the fluid pressure. Therefore, the power
P produced by the left ventricle of the heart is
P 83.4 � 1.63 � 105
1.35 � 107
erg/sec 1.35 W
Exercise 8-9 shows that during intense physical activity when the flow rate
increases to 25 liters/min, the peak power output of the left ventricle increases
to 10.1 W.
Section 8.11 Measurement of Blood Pressure 113
The flow rate through the right ventricle, which pumps the blood through
the lungs, is the same as the flow through the left ventricle. Here, however, the
blood pressure is only one sixth the pressure in the aorta. Therefore, as shown
in Exercise 8-10, the power output of the right ventricle is 0.25 W at rest
and 4.5 W during intense physical activity. Thus, the total peak power output
of the heart is between 1.9 and 14.6 W, depending on the intensity of the
physical activity. While in fact the systolic blood pressure rises with increased
blood flow, in these calculations we have assumed that it remains
at 120 torr.
8.11 Measurement of Blood Pressure
The arterial blood pressure is an important indicator of the health of an individual.
Both abnormally high and abnormally low blood pressures indicate
some disorders in the body that require medical attention. High blood pressure,
which may be caused by constrictions in the circulatory system, certainly
implies that the heart is working harder than usual and that it may be endangered
by the excess load. Blood pressure can be measured most directly by
inserting a vertical glass tube into an artery and observing the height to which
the blood rises (see Fig. 8.5). This was, in fact, the way blood pressure was
first measured in 1733 by Reverend Stephen Hales, who connected a long vertical
glass tube to an artery of a horse. Although sophisticated modifications
of this technique are still used in special cases, this method is obviously not
satisfactory for routine clinical examinations. Routine measurements of blood
pressure are now most commonly performed by the cut-off method. Although
this method is not as accurate as direct measurements, it is simple and in most
cases adequate. In this technique, a cuff containing an inflatable balloon is
placed tightly around the upper arm. The balloon is inflated with a bulb, and
the pressure in the balloon is monitored by a pressure gauge. The initial pressure
in the balloon is greater than the systolic pressure, and the flow of blood
through the artery is therefore cut off. The observer then allows the pressure in
the balloon to fall slowly by releasing some of the air. As the pressure drops,
she listens with a stethoscope placed over the artery downstream from the cuff.
No sound is heard until the pressure in the balloon decreases to the systolic
pressure. Just below this point the blood begins to flow through the artery;
however, since the artery is still partially constricted, the flow is turbulent and
is accompanied by a characteristic sound. The pressure recorded at the onset
of sound is the systolic blood pressure. As the pressure in the balloon drops
further, the artery expands to its normal size, the flow becomes laminar, and
the noise disappears. The pressure at which the sound begins to fade is taken
as the diastolic pressure.
114 Chapter 8 The Motion of Fluids
In clinical measurements, the variation of the blood pressure along the
body must be considered. The cut-off blood pressure measurement is taken
with the cuff placed on the arm approximately at heart level.
EXERCISES
8-1. Calculate the pressure drop per centimeter length of the aorta when the
blood flow rate is 25 liter/min. The radius of the aorta is about 1 cm,
and the coefficient of viscosity of blood is 4 � 10−2 poise.
8-2. Compute the drop in blood pressure along a 30-cm length of artery
of radius 0.5 cm. Assume that the artery carries blood at a rate of
8 liter/min.
8-3. How high a column of blood can an arterial pressure of 100 torr
support? (The density of blood is 1.05 g/cm3.)
8-4. (a) Calculate the arterial blood pressure in the head of an erect person.
Assume that the head is 50 cm above the heart. (The density of blood
is 1.05 g/cm3.) (b) Compute the average arterial pressure in the legs of
an erect person, 130 cm below the heart.
8-5. (a) Show that if the pressure drop remains constant, reduction of the
radius of the arteriole from 0.1 to 0.08 mm decreases the blood flow
by more than a factor of 2. (b) Calculate the decrease in the radius
required to reduce the blood flow by 90%.
8-6. Compute the average velocity of the blood in the aorta of radius 1 cm
if the flow rate is 5 liter/min.
8-7. When the rate of blood flow in the aorta is 5 liter/min, the velocity
of the blood in the capillaries is about 0.33 mm/sec. If the average
diameter of a capillary is 0.008 mm, calculate the number of capillaries
in the circulatory system.
8-8. Compute the decrease in the blood pressure of the blood flowing
through an artery the radius of which is constricted by a factor of 3.
Assume that the average flow velocity in the unconstricted region is
50 cm/sec.
8-9. Using information provided in the text, calculate the power generated
by the left ventricle during intense physical activity when the flow rate
is 25 liter/min.
8-10. Using information provided in the text, calculate the power generated
by the right ventricle during (a) restful state; blood flow 5 liter/min,
and (b) intense activity; blood flow 25 liter/min.
Chapter 8 Exercises 115
8-11. During each heartbeat, the blood from the heart is ejected into the aorta
and the pulmonary artery. Since the blood is accelerated during this
part of the heartbeat, a force in the opposite direction is exerted on
the rest of the body. If a person is placed on a sensitive scale (or
other force-measuring device), this reaction force can be measured.
An instrument based on this principle is called the ballistocardiograph.
Discuss the type of information that might be obtained from measurements
with a ballistocardiograph, and estimate the magnitude of the
forces measured by this instrument.
Chapter 9
Heat and Kinetic Theory
9.1 Heat and Hotness
The sensation of hotness is certainly familiar to all of us. We know from
experience that when two bodies, one hot and the other cold, are placed in
an enclosure, the hotter body will cool and the colder body will heat until the
degree of hotness of the two bodies is the same. Clearly something has been
transferred from one body to the other to equalize their hotness. That which
has been transferred from the hot body to the cold body is called heat. Heat
may be transformed into work, and therefore it is a form of energy. Heated
water, for example, can turn into steam, which can push a piston. In fact, heat
can be defined as energy being transferred from a hotter body to a colder body.
In this chapter, we will discuss various properties associated with heat.
We will describe the motion of atoms and molecules due to thermal energy
and then discuss diffusion in connection with the functioning of cells and the
respiratory system.
9.2 Kinetic Theory of Matter
To understand the present-day concept of heat, we must briefly explain the
structure of matter. Matter is made of atoms and molecules, which are in
continuous chaotic motion. In a gas, the atoms (or molecules) are not bound
together. They move in random directions and collide frequently with one
another and with the walls of the container. In addition to moving linearly, gas
molecules vibrate and rotate, again in random directions. In a solid, where the
116
Section 9.2 Kinetic Theory of Matter 117
atoms are bound together, the random motion is more restricted. The atoms
are free only to vibrate and do so, again randomly, about some average position
to which they are locked. The situation with regard to liquids is between
these two extremes. Here the molecules can vibrate, but they also have some
freedom to move and to rotate.
Because of their motion, the moving particles in a material possess kinetic
energy. This energy of motion inside materials is called internal energy, and
the motion itself is called thermal motion. What we have so far qualitatively
called the hotness of a body is a measure of the internal energy; that is, in hotter
bodies, the random motion of atoms and molecules is faster than in colder
bodies. Therefore, the hotter an object, the greater is its internal energy. The
physical sensation of hotness is the effect of this random atomic and molecular
motion on the sensory mechanism. Temperature is a quantitative measure of
hotness. The internal energy of matter is proportional to its temperature.
Using these concepts, it is possible to derive the equations that describe
the behavior of matter as a function of temperature. Gases are the simplest
to analyze. The theory considers a gas made of small particles (atoms or
molecules) which are in continuous random motion. Each particle travels in
a straight line until it collides with another particle or with the walls of the
container. After a collision, the direction and speed of the particle is changed
randomly. In this way kinetic energy is exchanged among the particles.
The colliding particles exchange energy not only among themselves but
also with the wall of the container (Fig. 9.1). For example, if initially the
walls of the container are hotter than the gas, the particles colliding with the
wall on the average pick up energy from the vibrating molecules in the wall.
As a result of the wall collisions, the gas is heated until it is as hot as the walls.
After that, there is no net exchange of energy between the walls and the gas.
This is an equilibrium situation in which, on the average, as much energy is
delivered to the wall by the gas particles as is picked up from it.
The speed and corresponding kinetic energy of the individual particles in
a gas vary over a wide range. Still it is possible to compute an average kinetic
energy for the particles by adding the kinetic energy of all the individual particles
in the container and dividing by the total number of particles (for details,
see [11-7]). Many of the properties of a gas can be simply derived by assuming
that each particle has this same average energy.
The internal energy in an ideal gas is in the form of kinetic energy,1 and
therefore the average kinetic energy 1
2mv2
av
is proportional to the temperature.
The proportionality can be changed to an equality by multiplying
the temperature T by a suitable constant which relates the temperature to the
internal energy. The constant is designated by the symbol k, which is called
1
The simple theory neglects the vibrational and rotational energy of the molecules.
118 Chapter 9 Heat and Kinetic Theory
FIGURE 9.1 Collisions in a gas.
Boltzmann constant. For historical reasons, Boltzmann constant has been so
defined that it has to be multiplied by a factor of 3
2 to relate temperature to the
average kinetic energy of a molecule; thus,
1
2
mv2
av
3
2
kT (9.1)
The temperature in this equation is measured on the absolute temperature scale
in degrees Kelvin. The size of the degree division on the absolute scale is equal
to the Celsius, or centigrade, degree, but the absolute scale is transposed so that
0◦C 273.15 K. Since our calculations are carried only to three significant figures,
we will use simply 0◦C 273◦K. The value of Boltzmann constant is
k 1.38 � 10−23
J/molecule K
The velocity defined by Eq. 9.1 is called thermal velocity.
Each time a molecule collides with the wall, momentum is transferred to
the wall. The change in momentum per unit time is a force. The pressure
exerted by a gas on the walls of its container is due to the numerous collisions
of the gas molecules with the container. The following relationship between
pressure P, volume V, and temperature is derived in most basic physics texts:
PV NkT (9.2)
Here N is the total number of gas molecules in the container of volume V, and
the temperature is again measured on the absolute scale.
Section 9.3 Definitions 119
TABLE 9.1 Specific Heat for Some Substances
Substance Specific heat(cal/g◦C)
Water 1
Ice 0.480
Average for human body 0.83
Soil 0.2 to 0.8, depending on water
content
Aluminum 0.214
Protein 0.4
In a closed container, the total number of particles N is fixed; therefore,
if the temperature is kept unchanged, the product of pressure and volume is a
constant. This is known as Boyle’s law. (See Exercises 9-1 and 9-2.)
9.3 Definitions
9.3.1 Unit of Heat
As discussed in Appendix A, heat is measured in calories. One calorie (cal) is
the amount of heat required to raise the temperature of 1 g of water by 1 C◦.2
Actually, because this value depends somewhat on the initial temperature of
the water, the calorie is defined as the heat required to raise the temperature
of 1 g of water from 14.5◦C to 15.5◦C. One calorie is equal to 4.184 J. In
the life sciences, heat is commonly measured in kilocalorie units, abbreviated
Cal; 1 Cal is equal to 1000 cal.
9.3.2 Specific Heat
Specific heat is the quantity of heat required to raise the temperature of 1 g of
a substance by 1 degree. The specific heats of some substances are shown in
Table 9.1.
The human body is composed of water, proteins, fat, and minerals. Its
specific heat reflects this composition. With 75% water and 25% protein, the
specific heat of the body would be
Specific heat 0.75 � 1 + 0.25 � 0.4 0.85
The specific heat of the average human body is closer to 0.83 due to its fat and
mineral content, which we have not included in the calculation.
2
For the symbol ◦
C read degree Celsius. For the symbol C◦
, read Celsius degree.
120 Chapter 9 Heat and Kinetic Theory
FIGURE 9.2 Heat is transferred from one region to another by (a) conduction, by
(b) convection, and by (c) radiation.
9.3.3 Latent Heats
In order to convert a solid to a liquid at the same temperature or to convert a
liquid to a gas, heat energy must be added to the substance. This energy is
called latent heat. The latent heat of fusion is the amount of energy required
to change 1 g of solid matter to liquid. The latent heat of vaporization is the
amount of heat required to change 1 g of liquid to gas.
9.4 Transfer of Heat
Heat is transferred from one region to another in three ways: by conduction,
convection, and radiation (Fig. 9.2).
9.4.1 Conduction
If one end of a solid rod is placed in the proximity of a heat source such as a
fire, after some time the other end of the rod will become hot. In this case, heat
has been transferred from the fire through the rod by conduction. The process
of heat conduction involves the increase of internal energy in the material. The
heat enters one end of the rod and increases the internal energy of the atoms
near the heat source. In a solid material, the internal energy is in the vibration
of the bound atoms and in the random motion of free electrons, which exist
in some materials. The addition of heat increases both the random atomic
vibrations and the speed of the electrons. The increased vibrational motion is
transferred down the rod through collisions with neighboring atoms. However,
Section 9.4 Transfer of Heat 121
FIGURE 9.3 Heat flow through a block of material.
because the atoms in the solid are tightly bound, their motion is restricted.
Therefore, heat transfer via atomic vibrations is slow.
In some materials, the electrons in the atoms have enough energy to break
loose from a specific nucleus and move freely through the material. The electrons
move rapidly through the material so that, when they gain energy, they
transfer it quickly to adjacent electrons and atoms. In this way, free electrons
transfer the increase in the internal energy down the rod. Materials such as
metals, which contain free electrons, are good conductors of heat; materials
such as wood, which do not have free electrons, are insulators.
The amount of heat Hc conducted per second through a block of material
(see Fig. 9.3) is given by
Hc
KcA
L
(T1 − T2) (9.3)
Here A is the area of the block, L is its length, and T1 − T2 is the temperature
difference between the two ends. The constant Kc is the coefficient of thermal
conductivity. In physics texts, Kc is usually given in units of cal cm/seccm2-C◦.
However, for problems involving living systems, it is often more
convenient to express Kc in units of Cal cm/m2-hr-C◦. This is the amount of
heat (in Cal units) per hour which flows through a slab of material 1 cm thick
and 1 m square per C◦ temperature difference between the faces of the slab.
The thermal conductivity of a few materials is given in Table 9.2.
9.4.2 Convection
In solids, heat transfer occurs by conduction; in fluids (gases and liquids), heat
transfer proceeds primarily by convection. When a liquid or a gas is heated,
122 Chapter 9 Heat and Kinetic Theory
TABLE 9.2 Thermal Conductivity of
Some Materials
Thermal conductivity, Kc
Material (Cal cm/m2
-hr-C◦)
Silver 3.6 � 104
Cork 3.6
Tissue (unperfused) 18
Felt and down 0.36
Aluminum 1.76 � 104
the molecules near the heat source gain energy and tend to move away from the
heat source. Therefore, the fluid near the heat source becomes less dense. Fluid
from the denser region flows into the rarefied region, causing convection currents.
These currents carry energy away from the heat source. When the energetic
molecules in the heated convection current come in contact with a solid
material, they transfer some of their energy to the atoms of the solid, increasing
the internal energy of the solid. In this way, heat is coupled into a solid. The
amount of heat transferred by convection per unit time Hc is given by
Hc KcA(T1 − T2) (9.4)
Here A is the area exposed to convective currents, T1 − T2 is the temperature
difference between the surface and the convective fluid, and Kc is the
coefficient of convection, which is usually a function of the velocity of the
convective fluid.
9.4.3 Radiation
Vibrating electrically charged particles emit electromagnetic radiation, which
propagates away from the source at the speed of light. Electromagnetic radiation
is itself energy (called electromagnetic energy), which in the case of a
moving charge is obtained from the kinetic energy of the charged particle.
Because of internal energy, particles in a material are in constant random
motion. Both the positively charged nuclei and the negatively charged electrons
vibrate and, therefore, emit electromagnetic radiation. In this way, internal
energy is converted into radiation, called thermal radiation. Due to the
loss of internal energy, the material cools. The amount of radiation emitted
by vibrating charged particles is proportional to the speed of vibration.
Hot objects, therefore, emit more radiation than cold ones. Because the electrons
are much lighter than the nuclei, they move faster and emit more radiant
energy than the nuclei.
Section 9.4 Transfer of Heat 123
When a body is relatively cool, the radiation from it is in the long-wavelength
region to which the eye does not respond. As the temperature (i.e.,
the internal energy) of the body increases, the wavelength of the radiation
decreases. At high temperatures, some of the electromagnetic radiation is in
the visible region, and the body is observed to glow.
When electromagnetic radiation impinges on an object, the charged particles
(electrons) in the object are set into motion and gain kinetic energy.
Electromagnetic radiation is, therefore, transformed into internal energy. The
amount of radiation absorbed by a material depends on its composition. Some
materials, such as carbon black, absorb most of the incident radiation. These
materials are easily heated by radiation. Other materials, such as quartz and
certain glasses, transmit the radiation without absorbing much of it. Metallic
surfaces also reflect radiation without much absorption. Such reflecting and
transmitting materials cannot be heated efficiently by radiation. The rate of
emission of radiant energy Hr by a unit area of a body at temperature T is
Hr eσT 4
(9.5)
Here σ is the Stefan-Boltzmann constant, which is 5.67 � 10−8 W/m2-K4 or
5.67 � 10−5 erg/cm2-◦K4-sec. The temperature is measured on the absolute
scale, and e is the emissivity of the surface, which depends on the temperature
and nature of the surface. The value of the emissivity varies from 0 to 1.
Emission and absorption of radiation are related phenomena; surfaces that are
highly absorptive are also efficient emitters of radiation and have an emissivity
close to 1. Conversely, surfaces that do not absorb radiation are poor emitters
with a low value of emissivity.
A body at temperature T1 in an environment at temperature T2 will both
emit and absorb radiation. The rate of energy emitted per unit area is eσT 4
1 ,
and the rate of energy absorbed per unit is eσT 4
2 . The values for e and σ are
the same for both emission and absorption.
If a body at a temperature T1 is placed in an environment at a lower temperature
T2, the net loss of energy from the body is
Hr eσ T 4
1 − T 4
2 (9.6)
If the temperature of the body is lower than the temperature of the environment,
the body gains energy at the same rate.
9.4.4 Diffusion
If a drop of colored solution is introduced into a still liquid, we observe that the
color spreads gradually throughout the volume of the liquid. The molecules of
124 Chapter 9 Heat and Kinetic Theory
FIGURE 9.4 Random walk.
color spread from the region of high concentration (of the initially introduced
drop) to regions of lower concentration. This process is called diffusion.
Diffusion is the main mechanism for the delivery of oxygen and nutrients
into cells and for the elimination of waste products from cells. On a large
scale, diffusive motion is relatively slow (it may take hours for the colored
solution in our example to diffuse over a distance of a few centimeters), but
on the small scale of tissue cells, diffusive motion is fast enough to provide
for the life function of cells.
Diffusion is the direct consequence of the random thermal motion of molecules.
Although a detailed treatment of diffusion is beyond our scope, some
of the features of diffusive motion can be deduced from simple kinetic theory.
Consider a molecule in a liquid or a gas which is moving away from the
starting point 0. The molecule has a thermal velocity v and travels on the
average a distance L before colliding with another molecule (see Fig. 9.4).
As a result of the collision, the direction of motion of the molecule is changed
randomly. The path may be only slightly deflected, or it may be altered substantially.
On the average, however, after a certain number of collisions the
molecule will be found a distance S from the starting point. A statistical analysis
of this type of motion shows that after N collisions the distance of the
molecule from the starting point is, on the average,
S L N (9.7)
The average distance (L) traveled between collisions is called the mean free
path. This type of diffusive motion is called a random walk.
A frequently used illustration of the random walk examines the position
of a drunkard walking away from a lamppost. He starts off in a particular
Section 9.4 Transfer of Heat 125
direction, but with each step he changes his direction of motion randomly.
If the length of each step is 1 m, after taking 100 steps he will be only 10 m
away from the lamppost although he has walked a total of 100 m. After 10,000
steps, having walked 10 km, he will be still only 100 m (on the average) from
his starting point.
Let us now calculate the length of time required for a molecule to diffuse
a distance S from the starting point. From Eq. 9.7 the number of steps or
collisions that take place while diffusing through a distance S is
N
S2
L2
(9.8)
The total distance traveled is the product of the number of steps and the length
of each step; that is,
Total distance NL
S2
L
(9.9)
If the average velocity of the particle is v, the time t required to diffuse a
distance S is
t
Total distance
v
S2
Lv
(9.10)
Although our treatment of diffusion has been simplified, Eq. 9.10 does
lead to reasonable estimates of diffusion times. In a liquid such as water,
molecules are close together. Therefore, the mean free path of a diffusing
molecule is short, about 10−8 cm (this is approximately the distance between
atoms in a liquid). The velocity of the molecule depends on the temperature
and on its mass. At room temperature, the velocity of a light molecule may be
about 104 cm/sec. From Eq. 9.10, the time required for molecules to diffuse a
distance of 1 cm is
t
S2
Lv
(1)2
10−8 � 104
104
sec 2.8 hr
However, the time required to diffuse a distance of 10−3 cm, which is the
typical size of a tissue cell, is only 10−2 sec (see Exercise 9-3a).
Gases are less densely packed than liquids; consequently, in gases
the mean free path is longer and the diffusion time shorter. In a gas at 1 atm
pressure, the mean free path is on the order of 10−5—the exact value depends
on the specific gas. The time required to diffuse a distance of 1 cm is about
10 sec. Diffusion through a distance of 10−3 cm takes only 10−5 sec (see
Exercise 9-3b).
126 Chapter 9 Heat and Kinetic Theory
9.5 Transport of Molecules by Diffusion
We will now calculate the number of molecules transported by diffusion from
one region to another. Consider a cylinder containing a nonuniform distribution
of diffusing molecules or other small particles (see Fig. 9.5). At position
x 0, the density of the diffusing molecules is C1. At a small distance x
away from this point, the concentration is C2. We can define a diffusion velocity
VD as the average speed of diffusion from x 0 to x x. This velocity
is simply the distance x divided by the average time for diffusion t; that is,
VD
x
t
Substituting t ( x)2/Lv from Eq. 9.10, we obtain
VD
x
( x)2/Lv
Lv
x
(9.11)
(Remember that v here is the thermal velocity.) The number of molecules J
arriving per second per unit area, from region 1 where the density is C1 to
region 2 (see Exercise 9-4) is
J1
VDC1
2
(9.12)
The factor of 2 in the denominator accounts for the fact that molecules are
diffusing both toward and away from region 2. The term J is called the flux,
and it is in units (cm−2s−1).
FIGURE 9.5 Diffusion.
Section 9.5 Transport of Molecules by Diffusion 127
At the same time, molecules are also diffusing from region 2 where the
density is C2 toward region 1. This flux J2 is
J2
VDC2
2
The net flux of molecules into region 2 is the difference between the arriving
and the departing flux, which is
J J1 − J2
VD (C1 − C2)
2
Substituting for VD Lv/ x, we obtain
J
Lv (C1 − C2)
2 x
(9.13)
This derivation assumes that the velocities v in the two regions are the same.
Although this solution for the diffusion problem is not exact, it does illustrate
the nature of the diffusion process. (For a more rigorous treatment, see, for
example, [11-7]). The net flux from one region to another depends on the
difference in the density of the diffusing particles in the two regions. The flux
increases with thermal velocity v and decreases with the distance between the
two regions.
Equation 9.13 is usually written as
J
D
x
(C1 − C2) (9.14)
where D is called the diffusion coefficient. In our case, the diffusion coefficient
is simply
D
Lv
2
(9.15)
In general, however, the diffusion coefficient is a more complex function
because the mean free path L depends on the size of the molecule and the
viscosity of the diffusing medium. In our previous illustration of diffusion
through a fluid, where L 10−8 cm and v 104 cm/sec, the diffusion
coefficient calculated from Eq. 9.15 is 5 � 10−5 cm2/sec. By comparison,
the measured diffusion coefficient of salt (NaCl) in water, for example, is
1.09 � 10−5 cm2/sec. Thus, our simple calculation gives a reasonable estimate
for the diffusion coefficient. Larger molecules, of course, have a smaller
diffusion coefficient. The diffusion coefficients for biologically important
molecules are in the range from 10−7 to 10−6 cm2/sec.
128 Chapter 9 Heat and Kinetic Theory
9.6 Diffusion through Membranes
So far we have discussed only free diffusion through a fluid, but the cells
constituting living systems are surrounded by membranes which impede free
diffusion. Oxygen, nutrients, and waste products must pass through these
membranes to maintain the life functions. In the simplest model, the biological
membrane can be regarded as porous, with the size and the density of the
pores governing the diffusion through the membrane. If the diffusing molecule
is smaller than the size of the pores, the only effect of the membrane is to reduce
the effective diffusion area and thus decrease the diffusion rate. If the diffusing
molecule is larger than the size of the pores, the flow of molecules through the
membranemaybebarred. (SeeFig.9.6.) (Somemoleculesmaystillgetthrough
the membrane, however, by dissolving into the membrane material.)
The net flux of molecules J flowing through a membrane is given in terms
of the permeability of the membrane P
J P (C1 − C2) (9.16)
This equation is similar to Eq. 9.14 except that the term D is replaced by the
permeability P, which includes the diffusion coefficient as well as the effective
thickness x of the membrane. The permeability depends, of course, on the
type of membrane as well as on the diffusing molecule. Permeability may be
nearly zero (if the molecules cannot pass through the membrane) or as high
as 10−4 cm/sec.
FIGURE 9.6 Diffusion through a membrane.
Section 9.7 The Respiratory System 129
The dependence of permeability on the diffusing species allows the cell
to maintain a composition different from that of the surrounding environment.
Many membranes, for example, are permeable to water but do not
pass molecules dissolved in water. As a result water can enter the cell, but the
components of the cell cannot pass out of the cell. Such a one-way passage of
water is called osmosis.
In the type of diffusive motion we have discussed so far, the movement of
the molecules is due to their thermal kinetic energy. Some materials, however,
are transported through membranes with the aid of electric fields that are generated
by charge differences across the membrane. This type of a transport
will be discussed in Chapter 13.
We have shown that over distances larger than a few millimeters diffusion
is a slow process. Therefore, large living organisms must use circulating systems
to transport oxygen nutrients and waste products to and from the cells.
The evolution of the respiratory system in animals is a direct consequence of
the inadequacy of diffusive transportation over long distances.
9.7 The Respiratory System
As will be shown in the following two chapters, animals require energy to
function. This energy is provided by food, which is oxidized by the body.
On the average, 0.207 liter of oxygen at 760 torr are required for every Cal of
energy released by the oxidation of food in the body. At rest, an average 70-kg
adult requires about 70 Cal of energy per hour, which implies a consumption
of 14.5 liter of O2 per hour, which is about 1020 oxygen molecules per second
(see Exercise 9-5).
The simplest way to obtain the required oxygen is by diffusion through the
skin. This method, however, cannot supply the needs of large animals. It has
been determined that in a person only about 2% of oxygen consumed at rest
is obtained by diffusion through the skin. The rest of the oxygen is obtained
through the lungs.
The lungs can be thought of as an elastic bag suspended in the chest cavity
(see Fig. 9.7). When the diaphragm descends, the volume of the lungs
increases, causing a reduction in gas pressure inside the lungs. As a result,
air enters the lungs through the trachea. The trachea branches into smaller
and smaller tubes, which finally terminate at tiny cavities called alveoli. It is
here that gas is exchanged by diffusion between the blood and the air in the
lungs. The lungs of an adult contain about 300 million alveoli with diameters
ranging between 0.1 and 0.3 mm. The total alveolar area of the lungs is about
100 m2, which is about 50 times larger than the total surface area of the skin.
The barrier between the alveolar air and the blood in the capillaries is very
130 Chapter 9 Heat and Kinetic Theory
FIGURE 9.7 Lungs.
thin, only about 4 � 10−5 cm. Therefore, the gas exchange of oxygen into the
blood and CO2 out of the blood is very fast.
The lungs are not fully emptied and filled with each breath. In fact, the full
volume of the lungs is about 6 liter, and at rest only about 1
2 liter is exchanged
during each breath. The composition of inspired and expired air is shown in
Table 9.3.
Using the experimental data in Table 9.3, we can easily show that about
10.5 breaths per minute satisfy the oxygen requirements for a resting person
(see Exercise 9-6). The oxygen requirement, of course, rises with increased
physical activity, which results in both faster and deeper breathing. During
Section 9.7 The Respiratory System 131
TABLE 9.3 The Percentage of N2, O2,
and CO2 in Inspired and Expired Air for
a Resting Person
N2 O2 CO2
Inspired air 79.02 20.94 0.04
Expired air 79.2 16.3 4.5
deep breathing, as much as 70% of the air in the lungs is exchanged in each
breath.
While diffusion through the skin can supply only a small fraction of the
oxygen required by large animals, the oxygen needs of small animals may
be completely satisfied through this channel. This can be deduced from the
following considerations. The energy consumption and, hence, the oxygen
requirement of an animal is approximately proportional to its mass.3 The
mass in turn is proportional to the volume of the animal. The amount of
oxygen diffusing through the skin is proportional to the surface area of the
skin. Now, if R is a characteristic linear dimension of the animal, the volume
is proportional to R3, and the skin surface area is proportional to R2. The
surface to volume ratio is given by
Surface area
Volume
R2
R3
1
R
(9.17)
Therefore, as the size of the animal R decreases, its surface-to-volume
ratio increases; that is, for a unit volume, a small animal has a greater surface
area than a large animal.
It is possible to obtain an estimate for the maximum size of the animal
that can get its oxygen entirely by skin diffusion. A highly simplified calculation
outlined in Exercise 9-7 shows that the maximum linear size of such an
animal is about 0.5 cm. Therefore, only small animals, such as insects, can
rely entirely on the diffusion transfer to provide them with oxygen. However,
during hibernation when the oxygen requirements of the animal are reduced
to a very low value, larger animals such as frogs can obtain all the necessary
oxygen through their skin. In fact some species of frog hibernate through
the winter at the bottom of lakes where the temperature is constant at 4◦C.
3
This is an approximation. A more detailed discussion is found in [11-10].
132 Chapter 9 Heat and Kinetic Theory
The required oxygen enters the frog’s body by diffusion from the surrounding
water, which contains dissolved oxygen.
9.8 Surfactants and Breathing
The discussion in the previous section neglected an important aspect of
breathing, the size of the alveoli. As was stated, the diameters of the alveoli
range from about 0.1 to 0.3 mm (radius R from 0.05 to 0.15 mm). The inner
wall of the alveoli is coated with a thin layer of water that protects the tissue.
The surface tension of this water layer tends to minimize the surface thereby
shrinking the alveolar cavity. When the diaphragm descends, the incoming air
has to enter the alveoli and expand them to their full size. Because the alveoli
are embedded in a moist medium, expanding the alveoli is analogous to creating
a bubble inside a liquid. As was discussed in Section 7.7, to create gas
bubble of radius R in a liquid with surface tension T, the pressure of the gas
injected into the liquid must be greater than the pressure of the surrounding
liquid by P as given in Eq. 7.21. As is shown in Exercise 9-8, P required
to expand a 0.05 diameter alveolus to its full volume is 2.9 atmospheres. This
is the minimum pressure required to open a 0.05 diameter alveolus that has
its walls coated with plain water. Clearly the incoming air at one atmosphere
cannot open the small alveoli and can barely begin to expand the larger ones.
Breathing is made possible by surfactants that cover the alveolar water
layer and greatly reduce its surface tension. These surfactant molecules are a
complex mixture of lipids and proteins produced by special cells in the alveoli
and they can reduce surface tension by as much as a factor of 70 (to about
1 dyn/cm).
The lungs of premature infants often fail to produce adequate amounts of
surfactants required for breathing. This life threatening condition called Infant
Respiratory Distress Syndrome can now be treated with artificial lung surfactants
developed in the 1980s. When introduced into the lungs of the infant
these surfactants often stabilize breathing till the alveoli begin to produce surfactants
on their own.
Cold-blooded animals such as frogs, snakes and lizards do not need
lung surfactants for breathing. Such animals do not use energy to heat their
bodies. As a result they require about a factor of ten less oxygen than warmblooded
animals of comparable size. Therefore, cold-blooded animals can
function with correspondingly smaller lung surface area. The alveolal radii
of these animals are ten times larger than those of warm-blooded animals
(see Exercise 9-9). An alveolus of larger radius requires correspondingly
lower pressure to overcome surface tension eliminating the need for lung
surfactants.
Chapter 9 Exercises 133
9.9 Diffusion and Contact Lenses
Most parts of the human body receive the required oxygen from the circulating
blood. However, the cornea, which is the transparent surface layer of the eye,
does not contain blood vessels (this allows it to be transparent). The cells in
the cornea receive oxygen by diffusion from the surface layer of tear fluid,
which contains oxygen. This fact allows us to understand why most contact
lenses should not be worn during sleep. The contact lens is fitted so that
blinking rocks the lens slightly. This rocking motion brings fresh oxygen-rich
tear fluid under the lens. Of course, when people sleep they do not blink;
therefore, the corneas under their contact lenses are deprived of oxygen. This
may result in a loss of corneal transparency.
EXERCISES
9-1. Fish using air bladders to control their buoyancy are less stable than
those using porous bones. Explain this phenomenon using the gas
equation (Eq. 9.2). (Hint: What happens to the air bladder as the fish
sinks to a greater depth?)
9-2. A scuba diver breathes air from a tank which has a pressure regulator
that automatically adjusts the pressure of the inhaled air to the ambient
pressure. If a diver 40 m below the surface of a deep lake fills his lungs
to the full capacity of 6 liters and then rises quickly to the surface, to
what volume will his lungs expand? Is such a rapid ascent advisable?
9-3. (a) Calculate the time required for molecules to diffuse in a liquid a
distance of 10−3 cm. Assume that the average velocity of the molecules
is 104 cm/sec and that the mean free path is 10−8 cm. (b) Repeat the
calculation for diffusion in a gas at 1 atm pressure, where the mean free
path is 10−5 cm.
9-4. Consider a beam of particles traveling at a velocity VD. If the area of the
beam is A and the density of particle in the beam is C, show that the number
of particles that pass by a given point each second is VD � C � A.
9-5. A consumption of 14.5 liters of oxygen per hour is equivalent to how
many molecules per second? (The number of molecules per cubic
centimeter at 0 ◦C and 760 torr is 2.69 � 1019.)
9-6. Using the data in the text and in Table 9.3, calculate the number of
breaths per minute required to satisfy the oxygen needs of a resting
person.
134 Chapter 9 Heat and Kinetic Theory
9-7. (a) We stated in the text that the oxygen consumption at rest for a 70-kg
person is 14.5 liter/h and that 2% of this requirement is provided by the
diffusion of oxygen through the skin. Assuming that the skin surface area
of the person is 1.7 m2, calculate the diffusion rate for oxygen through
the skin in liter/h-cm2. (b) What is the maximum linear size of an animal
whose oxygen requirements at rest can be provided by diffusion through
the skin?
Use the following assumptions:
(i) The density of animal tissue is 1 g/cm3.
(ii) Per unit volume, all animals require the same amount of oxygen.
(iii) The animal is spherical in shape.
9-8. Calculate the excess pressure P required to expand a 0.05 diameter
alveolus to its full volume.
9-9. Show that if the oxygen requirement of an animal is reduced by a
factor of 10, then within the same lung volume, alveolar radius can be
increased by a factor of 10.
Chapter 10
Thermodynamics
Thermodynamics is the study of the relationship between heat, work, and
the associated flow of energy. After many decades of experience with heat
phenomena, scientists formulated two fundamental laws as the foundation of
thermodynamics. The First Law of Thermodynamics states that energy, which
includes heat, is conserved; that is, one form of energy can be converted into
another, but energy can neither be created nor destroyed. This implies that the
total amount of energy in the universe is a constant.1 The second law, more
complex than the first, can be stated in a number of ways which, although they
appear different, can be shown to be equivalent. Perhaps the simplest statement
of the Second Law of Thermodynamics is that spontaneous change in
nature occurs from a state of order to a state of disorder.
10.1 First Law of Thermodynamics
One of the first to state the law of energy conservation was the German physician
Robert Mayer (1814–1878). In 1840 Mayer was the physician on the
schooner Java, which sailed for the East Indies. While aboard ship, he was
reading a treatise by the French scientist Laurent Lavoisier in which Lavoisier
suggested that the heat produced by animals is due to the slow combustion of
food in their bodies. Lavoisier further noted that less food is burned by the
body in a hot environment than in a cold one.
1
It has been shown by the theory of relativity that the conservation law must include matter
which is convertible to energy.
135
136 Chapter 10 Thermodynamics
FIGURE 10.1 The energetics of the body.
When the ship reached the tropics, many of its crew became sick with
fever. Applying the usual remedy for fever, Mayer bled his patients. He
noticed that the venous blood, which is normally dark red, was nearly as red
as arterial blood. He considered this a verification of Lavoisier’s suggestion.
Because in the tropics less fuel is burned in the body, the oxygen content of
the venal blood is high, giving it the brighter color. Mayer then went beyond
Lavoisier’s theory and suggested that in the body there is an exact balance of
energy (which he called force). The energy released by the food is balanced
by the lost body heat and the work done by the body. Mayer wrote in an article
published in 1842, “Once in existence, force [energy] cannot be annihilated—
it can only change its form.”
Considerably more evidence had to be presented before conservation
of energy was accepted as a law, but it is interesting that such a fundamental
physical law was first suggested from the observation of human
physiology.
Conservation of energy is implicit in all our calculations of energy balance
in living systems. Consider, for example, the energetics for the functioning
of an animal (see Fig. 10.1). The body of an animal contains internal thermal
energy Et, which is the product of the mass and specific heat, and chemical
energy Ec stored in the tissue of the body. In terms of energy, the activities of an
animal consist of simply eating, working, and rejecting excess heat by means
of various cooling mechanisms (radiation, convection, etc.). Without going
into detailed calculations, the first law allows us to draw some conclusions
about the energetics of the animal. For example, if the internal temperature
and the weight of the animal are to remain constant (i.e., Ec and Et constant),
over a given period of time the energy intake must be exactly equal to the
sum of the work done and the heat lost by the body. An imbalance between
intake and output energy implies a change in the sum Ec + Et. The First Law
of Thermodynamics is implicit in all the numerical calculations presented in
Chapter 11.
Section 10.2 Second Law of Thermodynamics 137
10.2 Second Law of Thermodynamics
There are many imaginable phenomena that are not forbidden by the First Law
of Thermodynamics but still do not occur. For example, when an object falls
from a table to the ground, its potential energy is first converted into kinetic
energy; then, as the object comes to rest on the ground, the kinetic energy is
converted into heat. The First Law of Thermodynamics does not forbid the
reverse process, whereby the heat from the floor would enter the object and be
converted into kinetic energy, causing the object to jump back on the table. Yet
this event does not occur. Experience has shown that certain types of events
are irreversible. Broken objects do not mend by themselves. Spilled water
does not collect itself back into a container. The irreversibility of these types
of events is intimately connected with the probabilistic behavior of systems
comprised of a large ensemble of subunits.
As an example, consider three coins arranged heads up on a tray. We
will consider this an ordered arrangement. Suppose that we now shake the
tray so that each coin has an equal chance of landing on the tray with either
head or tail up. The possible arrangements of coins that we may obtain are
shown in Table 10.1. Note that there are eight possible outcomes of tossing
the three coins. Of these, only one yields the original ordered arrangement
of three heads (H,H,H). Because the probabilities of obtaining any one of the
coin arrangements in Table 10.1 are the same, the probability of obtaining the
three-head arrangement after shaking the tray once is 1/8, or 0.125; that is, on
the average, we must toss the coins eight times before we can expect to see
the three-head arrangement again.
As the number of coins in the experiment is increased, the probability of
returning to the ordered arrangement of all heads decreases. With 10 coins on
the tray, the probability of obtaining all heads after shaking the tray is 0.001.
TABLE 10.1 The Ordering
of Three Coins
Coin 1 Coin 2 Coin 3
H H H
H H T
H T H
T H H
H T T
T H T
T T H
T T T
138 Chapter 10 Thermodynamics
With 1000 coins, the probability of obtaining all heads is so small as to be
negligible. We could shake the tray for many years without seeing the ordered
arrangement again. In summary, the following is to be noted from this illustration:
The number of possible coin arrangements is large, and only one of
them is the ordered arrangement; therefore, although any one of the coin
arrangements—including the ordered one—is equally likely, the probability
of returning to an ordered arrangement is small. As the number of coins in the
ensemble increases, the probability of returning to an ordered arrangement
decreases. In other words, if we disturb an ordered arrangement, it is likely
to become disordered. This type of behavior is characteristic of all events that
involve a collective behavior of many components.
The Second Law of Thermodynamics is a statement about the type of probabilistic
behavior illustrated by our coin experiment. One statement of the
second law is: The direction of spontaneous change in a system is from an
arrangement of lesser probability to an arrangement of greater probability;
that is, from order to disorder. This statement may seem to be so obvious as to
be trivial, but, once the universal applicability of the second law is recognized,
its implications are seen to be enormous. We can deduce from the second law
the limitations on information transmission, the meaning of time sequence,
and even the fate of the universe. These subjects, however, are beyond the
scope of our discussion.
One important implication of the second law is the limitation on the conversion
of heat and internal energy to work. This restriction can be understood
by examining the difference between heat and other forms of energy.
10.3 Difference between Heat and Other Forms of
Energy
We defined heat as energy being transferred from a hotter to a colder body. Yet
when we examined the details of this energy transfer, we saw that it could be
attributed to transfer of a specific type of energy such as kinetic, vibrational,
electromagnetic, or any combination of these (see Chapter 9). For this reason,
it may not seem obvious why the concept of heat is necessary. It is, in
fact, possible to develop a theory of thermodynamics without using the concept
of heat explicitly, but we would then have to deal with each type of energy
transfer separately, and this would be difficult and cumbersome. In many cases,
energy is being transferred to or from a body by different methods, and keeping
track of each of these is often not possible and usually not necessary. No matter
how energy enters the body, its effect is the same. It raises the internal energy
of the body. The concept of heat energy is, therefore, very useful.
The main feature that distinguishes heat from other forms of energy is
the random nature of its manifestations. For example, when heat flows via
Section 10.3 Difference between Heat and Other Forms of Energy 139
FIGURE 10.2 The motion of a piston.
conduction from one part of the material to another part, the flow occurs
through the sequential increase in the internal energy along the material. This
internal energy is in the form of random chaotic motion of atoms. Similarly,
when heat is transferred by radiation, the propagating waves travel in random
directions. The radiation is emitted over a wide wavelength (color) range, and
the phases of the wave along the wave front are random. By comparison, other
forms of energy are more ordered. Chemical energy, for example, is present
by virtue of specific arrangements of atoms in a molecule. Potential energy is
due to the well-defined position, or configuration, of an object.
While one form of energy can be converted to another, heat energy,
because of its random nature, cannot be completely converted to other forms
of energy. We will use the behavior of a gas to illustrate our discussion. First,
let us examine how heat is converted to work in a heat engine (for example,
the steam engine). Consider a gas in a cylinder with a piston (see Fig. 10.2).
Heat flows into the gas; this increases the kinetic energy of the gas molecules
and, therefore, raises the internal energy of the gas. The molecules moving in
the direction of the piston collide with the piston and exert a force on it. Under
the influence of this force, the piston moves. In this way, heat is converted into
work via internal energy.
The heat added to the gas causes the molecules in the cylinder to move
in random directions, but only the molecules that move in the direction of
the piston can exert a force on it. Therefore, the kinetic energy of only the
molecules that move toward the piston can be converted into work. For the
added heat to be completely converted into work, all the gas molecules would
have to move in the direction of the piston motion. In a large ensemble of
molecules, this is very unlikely.
The odds against the complete conversion of 1 cal of heat into work can
be expressed in terms of a group of monkeys who are hitting typewriter keys
at random and who by chance type out the complete works of Shakespeare
without error. The probability that 1 cal of heat would be completely converted
to work is about the same as the probability that the monkeys would type
140 Chapter 10 Thermodynamics
FIGURE 10.3 Conversion of heat to work.
Shakespeare’s works 15 quadrillion times in succession. (This example is
taken from [11-2].)
The distinction between work and heat is this: In work, the energy is in
an ordered motion; in heat, the energy is in random motion. Although some
of the random thermal motion can be ordered again, the ordering of all the
motion is very improbable. Because the probability of completely converting
heat to work is vanishingly small, the Second Law of Thermodynamics states
categorically that it is impossible.
Heat can be partially converted to work as it flows from a region of higher
temperature T1 to a region of lower temperature T2 (see Fig. 10.3). A quantitative
treatment of thermodynamics shows (see, for example, [11-5]) that the
maximum ratio of work to the input heat is
Work
Heat input
1 −
T2
T1
(10.1)
Here the temperature is measured on the absolute scale.
From this equation, it is evident that heat can be completely converted into
work only if the heat is rejected into a reservoir at absolute zero temperature.
Although objects can be cooled to within a very small fraction of absolute
zero, absolute zero cannot be attained. Therefore, heat cannot be completely
converted into work.
10.4 Thermodynamics of Living Systems
It is obvious that animals need food to live, but the reason for this is less
obvious. The idea that animals need energy because they consume energy is,
Section 10.4 Thermodynamics of Living Systems 141
strictly speaking, incorrect. We know from the First Law of Thermodynamics
that energy is conserved. The body does not consume energy, it changes it
from one form to another. In fact, the first law could lead us to the erroneous
conclusion that animals should be able to function without a source of external
energy. The body takes in energy that is in the chemical bonds of the food
molecules and converts it to heat. If the weight and the temperature of the
body remain constant and if the body performs no external work, the energy
input to the body equals exactly the heat energy leaving the body. We may
suppose that if the heat outflow could be stopped—by good insulation, for
example—the body could survive without food. As we know, this supposition
is wrong. The need for energy is made apparent by examining the functioning
of the body in the light of the Second Law of Thermodynamics.
The body is a highly ordered system. A single protein molecule in the
body may consist of a million atoms bound together in an ordered sequence.
Cells are more complex still. Their specialized functions within the body
depend on a specific structure and location. We know from the Second Law
of Thermodynamics that such a highly ordered system, left to itself, tends
to become disordered, and once it is disordered, it ceases to function. Work
must be done on the system continuously to prevent it from falling apart. For
example, the blood circulating in veins and arteries is subject to friction, which
changes kinetic energy to heat and slows the flow of blood. If a force were not
applied to the blood, it would stop flowing in a few seconds. The concentration
of minerals inside a cell differs from that in the surrounding environment.
This represents an ordered arrangement. The natural tendency is toward an
equalization with the environment. Work must be done to prevent the contents
of the cell from leaking out. Finally, cells that die must be replaced, and if the
animal is growing, new tissue must be manufactured. For such replacement
and growth, new proteins and other cell constituents must be put together
from smaller, relatively more random subcomponents. Thus, the process of
life consists of building and maintaining ordered structures. In the face of the
natural tendency toward disorder, this activity requires work. The situation is
somewhat analogous to a pillar made of small, slippery, uneven blocks that
tend to slide out of the structure. The pillar remains standing only if blocks
are continuously pushed back.
The work necessary to maintain the ordered structures in the body is
obtained from the chemical energy in food. Except for the energy utilized
in external work done by the muscles, all the energy provided by food is ultimately
converted into heat by friction and other dissipative processes in the
body. Once the temperature of the body is at the desired level, all the heat
generated by the body must leave through the various cooling mechanisms
of the body (see Chapter 11). The heat must be dissipated because, unlike
heat engines (such as the turbine or the steam engine), the body does not have
the ability to obtain work from heat energy. The body can obtain work only
142 Chapter 10 Thermodynamics
from chemical energy. Even if the body did have mechanisms for using heat
to perform work, the amount of work it could obtain in this way would be
small. Once again, the second law sets the limit. The temperature differences
in the body are small—not more than about 7 C◦ between the interior and the
exterior. With the interior temperature T1 at 310 K (37◦C) and the exterior
temperature T1 at 303 K, the efficiency of heat conversion to work would be
(from Eq. 10.1) at most only about 2%.
Of all the various forms of energy, the body can utilize only the chemical
binding energy of the molecules which constitute food. The body does not
have a mechanism to convert the other forms of energy into work. A person
could bask in the sun indefinitely, receiving large quantities of radiant energy,
and yet die of starvation. Plants, on the other hand, are able to utilize radiant
energy. As animals use chemical energy, so plants utilize solar radiation to
provide the energy for the ordering processes necessary for life.
The organic materials produced in the life cycle of plants provide food
energy for herbivorous animals, which in turn are food for the carnivorous
animals that eat them. The sun is, thus, the ultimate source of energy for life
on Earth.
Since living systems create order out of relative disorder (for example, by
synthesizing large complex molecules out of randomly arranged subunits), it
may appear at first glance that they violate the Second Law of Thermodynamics,
but this is not the case. To ascertain that the second law is valid, we must
examine the whole process of life, which includes not only the living unit but
also the energy that it consumes and the by-products that it rejects. To begin
with, the food that is consumed by an animal contains a considerable degree
of order. The atoms in the food molecules are not randomly arranged but
are ordered in specific patterns. When the chemical energy in the molecular
bindings of the food is released, the ordered structures are broken down. The
eliminated waste products are considerably more disordered than the food
taken in. The ordered chemical energy is converted by the body into disordered
heat energy.
The amount of disorder in a system can be expressed quantitatively by
means of a concept called entropy. Calculations show that, in all cases, the
increase in the entropy (disorder) in the surroundings produced by the living
system is always greater than the decrease in entropy (i.e., ordering) obtained
in the living system itself. The total process of life, therefore, obeys the second
law. Thus, living systems are perturbations in the flow toward disorder. They
keep themselves ordered for a while at the expense of the environment. This
is a difficult task requiring the use of the most complex mechanisms found in
nature. When these mechanisms fail, as they eventually must, the order falls
apart, and the organism dies.
Section 10.5 Information and the Second Law 143
10.5 Information and the Second Law
We have stressed earlier that work must be done to create and maintain the
highly ordered local state of life. We now turn to the question, what else is
needed for such local ordering to occur? Perhaps we can get an insight into
this issue from a simple everyday experience. In the course of time, our apartment
becomes disordered. Books, which had been placed neatly, in alphabetical
order, on a shelf in the living room, are now strewn on the table and some
are even under the bed. Dishes that were clean and neatly stacked in the cupboard,
are now dirty with half-eaten food and are on the living room table. We
decide to clean up, and in 15 minutes or so the apartment is back in order. The
books are neatly shelved, and the dishes are clean and stacked in the kitchen.
The apartment is clean.
Two factors were necessary for this process to occur. First, as was already
stated, energy was required to do the work of gathering and stacking the books
and cleaning and ordering the dishes. Second, and just as important, information
was required to direct the work in the appropriate direction. We had to
know where to place the books and how to clean the dishes and stack them
just so. The concept of information is of central importance here.
In the 1940s, Claude Shannon developed a quantitative formulation for
the amount of information available in a given system. Shannon’s formula for
information content is shown to be equivalent to the formula for entropy—the
measure of disorder—except, with a negative sign. This mathematical insight
formally shows that if energy and information are available, the entropy in
a given locality can be decreased by the amount of information available to
engage in the process of ordering. In other words, as in our example of the
messy living room, order can be created in a disordered system by work that
is directed by appropriate information. The second law, of course, remains
valid: the overall entropy of the universe increases. The work required to
perform the ordering, one way or another, causes a greater disorder in the
surroundings than the order that was created in the system itself. It is the
availability of information and energy that allows living systems to replicate,
grow, and maintain their structures.
The chain of life begins with plants that possess information in their genetic
material on how to utilize the energy from the sun to make highly ordered complex
structures from the simple molecules available to them: principally water,
carbon dioxide, and an assortment of minerals. The process is, in essence,
similar in human beings and other animals. All the information required for
the function of the organism is contained in the intricate structure of DNA. The
human DNA consists of about a billion molecular units in a well-determined
sequence. Utilizing the energy obtained from the food that is consumed by
144 Chapter 10 Thermodynamics
the organism, the information in the DNA guides the assembly of the various
proteins and enzymes required for the functioning of the organism.
EXERCISES
10-1. Explain how the second law of thermodynamics limits conversion of heat
to work.
10-2. From your own experience, give an example of the second law of ther-
modynamics.
10-3. Describe the connections between information, the second law of
thermodynamics, and living systems.
Chapter 11
Heat and Life
The degree of hotness, or temperature, is one of the most important environmental
factors in the functioning of living organisms. The rates of the
metabolic processes necessary for life, such as cell divisions and enzyme reactions,
depend on temperature. Generally the rates increase with temperature.
A 10 degree change in temperature may alter the rate by a factor of 2.
Because liquid water is an essential component of living organisms as we
know them, the metabolic processes function only within a relatively narrow
range of temperatures, from about 2◦C to 120◦C. Only the simplest of living
organisms can function near the extremes of this range.1 Large-scale living
systems are restricted to a much narrower range of temperatures.
The functioning of most living systems, plants and animals, is severely
limited by seasonal variations in temperature. The life processes in reptiles,
for example, slow down in cold weather to a point where they essentially cease
to function. On hot sunny days these animals must find shaded shelter to keep
their body temperatures down.
For a given animal, there is usually an optimum rate for the various metabolic
processes. Warm-blooded animals (mammals and birds) have evolved
methods for maintaining their internal body temperatures at near constant levels.
As a result, warm-blooded animals are able to function at an optimum
level over a wide range of external temperatures. Although this temperature
regulation requires additional expenditures of energy, the adaptability
achieved is well worth this expenditure.
1
In deep oceans, the pressure is high and so is the boiling point of water. Here certain
thermophilic bacteria can survive near thermal vents at significantly higher temperatures.
145
146 Chapter 11 Heat and Life
TABLE 11.1 Metabolic Rates for Selected Activities
Metabolic rate
Activity (Cal/m2
-hr)
Sleeping 35
Lying awake 40
Sitting upright 50
Standing 60
Walking (3 mph) 140
Moderate physical work 150
Bicycling 250
Running 600
Shivering 250
In this chapter, we will examine energy consumption, heat flow, and temperature
control in animals. Although most of our examples will be specific
to people, the principles are generally applicable to all animals.
11.1 Energy Requirements of People
All living systems need energy to function. In animals, this energy is used to
circulate blood, obtain oxygen, repair cells, and so on. As a result, even at
complete rest in a comfortable environment, the body requires energy to sustain
its life functions. For example, a man weighing 70 kg lying quietly awake
consumes about 70 Cal/h (1 cal 4.18 J; 1,000 cal 1 Cal; 1 Cal/h 1.16 W).
Of course, the energy expenditure increases with activity.
The amount of energy consumed by a person depends on the person’s
weight and build. It has been found, however, that the amount of energy
consumed by a person during a given activity divided by the surface area of
the person’s body is approximately the same for most people. Therefore, the
energy consumed for various activities is usually quoted in Cal/m2-hr. This
rate is known as the metabolic rate. The metabolic rates for some human
activities are shown in Table 11.1. To obtain the total energy consumption per
hour, we multiply the metabolic rate by the surface area of the person. The
following empirical formula yields a good estimate for the surface area.
Area (m2
) 0.202 � W0.425
� H0.725
(11.1)
Here W is the weight of the person in kilograms, and H is the height of the
person in meters.
Section 11.2 Energy from Food 147
The surface area of a 70-kg man of height 1.55 m is about 1.70 m2. His
metabolic rate at rest is therefore (40 Cal/m2-hr) � 1.70 m2 68 Cal/hr, or
about 70 Cal/hr as stated in our earlier example. This metabolic rate at rest is
called the basal metabolic rate.
11.2 Energy from Food
The chemical energy used by animals is obtained from the oxidation of food
molecules. The glucose sugar molecule, for example, is oxidized as follows:
C6H12O6 + 6O2 → 6CO2 + 6H2O + energy (11.2)
For every gram of glucose ingested by the body, 3.81 Cal of energy is released
for metabolic use.
The caloric value per unit weight is different for various foods. Measurements
show that, on the average, carbohydrates (sugars and starches) and proteins
provide about 4 Cal/g; lipids (fats) produce 9 Cal/g, and the oxidation of
alcohol produces 7 Cal/g.2
The oxidation of food, which releases energy, does not occur spontaneously
at normal environmental temperatures. For oxidation to proceed at body
temperature, a catalyst must promote the reaction. In living systems, complex
molecules, called enzymes, provide this function.
In the process of obtaining energy from food, oxygen is always consumed.
It has been found that, independent of the type of food being utilized, 4.83 Cal
of energy are produced for every liter of oxygen consumed. Knowing this
relationship, one can measure with relatively simple techniques the metabolic
rate for various activities (see Exercise 11-1).
The daily food requirements of a person depend on his or her activities.
A sample schedule and the associated metabolic energy expenditure per square
meter are shown in Table 11.2. Assuming, as before, that the surface area of
the person whose activities are shown in Table 11.2 is 1.7 m2, his/her total
energy expenditure is 3940 Cal/day. If the person spent half the day sleeping
and half the day resting in bed, the daily energy expenditure would be only
1530 Cal.
For most people the energy expenditure is balanced by the food intake.
For example, the daily energy needs of the person whose activities are shown
2
The high caloric content of alcohol presents a problem for people who drink heavily. The
body utilizes fully the energy released by the oxidation of alcohol. Therefore, people who obtain
a significant fraction of metabolic energy from this source reduce their intake of conventional
foods. Unlike other foods, however, alcohol does not contain vitamins, minerals, and other
substances necessary for proper functioning. As a result, chronic alcoholics often suffer from
diseases brought about by nutritional deficiencies.
148 Chapter 11 Heat and Life
TABLE 11.2 One Day’s Metabolic Energy Expenditure
Energy expenditure
Activity (Cal/m2
)
8 hr sleeping (35 Cal/m2
-hr) 280
8 hr moderate physical labor (150 Cal/m2
-hr) 1200
4 hr reading, writing, TV watching (60 Cal/m2
-hr) 240
1 hr heavy exercise (300 Cal/m2
-hr) 300
3 hr dressing, eating (100 Cal/m2
-hr) 300
Total expenditure 2320
TABLE 11.3 Composition and Energy Content of Some Common
Foods
Total Protein Carbohydrate Fat Total
Food weight (g) weight (g) weight (g) weight (g) energy (Cal)
Whole milk,
1 quart 976 32 48 40 660
Egg, 1 50 6 0 12 75
Hamburger, 1 85 21 0 17 245
Carrots, 1 cup 150 1 10 0 45
Potato (1 med.,
baked) 100 2 22 0 100
Apple 130 0 18 0 70
Bread, rye,
1 slice 23 2 12 0 55
Doughnut 33 2 17 7 135
in Table 11.2 (surface area 1.7 m2) are met by the consumption of 400 g of
carbohydrates, 200 g of protein, and 171 g of fat.
The composition and energy content of some common foods are shown in
Table 11.3. Note that the sum of the weights of the protein, carbohydrates, and
fat is smaller than the total weight of the food. The difference is due mostly
to the water content of the food. The energy values quoted in the table reflect
the fact that the caloric content of different proteins, carbohydrates, and fats
deviate somewhat from the average values stated in the text.
If an excess of certain substances, such as water and salt, is ingested, the
body is able to eliminate it. The body has no mechanism, however, for eliminating
an excess in caloric intake. Over a period of time the excess energy
is used by the body to manufacture additional tissue. If the consumption of
excess food occurs simultaneously with heavy exercise, the energy may be
Section 11.3 Regulation of Body Temperature 149
utilized to increase the weight of the muscles. Most often, however, the excess
energy is stored in fatty tissue that is manufactured by the body. Conversely,
if the energy intake is lower than the demand, the body consumes its own tissue
to make up the deficit. While the supply lasts, the body first utilizes its
stored fat. For every 9 Cal of energy deficit, about 1 g of fat is used. Under
severe starvation, once the fat is used up, the body begins to consume its own
protein. Each gram of consumed protein yields about 4 Cal. Consumption of
body protein results in the deterioration of body functions, of course. A relatively
simple calculation (see Exercise 11-4) shows that an average healthy
person can survive without food but with adequate water up to about 50 days.
Overweight people can do better, of course. The “Guinness Book of World
Records” states that Angus Barbieri of Scotland fasted from June, 1965, to
July, 1966, consuming only tea, coffee, and water. During this period, his
weight declined from 472 lb to 178 lb.
For a woman, the energy requirements increase somewhat during pregnancy
due to the growth and metabolism of the fetus. As the following calculation
indicates, the energy needed for the growth of the fetus is actually rather
small. Let us assume that the weight gain of the fetus during the 270 days of
gestation is uniform.3 If at birth the fetus weighs 3 kg, each day it gains 11 g.
Because 75% of tissue consists of water and inorganic minerals, only 2.75 g of
the daily mass increase is due to organic materials, mainly protein. Therefore,
the extra Calories per day required for the growth of the fetus is
Calories required
2.75 g protein
day
�
4 Cal
g protein
11 Cal/day
To this number, we must add the basal metabolic consumption of the fetus.
At birth, the surface area of the fetus is about 0.13 m2 (from Eq. 11.1); therefore,
at most, the basal metabolic consumption of the fetus per day is about
0.13 � 40 � 24 125 Cal. Thus, the total increase in the energy requirement
of a pregnant woman is only about (125 + 11) Cal/day 136 Cal/day. Actually,
it may not even be necessary for a pregnant woman to increase her food
intake, as the energy requirements of the fetus may be balanced by decreased
physical activity during pregnancy. Various other aspects of metabolic energy
balance are examined in Exercises 11-2 to 11-5.
11.3 Regulation of Body Temperature
People and other warm-blooded animals must maintain their body temperatures
at a nearly constant level. For example, the normal internal body
3
This is a simplification because the weight gain is not uniform. It is greatest toward the end
of gestation.
150 Chapter 11 Heat and Life
temperature of a person is about 37◦C. A deviation of one or two degrees
in either direction may signal some abnormality. If the temperature-regulating
mechanisms fail and the body temperature rises to 44◦ or 45◦C, the protein
structures are irreversibly damaged. A fall in body temperature below about
28◦C results in heart stoppage.
The body temperature is sensed by specialized nerve centers in the brain
and by receptors on the surface of the body. The various cooling or heating
mechanisms of the body are then activated in accord with the temperature. The
efficiency of muscles in performing external work is at best 20%. Therefore, at
least 80% of the energy consumed in the performance of a physical activity is
converted into heat inside the body. In addition, the energy consumed to maintain
the basic metabolic processes is ultimately all converted to heat. If this heat
were not eliminated, the body temperature would quickly rise to a dangerous
level. For example, during moderate physical activity, a 70-kg man may consume
260 Cal/hr. Of this amount, at least 208 Cal is converted to heat. If this
heat remained within the body, the body temperature would rise by 3 C◦/hr.
Two hours of such an activity would cause complete collapse. Fortunately, the
body possesses a number of highly efficient methods for controlling the heat
flow out of the body, thereby maintaining a stable internal temperature.
Most of the heat generated by the body is produced deep in the body, far
from the surfaces. In order to be eliminated, this heat must first be conducted
to the skin. For heat to flow from one region to another, there must be a
temperature difference between the two regions. Therefore, the temperature
of the skin must be lower than the internal body temperature. In a warm
environment, the temperature of the human skin is about 35◦C. In a cold
environment, the temperature of some parts of the skin may drop to 27◦C.
The tissue of the body, without blood flowing through it, is a poor conductor.
Its thermal conductivity is comparable to that of cork (see Table 9.2).
(Kc for tissue without blood is 18 Cal-cm/m2-hr-C◦.) Simple thermal conductivity
through tissue is inadequate for elimination of the excess heat generated
by the body. The following calculation illustrates this point. Assume that the
thickness of the tissue between the interior and the exterior of the body is 3 cm
and that the average area through which conduction can occur is 1.5 m2. With
a temperature difference T between the inner body and the skin of 2◦C, the
heat flow H per hour is, from Eq. 9.3,
H
KcA T
L
18 � 1.5 � 2
3
18 Cal/hr (11.3)
In order to increase the conductive heat flow to a moderate level of say
150 Cal/hr, the temperature difference between the interior body and the skin
would have to increase to about 17 C◦.
Section 11.5 Convection 151
Fortunately the body possesses another method for transferring heat. Most
of the heat is transported from the inside of the body by blood in the circulatory
system. Heat enters the blood from an interior cell by conduction. In this case,
heat transfer by conduction is relatively fast because the distances between the
capillariesandtheheat-producingcellsaresmall. Thecirculatorysystemcarries
the heated blood near to the surface skin. The heat is then transferred to the
outside surface by conduction. In addition to transporting heat from the interior
of the body, the circulatory system controls the insulation thickness of the body.
When the heat flow out of the body is excessive, the capillaries near the surface
becomeconstrictedandthebloodflowtothesurfaceisgreatlyreduced. Because
tissue without blood is a poor heat conductor, this procedure provides a heatinsulating
layer around the inner body core.
11.4 Control of Skin Temperature
As was stated, for heat to flow out of the body, the temperature of the skin must
be lower than the internal body temperature. Therefore, heat must be removed
from the skin at a sufficient rate to ensure that this condition is maintained.
Because the heat conductivity of air is very low (2.02 Cal-cm/m2-hr-C◦), if
the air around the skin is confined—for example, by clothing—the amount
of heat removed by conduction is small. The surface of the skin is cooled
primarily by convection, radiation, and evaporation. However, if the skin is in
contact with a good thermal conductor such as a metal, a considerable amount
of heat can be removed by conduction (see Exercise 11-6).
11.5 Convection
When the skin is exposed to open air or some other fluid, heat is removed
from it by convection currents. The rate of heat removal is proportional to the
exposed surface area and to the temperature difference between the skin and
the surrounding air. The rate of heat transfer by convection Hc (see Eq. 9.4)
is given by
Hc KcAc(Ts − Ta) (11.4)
where Ac is the skin area exposed to the open air; Ts and Ta are the skin and
air temperatures, respectively; and Kc is the convection coefficient, which has
a value that depends primarily on the prevailing wind velocity. The value of
Kc as a function of air velocity is shown in Fig. 11.1. As the plot shows, the
convection coefficient initially increases sharply with wind velocity, and then
the increase becomes less steep (see Exercise 11-7).
152 Chapter 11 Heat and Life
FIGURE 11.1 Convection coefficient as a function of air velocity.
The exposed area Ac is generally smaller than the total surface area of the
body. For a naked person standing with legs together and arms close to the
body, about 80% of the surface area is exposed to convective air currents.
(The exposed area can be reduced by curling up the body.)
Note that heat flows from the skin to the environment only if the air is
colder than the skin. If the opposite is the case, the skin is actually heated by
the convective air flow.
Let us now calculate the amount of heat removed from the skin by convection.
Consider a naked person whose total surface area is 1.7 m2. Standing
straight, the exposed area is about 1.36 m2. If the air temperature is 25◦C and
the average skin temperature is 33◦C, the amount of heat removed is
Hc 1.36Kc � 8 10.9Kc Cal/hr
Under nearly windless conditions, Kc is about 6 Cal/m2-hr-C◦ (see Fig. 11.1),
and the convective heat loss is 65.4 Cal/hr. During moderate work, the energy
consumption for a person of this size is about 170 Cal/hr. Clearly, convection
in a windless environment does not provide adequate cooling. The wind
Section 11.7 Radiative Heating by the Sun 153
velocity has to increase to about 1.5 m/sec to provide cooling at a rate of
170 Cal/hr.
11.6 Radiation
Equation 9.6 shows that the energy exchange by radiation Hr involves the
fourth power of temperature; that is,
Hr eσ T 4
1 − T 4
2
However, because in the environment encountered by living systems the temperature
on the absolute scale seldom varies by more than 15%, it is possible to
use, without much error, a linear expression for the radiative energy exchange
(see Exercise 11-8a and b); that is,
Hr KrAre(Ts − Tr) (11.5)
where Ts and Tr are the skin surface temperature and the temperature of the
nearby radiating surface, respectively; Ar is the area of the body participating
in the radiation; e is the emissivity of the surface; and Kr is the radiation
coefficient. Over a fairly wide range of temperatures, Kr is, on the average,
about 6.0 Cal/m2-hr-C◦ (see Exercise 11-8c).
The environmental radiating surface and skin temperatures are such that
the wavelength of the thermal radiation is predominantly in the infrared region
of the spectrum. The emissivity of the skin in this wavelength range is nearly
unity, independent of the skin pigmentation. For a person with Ar 1.5 m2,
Tr 25◦C and Ts 32◦C. The radiative heat loss is 63 Cal/hr.
If the radiating surface is warmer than the skin surface, the skin is heated
by radiation. A person begins to feel discomfort due to radiation if the temperature
difference between the exposed skin and the radiating environment
exceeds about 6 C◦. In the extreme case, when the skin is illuminated by
the sun or some other very hot object like a fire, the skin is heated intensely.
Because the temperature of the source is now much higher than the temperature
of the skin, the simplified expression in Eq. 11.5 no longer applies.
11.7 Radiative Heating by the Sun
The intensity of solar energy at the top of the atmosphere is about 1150 Cal/
m2-hr. Not all this energy reaches the surface of the Earth. Some of it is
reflected by airborne particles and water vapor. A thick cloud cover may reflect
as much as 75% of solar radiation. The inclination of the Earth’s axis of
154 Chapter 11 Heat and Life
FIGURE 11.2 Radiative heating by the sun.
rotation further reduces the intensity of solar radiation at the surface. However,
in dry equatorial deserts, nearly all the solar radiation may reach the surface.
Because the rays of the sun come from one direction only, at most half
the body surface is exposed to solar radiation. In addition the area perpendicular
to the solar flux is reduced by the cosine of the angle of incidence
(see Fig. 11.2). As the sun approaches the horizon, the effective area for the
interception of radiation increases, but at the same time the radiation intensity
decreases because the radiation passes through a thicker layer of air. Still, the
amount of solar energy heating the skin can be very large. Assuming that the
full intensity of solar radiation reaches the surface, the amount of heat Hr that
the human body receives from solar radiation is
Hr 1150/2 � e � A cos θ Cal/hr (11.6)
Here A is the skin area of the person, θ is the angle of incidence of sunlight,
and e is the emissivity of the skin. The emissivity of the skin in the
wavelength region of solar radiation depends on the pigmentation. Dark skin
absorbs about 80% of the radiation, and light skin absorbs about 60%. From
Eq. 11.6, a light-skinned person with a skin area of 1.7 m2, subject to intense
solar radiation incident at a 60◦ angle, receives heat at the rate of 294 Cal/hr.
Radiative heating is decreased by about 40% if the person wears light-colored
clothing. Radiative heating is also reduced by changing the orientation of the
body with respect to the sun. Camels resting in the shadeless desert face the
sun, which minimizes the skin area exposed to solar radiation.
Section 11.8 Evaporation 155
11.8 Evaporation
In a warm climate, convection and radiation cannot adequately cool a person
engaged in even moderate physical activity. A large fraction of cooling is
provided by the evaporation of sweat from the skin surfaces. At normal skin
temperatures, the latent heat of vaporization for water is 0.580 Cal/g. Therefore,
about 580 Cal of heat are removed for each liter of sweat that evaporates
from the skin. The body contains two types of sweat glands, the eccrine
and the apocrine. The eccrine glands are distributed over the whole surface
of the body, and they respond primarily to the nerve impulses generated by
the thermoregulatory system of the body. As the heat load on the body rises,
the sweat secreted by these glands increases proportionately. There is an
exception to this. The eccrine glands in the palms of the hand and the soles
of the feet are stimulated by elevated levels of adrenaline in the blood, which
may result from emotional stress.
The apocrine sweat glands, found mostly in the pubic regions, are not
associated with temperature control. They are stimulated by adrenaline in the
blood stream, and they secrete a sweat rich in organic matter. The decomposition
of these substances produces body odor.
The ability of the human body to secrete sweat is remarkable. For brief
periods of time, a person can produce sweat at a rate up to 4 liter/hr. Such a
high rate of sweating, however, cannot be maintained. For longer periods, up
to 6 hours, a sweating rate of 1 liter/hr is common in the performance of heavy
work in a hot environment.
During prolonged heavy sweating, adequate amounts of water must be
drunk; otherwise, the body becomes dehydrated. A person’s functioning is
severely limited when dehydration results in a 10% loss of body weight. Some
desert animals can endure greater dehydration than humans; a camel, for
example, may lose water amounting to 30% of its body weight without serious
consequences.
Only sweat that evaporates is useful in cooling the skin. Sweat that
rolls off or is wiped off does not provide significant cooling. Nevertheless,
excess sweat does ensure full wetting of the skin. The amount of sweat
that evaporates from the skin depends on ambient temperature, humidity,
and air velocity. Evaporative cooling is most efficient in a hot, dry, windy
environment.
There is another avenue for evaporative heat loss: breathing. The air leaving
the lungs is saturated by water vapor from the moist lining of the respiratory
system. At a normal human breathing rate, the amount of heat removed
by this avenue is small, less than 9 Cal/hr (see Exercise 11-9); however, for
furred animals that do not sweat, this method of heat removal is very important.
These animals can increase heat loss by taking short shallow breaths
156 Chapter 11 Heat and Life
(panting) that do not bring excessive oxygen into the lungs but do pick up
moisture from the upper respiratory tract.
By evaporative cooling, a person can cope with the heat generated by moderate
activity even in a very hot, sunny environment. To illustrate this, we will
calculate the rate of sweating required for a person walking nude in the sun at
a rate of 3 mph, with the ambient temperature at 47◦C (116.6◦F).
With a skin area of 1.7 m2, the energy consumed in the act of walking is
about 240 Cal/hr. Almost all this energy is converted to heat and delivered to
the skin. In addition, the skin is heated by convection and by radiation from
the environment and the sun. The heat delivered to the skin by convection is
Hc KcAc(Ts − Ta)
For a 1-m/sec wind, Kc is 13 Cal/m2-hr-C◦. The exposed area Ac is about
1.5 m2. If the skin temperature is 36◦C,
Hc 13 � 1.50 � (47 − 36) 215 Cal/hr
As calculated previously, the radiative heating by the sun is about
294 Cal/hr. The radiative heating by the environment is
Hc KrAre(Tr − Ts) 6 � 1.5 � (47 − 36) 99 Cal/hr
In this example, the only mechanism available for cooling the body is
the evaporation of sweat. The total amount of heat that must be removed
is (240 + 215 + 194 + 99) Cal/hr 848 Cal/hr. The evaporation of about
1.5 liter/hr of sweat will provide the necessary cooling. Of course, if the
person is protected by light clothing, the heat load is significantly reduced.
The human body is indeed very well equipped to withstand heat. In controlled
experiments, people have survived a temperature of 125◦C for a period of time
that was adequate to cook a steak.
11.9 Resistance to Cold
In a thermally comfortable environment, the body functions at a minimum
expenditure of energy. As the environment cools, a point is reached where the
basal metabolic rate increases to maintain the body temperature at a proper
level. The temperature at which this occurs is called the critical temperature.
This temperature is a measure of the ability of an animal to withstand cold.
Human beings are basically tropical animals. Unprotected, they are much
better able to cope with heat than with cold. The critical temperature for
humans is about 30◦C. By contrast, the critical temperature for the heavily
furred arctic fox is −40◦C.
Section 11.9 Resistance to Cold 157
The discomfort caused by cold is due primarily to the increased rate of
heat outflow from the skin. This rate depends not only on the temperature but
also on the wind velocity and humidity. For example, at 20◦C, air moving
with a velocity of 30 cm/sec removes more heat than still air at 15◦C. In this
case, a mild wind at 30 cm/sec is equivalent to a temperature drop of more
than 5 C◦.
The body defends itself against cold by decreasing the heat outflow and by
increasing the production of heat. When the temperature of the body begins to
drop, the capillaries leading to the skin become constricted, reducing the blood
flow to the skin. This results in a thicker thermal insulation of the body. In a
naked person, this mechanism is fully utilized when the ambient temperature
drops to about 19◦C. At this point, the natural insulation cannot be increased
any more.
Additional heat required to maintain the body temperature is obtained by
increasing the metabolism. One involuntary response that achieves this is
shivering. As shown in Table 11.1, shivering raises the metabolism to about
250 Cal/m2-hr. If these defenses fail and the temperature of the skin and
underlying tissue fall below about 5◦C, frostbite and eventually more serious
freezing occur.
The most effective protection against cold is provided by thick fur, feathers,
or appropriate clothing. At −40◦C, without insulation, the heat loss is
primarily convective and radiative. By convection alone in moderately moving
air, the rate of heat removal per square meter of skin surface is about
660 Cal/m2-hr (see Exercise 11-10). With a thick layer of fur or similar insulation
the skin is shielded from convection and the heat is transferred to the
environment by conduction only. The thermal conductivity of insulating materials
such as fur or down is Kc 0.36 Cal cm/m2-hr-C◦; therefore, the heat
transfer from the skin at 30◦C to the ambient environment at −40◦C through
1 cm of insulation is, from Eq. 9.3, 25.2 Cal/m2-hr. This is below the basal
metabolic rate for most animals. Although body heat is lost also through radiation
and evaporation, our calculation indicates that well-insulated animals,
including a clothed person, can survive in cold environments.
As stated earlier, at moderate temperatures the amount of heat removed
by breathing at a normal rate is small. At very cold temperatures, however,
the heat removed by this channel is appreciable. Although the heat removed
by the evaporation of moisture from the lungs remains approximately constant,
the amount of heat required to warm the inspired air to body temperature
increases as the ambient air temperature drops. For a person at an
ambient temperature of −40◦C, the amount of heat removed from the body
in the process of breathing is about 14.4 Cal/hr (see Exercise 11-11). For a
well-insulated animal, this heat loss ultimately limits its ability to withstand
cold.
158 Chapter 11 Heat and Life
11.10 Heat and Soil
Much of life depends directly or indirectly on biological activities near the
surface of the soil. In addition to plants, there are worms and insects whose
lives are soil-bound (1 acre of soil may contain 500 kg of earthworms). Soil is
also rich in tiny organisms such as bacteria, mites, and fungi whose metabolic
activities are indispensable for the fertility of the soil. To all this life, the
temperature of the soil is of vital importance.
The surface soil is heated primarily by solar radiation. Although some
heat is conducted to the surface from the molten core of the Earth, the amount
from this source is negligible compared to solar heating. The Earth is cooled
by convection, radiation, and the evaporation of soil moisture. On the average,
over a period of a year, the heating and cooling are balanced; and therefore,
over this period of time, the average temperature of the soil does not change
appreciably. However, over shorter periods of time, from night to day, from
winter to summer, the temperature of the top soil changes considerably; these
fluctuations govern the life cycles in the soil.
The variations in soil temperature are determined by the intensity of solar
radiation, the composition and moisture content of the soil, the vegetation
cover, and atmospheric conditions such as clouds, wind, and airborne particles
(see Exercises 11-12 and 11-13). Certain patterns, however, are general.
During the day while the sun is shining, more heat is delivered to the soil
than is removed by the various cooling mechanisms. The temperature of the
soil surface therefore rises during the day. In dry soil, the surface temperature
may increase by 3 or 4 C◦/hr. The surface heating is especially intense in dry,
unshaded deserts. Some insects living in these areas have evolved long legs to
keep them removed from the hot surface.
The heat that enters the surface is conducted deeper into the soil. It takes
some time, however, for the heat to propagate through the soil. Measurements
show that a temperature change at the surface propagates into the soil at a
rate of about 2 cm/h. At night, the heat loss predominates and the soil surface
cools. The heat that was stored in the soil during the day now propagates to
the surface and leaves the soil. Because of the finite time required for the heat
to propagate through the soil, the temperature a few centimeters below the surface
may still be rising while the surface is already cooling off. Some animals
take advantage of this lag in temperature between the surface and the interior
of the soil. They burrow into the ground to avoid the larger temperature
fluctuations at the surface.
At the usual temperatures the thermal radiation emitted by the soil is in
the infrared region of the spectrum, which is strongly reflected by water vapor
and clouds. As a result, on cloudy days the thermal radiation emitted by the
soil is reflected back, and the net outflow of heat from the soil is reduced—this
Chapter 11 Exercises 159
FIGURE 11.3 The greenhouse effect.
is called the greenhouse effect (see Fig. 11.3). A similar effect is produced by
the “greenhouse gases” in the atmosphere principally carbon dioxide (CO2),
methane (CH4) and ozone (O3). These gases absorb infrared radiation and
emit it back to the earth’s surface increasing the temperature of the planet.
EXERCISES
11-1. Design an experiment that would measure the metabolic rate of walking
at 5 km/h up a 20◦ slope.
11-2. How long can a man survive in an airtight room that has a volume of
27 m3. Assume that his surface area is 1.70 m2. Use data provided in
the text.
11-3. A submarine carries an oxygen tank that holds oxygen at a pressure of
100 atm. What must be the volume of the tank to provide adequate oxygen
for 50 people for 10 days? Assume that daily energy expenditure
is as given in Table 11.2 and the average surface area of each person is
1.70 m2.
11-4. Calculate the length of time that a person can survive without food but
with adequate water. Obtain a solution under the following assumptions:
(a) The initial weight and surface area of the person are 70 kg
160 Chapter 11 Heat and Life
and 1.70 m2, respectively. (b) The survival limit is reached when the
person loses one-half his or her body weight. (c) Initially the body
contains 5 kg of fatty tissue. (d) During the fast the person sleeps
8 hr/day and rests quietly the remainder of the time. (e) As the person
loses weight, his or her surface area decreases (see Eq. 11.1). However,
here we assume that the surface area remains unchanged.
11-5. Suppose that a person of weight 60 kg and height 1.4 m reduces her
sleep by 1 hr/day and spends this extra time reading while sitting
upright. If her food intake remains unchanged, how much weight will
she lose in one year?
11-6. Assume that a person is sitting naked on an aluminum chair with
400-cm2 area of the skin in contact with aluminum. If the skin temperature
is 38◦C and the aluminum is kept at 25◦C, compute the amount
of heat transfer per hour from the skin. Assume that the body contacting
the aluminum is insulated by a layer of unperfused fat tissue
0.5 cm thick (Kc 18 Cal cm/m2-hr-C◦) and that the heat conductivity
of aluminum is very large. Is this heat transfer significant in terms
of the metabolic heat consumption?
11-7. Explain qualitatively the functional dependence of Kc on the air
velocity (see Fig. 11.1).
11-8. (a) Show that (T 4
s − T 4
r ) (T 3
s + T 2
s Tr + TsT 2
r + T 3
r )(Ts − Tr).
(b) Compute percentage change in the term (T 3
s + T 2
s Tr + TsT 2
r + T 3
r )
as the radiative temperature of the environment changes from 0 to
40◦C. (Note that the temperatures in the computations must be
expressed on the absolute scale. However, if the expression contains
only the difference between two temperatures, either the absolute or
the centigrade scale may be used.) (c) Calculate the value of Kr
in Eq. 11.5 under conditions discussed in the text where Tr 25◦C
(298 K), Ts 32◦C (305 K), and Hr 63 Cal/hr.
11-9. A person takes about 20 breaths per minute with 0.5 liter of air in
each breath. How much heat is removed per hour by the moisture in
the exhaled breath if the incoming air is dry and the exhaled breath is
fully saturated? Assume that the water vapor pressure in the saturated
exhaled air is 24 torr. Use data in the Section 11.8.
11-10. Compute the heat loss per square meter of skin surface at −40◦C in
moderate wind (about 0.5 m /sec, Kc 10 Cal/m2-hr-◦C). Assume
that the skin temperature is 26◦C.
11-11. Calculate the amount of heat required per hour to raise the temperature
of inspired air from −40◦C to the body temperature of 37◦C. Assume
that the breathing rate is 600 liters of air per hour. (This is the breathing
Chapter 11 Exercises 161
rate specified in Exercise 11-9.) The amount of heat required to raise
the temperature of 1 mole of air (22.4 liter) by 1 C◦ at 1 atm is 29.2 J
(6.98 � 10−3 Cal).
11-12. Explain why the daily temperature fluctuations in the soil are smaller
(a) in wet soil than in dry soil, (b) in soil with a grass growth than in
bare soil, and (c) when the air humidity is high.
11-13. Explain why the temperature drops rapidly at night in a desert.
11-14. The therapeutic effects of heat have been known since ancient times.
Local heating, for example, relieves muscle pain and arthritic conditions.
Discuss some effects of heat on tissue that may explain its
therapeutic value.
Chapter 12
Waves and Sound
Most of the information about our physical surroundings comes to us through
our senses of hearing and sight. In both cases we obtain information about
objects without being in physical contact with them. The information is transmitted
to us in the first case by sound, in the second case by light. Although
sound and light are very different phenomena, they are both waves. A wave
can be defined as a disturbance that carries energy from one place to another
without a transfer of mass. The energy carried by the waves stimulates our
sensory mechanisms.
In this chapter, we will first explain briefly the nature of sound and then
review some general properties of wave motion applicable to both sound and
light. Using this background we will examine the process of hearing and some
other biological aspects of sound. Light will be discussed in Chapter 15.
12.1 Properties of Sound
Sound is a mechanical wave produced by vibrating bodies. For example, when
an object such as a tuning fork or the human vocal cords is set into vibrational
motion, the surrounding air molecules are disturbed and are forced to follow
the motion of the vibrating body. The vibrating molecules in turn transfer their
motion to adjacent molecules causing the vibrational disturbance to propagate
away from the source. When the air vibrations reach the ear, they cause the
eardrum to vibrate; this produces nerve impulses that are interpreted by the
brain.
162
Section 12.1 Properties of Sound 163
All matter transmits sound to some extent, but a material medium is needed
between the source and the receiver to propagate sound. This is demonstrated
by the well-known experiment of the bell in the jar. When the bell is set in
motion, its sound is clearly audible. As the air is evacuated from the jar, the
sound of the bell diminishes and finally the bell becomes inaudible.
The propagating disturbance in the sound-conducting medium is in the
form of alternate compressions and rarefactions of the medium, which are
initially caused by the vibrating sound source. These compressions and
rarefactions are simply deviations in the density of the medium from the
average value. In a gas, the variations in density are equivalent to pressure
changes.
Two important characteristics of sound are intensity, which is determined
by the magnitude of compression and rarefaction in the propagating medium,
and frequency, which is determined by how often the compressions and rarefactions
take place. Frequency is measured in cycles per second, which is
designated by the unit hertz after the scientist Heinrich Hertz. The symbol for
this unit is Hz. (1 Hz 1 cycle per second.)
The vibrational motion of objects can be highly complex (see Fig. 12.1),
resulting in a complicated sound pattern. Still, it is useful to analyze the
properties of sound in terms of simple sinusoidal vibrations such as would
be set up by a vibrating tuning fork (see Fig. 12.2). The type of simple sound
FIGURE 12.1 A complex vibrational pattern.
164 Chapter 12 Waves and Sound
FIGURE 12.2 Sinusoidal sound wave produced by a vibrating tuning fork.
pattern shown in Fig. 12.2 is called a pure tone. When a pure tone propagates
through air, the pressure variations due to the compressions and rarefactions
are sinusoidal in form.
If we were to take a “snapshot” of the sound at a given instant in time,
we would see pressure variations in space, which are also sinusoidal. (Such
pictures can actually be obtained with special techniques.) In such a picture
the distance between the nearest equal points on the sound wave is called the
wavelength λ.
The speed of the sound wave v depends on the material that propagates
the sound. In air at 20◦C, the speed of sound is about 3.3 � 104 cm/sec, and
in water it is about 1.4 � 105 cm/sec. In general, the relationship between
frequency, wavelength, and the speed of propagation is given by the following
equation:
v λf (12.1)
This relationship between frequency, wavelength, and speed is true for all
types of wave motions.
The pressure variations due to the propagating sound are superimposed on
the ambient air pressure. Thus, the total pressure in the path of a sinusoidal
sound wave is of the form
P Pa + Po sin 2πft (12.2)
Section 12.2 Some Properties of Waves 165
where Pa is the ambient air pressure (which at sea level at 0◦C is 1.01 �
105Pa 1.01 � 106 dyn/cm2), Po is the maximum pressure change due to
the sound wave, and f is the frequency of the sound. The amount of energy
transmitted by a sinusoidal sound wave per unit time through each unit area
perpendicular to the direction of sound propagation is called the intensity I
and is given by
I
P2
o
2ρv
(12.3)
Here ρ is the density of the medium, and v is the speed of sound propagation.
12.2 Some Properties of Waves
All waves, including sound and light, exhibit the phenomena of reflection,
refraction, interference, and diffraction. These phenomena, which play an
important role in both hearing and seeing, are described in detail in most basic
physics texts (see [12-8]). Here we will review them only briefly.
12.2.1 Reflection and Refraction
When a wave enters one medium from another, part of the wave is reflected at
the interface, and part of it enters the medium. If the interface between the two
media is smooth on the scale of the wavelength (i.e., the irregularities of the
interface surface are smaller than λ), the reflection is specular (mirrorlike). If
the surface has irregularities that are larger than the wavelength, the reflection
is diffuse. An example of diffuse reflection is light reflected from paper.
If the wave is incident on the interface at an oblique angle, the direction
of propagation of the transmitted wave in the new medium is changed (see
Fig. 12.3). This phenomenon is called refraction. The angle of reflection is
always equal to the angle of incidence, but the angle of the refracted wave is,
in general, a function of the properties of the two media. The fraction of the
energy transmitted from one medium to another depends again on the properties
of the media and on the angle of incidence. For a sound wave incident
perpendicular to the interface, the ratio of transmitted to incident intensity is
given by
It
Ii
4ρ1v1ρ2v2
(ρ1v1 + ρ2v2)2
(12.4)
where the subscripted quantities are the velocity and density in the two media.
The solution of Eq. 12.4 shows that when sound traveling in air is incident
166 Chapter 12 Waves and Sound
FIGURE 12.3 Illustration of reflection and refraction. (θ is the angle of incidence.)
perpendicular to a water surface, only about 0.1% of the sound energy enters
the water; 99.9% is reflected. The fraction of sound energy entering the water
is even smaller when the angle of incidence is oblique. Water is thus an efficient
barrier to sound.
12.2.2 Interference
When two (or more) waves travel simultaneously in the same medium, the
total disturbance in the medium is at each point the vectorial sum of the
individual disturbances produced by each wave. This phenomenon is called
interference. For example, if two waves are in phase, they add so that the
wave disturbance at each point in space is increased. This is called constructive
interference (see Fig. 12.4a). If two waves are out of phase by 180◦,
the wave disturbance in the propagating medium is reduced. This is called
destructive interference (Fig. 12.4b). If the magnitudes of two out-of-phase
waves are the same, the wave disturbance is completely canceled (Fig. 12.4c).
A special type of interference is produced by two waves of the same frequency
and magnitude traveling in opposite directions. The resultant wave
Section 12.2 Some Properties of Waves 167
FIGURE 12.4 (a) Constructive interference. (b, c) Destructive interference. R is
the resultant of the interference of the two waves A and B.
pattern is stationary in space and is called a standing wave. Such standing
sound waves are formed in hollow pipes such as the flute. It can be shown
that, in a given structure, standing waves can exist only at specific frequencies,
which are called resonant frequencies.
168 Chapter 12 Waves and Sound
12.2.3 Diffraction
Waves have a tendency to spread as they propagate through a medium. As a
result, when a wave encounters an obstacle, it spreads into the region behind
the obstacle. This phenomenon is called diffraction. The amount of diffraction
depends on the wavelength: The longer the wavelength, the greater is
the spreading of the wave. Significant diffraction into the region behind the
obstacle occurs only if the size of the obstacle is smaller than the wavelength.
For example, a person sitting behind a pillar in an auditorium hears the performer
because the long wavelength sound waves spread behind the pillar. But
the view of the performance is obstructed because the wavelength of light is
much smaller than the pillar, and, therefore, the light does not diffract into the
region behind the pillar.
Objects that are smaller than the wavelength do not produce a significant
reflection. This too is due to diffraction. The wave simply diffracts around the
small obstacle, much as flowing water spreads around a small stick.
Both light waves and sound waves can be focused with curved reflectors
and lenses. There is, however, a limit to the size of the focused spot. It can
be shown that the diameter of the focused spot cannot be smaller than about
λ/2. These properties of waves have important consequences in the process
of hearing and seeing.
12.3 Hearing and the Ear
The sensation of hearing is produced by the response of the nerves in the ear
to pressure variations in the sound wave. The nerves in the ear are not the
only ones that respond to pressure, as most of the skin contains nerves that
are pressure-sensitive. However, the ear is much more sensitive to pressure
variations than any other part of the body.
Figure 12.5 is a drawing of the human ear. (The ear construction of other
terrestrial vertebrates is similar.) For the purposes of description, the ear is
usually divided into three main sections: the outer ear, the middle ear, and the
inner ear. The sensory cells that convert sound to nerve impulses are located
in the liquid-filled inner ear.
The main purpose of the outer and middle ears is to conduct the sound into
the inner ear.
The outer ear is composed of an external flap called the pinna and the ear
canal, which is terminated by the tympanic membrane (eardrum). In many
animals the pinna is large and can be rotated toward the source of the sound;
this helps the animal to locate the source of sound. However, in humans the
pinna is fixed and so small that it does not seem to contribute significantly to
the hearing process.
Section 12.3 Hearing and the Ear 169
FIGURE 12.5 A semidiagrammatic drawing of the ear with various structures cut
away and simplified to show the basic relationships more clearly. The middle ear
muscles have been omitted.
The ear canal of an average adult is about 0.75 cm in diameter and 2.5 cm
long, a configuration that is resonant for sound waves at frequencies around
3000 Hz. This accounts in part for the high sensitivity of the ear to sound
waves in this frequency range.
For an animal to perceive sound, the sound has to be coupled from air to
the sensory cells that are in the fluid environment of the inner ear. We showed
earlier that direct coupling of sound waves into a fluid is inefficient because
most of the sound energy is reflected at the interface. The middle ear provides
an efficient conduction path for the sound waves from air into the fluid of the
inner ear.
The middle ear is an air-filled cavity that contains a linkage of three bones
called ossicles that connect the eardrum to the inner ear. The three bones are
called the hammer, the anvil, and the stirrup. The hammer is attached to the
inner surface of the eardrum, and the stirrup is connected to the oval window,
which is a membrane-covered opening in the inner ear.
170 Chapter 12 Waves and Sound
When sound waves produce vibrations in the eardrum, the vibrations are
transmitted by the ossicles to the oval window, which in turn sets up pressure
variations in the fluid of the inner ear. The ossicles are connected to the walls
of the middle ear by muscles that also act as a volume control. If the sound
is excessively loud, these muscles as well as the muscles around the eardrum
stiffen and reduce the transmission of sound to the inner ear.
The middle ear serves yet another purpose. It isolates the inner ear from
the disturbances produced by movements of the head, chewing, and the internal
vibrations produced by the person’s own voice. To be sure, some of the
vibrations of the vocal cords are transmitted through the bones into the inner
ear, but the sound is greatly attenuated. We hear ourselves talk mostly by
the sound reaching our eardrums from the outside. This can be illustrated by
talking with the ears plugged.
The Eustachian tube connects the middle ear to the upper part of the throat.
Air seeps in through this tube to maintain the middle ear at atmospheric pressure.
The movement of air through the Eustachian tube is aided by swallowing.
A rapid change in the external air pressure such as may occur during an
airplane flight causes a pressure imbalance on the two sides of the eardrum.
The resulting force on the eardrum produces a painful sensation that lasts until
the pressure in the middle ear is adjusted to the external pressure. The pain is
especially severe and prolonged if the Eustachian tube is blocked by swelling
or infection.
The conversion of sound waves into nerve impulses occurs in the cochlea,
which is located in the inner ear. The cochlea is a spiral cavity shaped like
a snail shell. The wide end of the cochlea, which contains the oval and the
round windows, has an area of about 4 mm2. The cochlea is formed into a
spiral with about 23
4 turns. If the cochlea were uncoiled, its length would be
about 35 mm.
Inside the cochlea there are three parallel ducts; these are shown in the
highly simplified drawing of the uncoiled cochlea in Fig. 12.6. All three ducts
are filled with a fluid. The vestibular and tympanic canals are joined at the
apex of the cochlea by a narrow opening called the helicotrema. The cochlear
duct is isolated from the two canals by membranes. One of these membranes,
called the basilar membrane, supports the auditory nerves.
The vibrations of the oval window set up a sound wave in the fluid filling
the vestibular canal. The sound wave, which travels along the vestibular canal
and through the helicotrema into the tympanic canal, produces vibrations in
the basilar membrane which stimulate the auditory nerves to transmit electrical
pulses to the brain (see Chapter 13). The excess energy in the sound wave
is dissipated by the motion of the round window at the end of the tympanic
canal.
Section 12.3 Hearing and the Ear 171
FIGURE 12.6 An uncoiled view of the cochlea.
12.3.1 Performance of the Ear
The nerve impulses evoke in the brain the subjective sensation of sound.
Loudness, pitch, and quality are some of the terms we use to describe the
sounds we hear. It is a great challenge for physiologists to relate these subjective
responses with the physical properties of sound such as intensity and
frequency. Some of these relationships are now well understood; others are
still subjects for research.
In most cases, the sound wave patterns produced by instruments and voices
are highly complex. Each sound has its own characteristic pattern. It would
be impossible to evaluate the effect of sound waves on the human auditory
system if the response to each sound pattern had to be analyzed separately.
Fortunately the problem is not that complicated. About 150 years ago, J. B.
J. Fourier, a French mathematician, showed that complex wave shapes can
be analyzed into simple sinusoidal waves of different frequencies. In other
words, a complex wave pattern can be constructed by adding together a sufficient
number of sinusoidal waves at appropriate frequencies and amplitudes.
Therefore, if we know the response of the ear to sinusoidal waves over a broad
range of frequencies, we can evaluate the response of the ear to a wave pattern
of any complexity.
An analysis of a wave shape into its sinusoidal components is shown in
Fig. 12.7. The lowest frequency in the wave form is called the fundamental,
and the higher frequencies are called harmonics. Figure 12.8, shows the
sound pattern for a specific note played by various instruments. It is the harmonic
content of the sound that differentiates one sound source from another.
172 Chapter 12 Waves and Sound
FIGURE 12.7 The analysis of a complex wave shape (a), into its sine components
(b). The point-by-point addition of the fundamental frequency sine wave and the
harmonic frequency sine waves yields the wave shape shown in (a).
For a given note played by the various instruments shown in Fig. 12.8, the
fundamental frequency is the same but the harmonic content of the wave is
different for each instrument.
12.3.2 Frequency and Pitch
The human ear is capable of detecting sound at frequencies between about
20 and 20,000 Hz. Within this frequency range, however, the response of the
ear is not uniform. The ear is most sensitive to frequencies between 200 and
4000 Hz, and its response decreases toward both higher and lower frequencies.
There are wide variations in the frequency response of individuals. Some
people cannot hear sounds above 8000 Hz, whereas a few people can hear
sounds above 20,000 Hz. Furthermore, the hearing of most people deteriorates
with age.
The sensation of pitch is related to the frequency of the sound. The pitch
increases with frequency. Thus, the frequency of middle C is 256 Hz, and the
Section 12.3 Hearing and the Ear 173
FIGURE 12.8 Wave forms of sound from different musical instruments sounding
the same note.
frequency of the A above is 440 Hz. There is, however, no simple mathematical
relationship between pitch and frequency.
12.3.3 Intensity and Loudness
The ear responds to an enormous range of intensities. At 3000 Hz, the lowest
intensity that the human ear can detect is about 10−16 W/cm2. The loudest
tolerable sound has an intensity of about 10−4 W/cm2. These two extremes
of the intensity range are called the threshold of hearing and the threshold of
pain, respectively. Sound intensities above the threshold of pain may cause
permanent damage to the eardrum and the ossicles.
The ear does not respond linearly to sound intensity; that is, a sound which
is a million times more powerful than another does not evoke a million times
higher sensation of loudness. The response of the ear to intensity is closer to
being logarithmic than linear.
Because of the nonlinear response of the ear and the large range of intensities
involved in the process of hearing, it is convenient to express sound
intensity on a logarithmic scale. On this scale, the sound intensity is measured
relative to a reference level of 10−16 W/cm2 (which is approximately the
174 Chapter 12 Waves and Sound
TABLE 12.1 Sound Levels Due to Various
Sources (representative values)
Sound level Sound level
Source of sound (dB) (W/cm2)
Threshold of pain 120 10−4
Riveter 90 10−7
Busy street traffic 70 10−9
Ordinary conversation 60 10−10
Quiet automobile 50 10−11
Quiet radio at home 40 10−12
Average whisper 20 10−14
Rustle of leaves 10 10−15
Threshold of hearing 0 10−16
lowest audible sound intensity). The logarithmic intensity is measured in units
of decibel (dB) and is defined as
Logarithmic intensity 10 log
Sound intensity in W/cm2
10−16 W/cm2
(12.5)
Thus, for example, the logarithmic intensity of a sound wave with a power
of 10−12 W/cm2 is
Logarithmic intensity 10 log
10−12
10−16
40 dB
Intensities of some common sounds are listed in Table 12.1.
At one time, it was believed that the ear responded logarithmically to
sound intensity. Referring to Table 12.1, a logarithmic response would imply
that, for example, a busy street sounds only six times louder than the rustle of
leaves even though the power of the street sounds is a million times greater.
Although it has been shown that the intensity response of the ear is not exactly
logarithmic, the assumption of a logarithmic response still provides a useful
guide for assessing the sensation of loudness produced by sounds at different
intensities (see Exercises 12-1 and 12-2).
The sensitivity of the ear is remarkable. At the threshold of hearing, in the
range of 2000–3000 Hz, the ear can detect a sound intensity of 10−16 W/cm2.
This corresponds to a pressure variation in the sound wave of only about
2.9 � 10−4 dyn/cm2 (see Exercise 12-3). Compare this to the background
atmospheric pressure, which is 1.013 � 106 dyn/cm2. This sensitivity appears
Section 12.4 Bats and Echoes 175
even more remarkable when we realize that the random pressure variations
in air due to the thermal motion of molecules are about 0.5 � 10−4 dyn/cm2.
Thus, the sensitivity of the ear is close to the ultimate limit at which it would
begin to detect the noise fluctuations in the air. The displacement of the
molecules corresponding to the power at the threshold of hearing is less than
the size of the molecules themselves.
The sensitivity of the ear is partly due to the mechanical construction of
the ear, which amplifies the sound pressure. Most of the mechanical amplification
is produced by the middle ear. The area of the eardrum is about 30
times larger than the oval window. Therefore, the pressure on the oval window
is increased by the same factor (see Exercise 12-4). Furthermore, the ossicles
act as a lever with a mechanical advantage of about 2. Finally, in the frequency
range around 3000 Hz, there is an increase in the pressure at the eardrum due
to the resonance of the ear canal. In this frequency range, the pressure is
increased by another factor of 2. Thus, the total mechanical amplification
of the sound pressure in the 3000-Hz range is about 2 � 30 � 2 120.
Because the intensity is proportional to pressure squared (see Eq. 12.3), the
intensity at the oval window is amplified by a factor of about 14,400.
The process of hearing cannot be fully explained by the mechanical construction
of the ear. The brain itself plays an important role in our perception
of sound. For example, the brain can effectively filter out ambient noise and
allow us to separate meaningful sounds from a relatively loud background din.
(This feature of the brain permits us to have a private conversation in the midst
of a loud party.) The brain can also completely suppress sounds that appear to
be meaningless. Thus, we may lose awareness of a sound even though it still
produces vibrations in our ear. The exact mechanism of interaction between
the brain and the sensory organs is not yet fully understood.
12.4 Bats and Echoes
The human auditory organs are very highly developed; yet, there are animals
that can hear even better than we can. Notable among these animals are the
bats. They emit high-frequency sound waves and detect the reflected sounds
(echoes) from surrounding objects. Their sense of hearing is so acute that they
can obtain information from echoes which is in many ways as detailed as the
information we can obtain with our sense of sight. The many different species
of bats utilize echoes in various ways. The Vespertilionidae family of bats emit
short chirps as they fly. The chirps last about 3 � 10−3 sec (3 msec) with a
time interval between chirps of about 70 msec. Each chirp starts at a frequency
of about 100 � 103 Hz and falls to about 30 � 103 Hz at the end. (The ears
of bats, of course, respond to these high frequencies.) The silent interval
176 Chapter 12 Waves and Sound
between chirps allows the bat to detect the weak echo without interference
from the primary chirp. Presumably the interval between the chirp and the
return echo allows the bat to determine its distance from the object. It is also
possible that differences in the frequency content of the chirp and the echo
allow the bat to estimate the size of the object (see Exercise 12-5). With a
spacing between chirps of 70 msec, an echo from an object as far as 11.5 m
can be detected before the next chirp (see Exercise 12-6). As the bat comes
closer to the object (such as an obstacle or an insect), both the duration of and
the spacing between chirps decrease, allowing the bat to localize the object
more accurately. In the final approach to the object, the duration of the chirps
is only about 0.3 msec, and the spacing between them is about 5 msec.
Experiments have shown that with echo location bats can avoid wire obstacles
with diameters down to about 0.1 mm, but they fail to avoid finer wires.
This is in accord with our discussion of wave diffraction (see Exercise 12-7).
Other animals, such as porpoises, whales, and some birds, also use echoes to
locate objects, but they are not able to do so as well as bats.
12.5 Sounds Produced by Animals
Animals can make sounds in various ways. Some insects produce sounds by
rubbing their wings together. The rattlesnake produces its characteristic sound
by shaking its tail. In most animals, however, sound production is associated
with the respiratory mechanism. In humans, the vocal cords are the primary
source of sound. These are two reeds, shaped like lips, attached to the upper
part of the trachea. During normal breathing the cords are wide open. To produce
a sound the edges of the cords are brought together. Air from the lungs
passes through the space between the edges and sets the cords into vibration.
The frequency of the sounds is determined by the tension on the vocal cords.
The fundamental frequency of the average voice is about 140 Hz for males and
about 230 Hz for females. The sound produced by the vocal cords is substantially
modified as it travels through the passages of the mouth and throat. The
tongue also plays an important role in the final sound. Many voice sounds are
produced outside the vocal cords (for example, the consonant s). The sounds
in a whispering talk are also produced outside the vocal cords.
12.6 Acoustic Traps
Electronically generated sounds that mimic those of animals and insects are
increasingly being used as lures to trap the creatures. Electronic fishing lures
are now commercially available. One such device mimics the distress call of
a mackerel and attracts marlin and other larger fish to the fishhook.
Section 12.8 Ultrasonic Waves 177
To obtain baseline data on bat populations, often the bats have to be
captured and examined. In one such study, the social call of a rare Bechstein’s
bat that inhabits the woodlands of southeast England was synthesized luring
the bats into the net. (The bats were released after examination.)
The Mediterranean fruit fly commonly called medfly is a pest that infests
fruits and other crops causing on the order of $1 billion damage worldwide.
At present, spraying of pesticides is the most common way of controlling the
medfly. An environmentally more friendly way of controlling the pest has
been sought for many years. Sound traps under development may provide a
viable alternative. The male medfly produces with its wings a vibration at a
fundamental frequency of about 350 Hz accompanied by complex harmonics.
The female medflys are attracted to this courtship call and can be lured into
a trap.
12.7 Clinical Uses of Sound
The most familiar clinical use of sound is in the analysis of body sounds with
a stethoscope. This instrument consists of a small bell-shaped cavity attached
to a hollow flexible tube. The bell is placed on the skin over the source of the
body sound (such as the heart or lungs). The sound is then conducted by the
pipe to the ears of the examiner who evaluates the functioning of the organ.
A modified version of the stethoscope consists of two bells that are placed on
different parts of the body. The sound picked up by one bell is conducted to
one ear, and the sound from the other bell is conducted to the other ear. The
two sounds are then compared. With this device, it is possible, for example, to
listen simultaneously to the heartbeats of the fetus and of the pregnant mother.
12.8 Ultrasonic Waves
With special electronically driven crystals, it is possible to produce mechanical
waves at very high frequencies, up to millions of cycles per second. These
waves, which are simply the extension of sound to high frequencies, are called
ultrasonic waves. Because of their short wavelength, ultrasonic waves can be
focused onto small areas and can be imaged much as visible light (see Exercise
12-8).
Ultrasonic waves penetrate tissue and are scattered and absorbed within it.
Using specialized techniques called ultrasound imaging, it is possible to form
visible images of ultrasonic reflections and absorptions. Therefore, structures
within living organisms can be examined with ultrasound, as with X-rays.
Ultrasonic examinations are safer than X-rays and often can provide as much
178 Chapter 12 Waves and Sound
information. In some cases, such as in the examination of a fetus and the heart,
ultrasonic methods can show motion, which is very useful in such displays.
The frequency of sound detected by an observer depends on the relative
motion between the source and the observer. This phenomenon is called the
Doppler effect. It can be shown (see Exercise 12-9) that if the observer is
stationary and the source is in motion, the frequency of the sound f detected
by the observer is given by
f f
v
v ∓ vs
(12.6)
where f is the frequency in the absence of motion, v is the speed of sound,
and vs is the speed of the source. The minus sign in the denominator is to be
used when the source is approaching the observer, and the plus sign when the
source is receding.
Using the Doppler effect, it is possible to measure motions within a body.
One device for obtaining such measurements is the ultrasonic flow meter,
which produces ultrasonic waves that are scattered by blood cells flowing
in the blood vessels. The frequency of the scattered sound is altered by the
Doppler effect. The velocity of blood flow is obtained by comparing the incident
frequency with the frequency of the scattered ultrasound.
Within the tissue, the mechanical energy in the ultrasonic wave is converted
to heat. With a sufficient amount of ultrasonic energy, it is possible to
heat selected parts of a patient’s body more efficiently and evenly than can be
done with conventional heat lamps. This type of treatment, called diathermy,
is used to relieve pain and promote the healing of injuries. It is actually possible
to destroy tissue with very high-intensity ultrasound. Ultrasound is now
routinely used to destroy kidney and gall stones (lithotripsy).
EXERCISES
12-1. The intensity of a sound produced by a point source decreases as the
square of the distance from the source. Consider a riveter as a point
source of sound and assume that the intensities listed in Table 12.1 are
measured at a distance 1 m away from the source. What is the maximum
distance at which the riveter is still audible? (Neglect losses due to
energy absorption in the air.)
12-2. Referring to Table 12.1, approximately how much louder does busy
street traffic sound than a quiet radio?
Chapter 12 Exercises 179
12-3. Calculate the pressure variation corresponding to a sound intensity of
10−16 W/cm2. (The density of air at 0◦C and 1 atm pressure is 1.29 �
10−3 g/cm3; for the speed of sound use the value 3.3 � 104 cm/sec.)
12-4. Explain why the relative sizes of the eardrum and the oval window result
in pressure magnification in the inner ear.
12-5. Explain how a bat might use the differences in the frequency content of
its chirp and echo to estimate the size of an object.
12-6. With a 70-msec space between chirps, what is the farthest distance at
which a bat can detect an object?
12-7. In terms of diffraction theory, discuss the limitations on the size of the
object that a bat can detect with its echo location.
12-8. Estimate the lower limit on the size of objects that can be detected with
ultrasound at a frequency of 2 � 106 Hz.
12-9. With the help of a basic physics textbook, explain the Doppler effect
and derive Eq. 12.6.
Chapter 13
Electricity
The word electricity usually evokes the image of a man-made technology
because we usually associate electricity with devices such as amplifiers, televisions,
and computers. This technology has certainly played an important role
in our understanding of living systems, as it has provided the major tools for
the study of life processes. However, many life processes themselves involve
electrical phenomena. The nervous system of animals and the control of muscle
movement, for example, are both governed by electrical interactions. Even
plants rely on electrical forces for some of their functions. In this chapter,
we will describe some of the electrical phenomena in living organisms, and in
Chapter 14 we will discuss the applications of electrical technology in biology
and medicine. A brief review of electricity in Appendix B summarizes the
concepts, definitions, and equations used in the text.
13.1 The Nervous System
The most remarkable use of electrical phenomena in living organisms is found
in the nervous system of animals. Specialized cells called neurons form a
complex network within the body which receives, processes, and transmits
information from one part of the body to another. The center of this network
is located in the brain, which has the ability to store and analyze information.
Based on this information, the nervous system controls various parts of the
body. The nervous system is very complex. The human nervous system, for
example, consists of about 1010 interconnected neurons. It is, therefore, not
surprising that, although the nervous system has been studied for more than
180
Section 13.1 The Nervous System 181
a hundred years, its functioning as a whole is still poorly understood. It is
not known how information is stored and processed by the nervous system;
nor is it known how the neurons grow into patterns specific to their functions.
Yet some aspects of the nervous system are now well known. Specifically,
during the past 40 years, the method of signal propagation through the nervous
system has been firmly established. The messages are electrical pulses
transmitted by the neurons. When a neuron receives an appropriate stimulus,
it produces electrical pulses that are propagated along its cablelike structure.
The pulses are constant in magnitude and duration, independent of the intensity
of the stimulus. The strength of the stimulus is conveyed by the number of
pulses produced. When the pulses reach the end of the “cable,” they activate
other neurons or muscle cells.
13.1.1 The Neuron
The neurons, which are the basic units of the nervous system, can be divided
into three classes: sensory neurons, motor neurons, and interneurons. The
sensory neurons receive stimuli from sensory organs that monitor the external
and internal environment of the body. Depending on their specialized functions,
the sensory neurons convey messages about factors such as heat, light,
pressure, muscle tension, and odor to higher centers in the nervous system for
processing. The motor neurons carry messages that control the muscle cells.
These messages are based on information provided by the sensory neurons and
by the central nervous system located in the brain. The interneurons transmit
information between neurons.
Each neuron consists of a cell body to which are attached input ends called
dendrites and a long tail called the axon which propagates the signal away
from the cell (see Fig. 13.1). The far end of the axon branches into nerve
endings that transmit the signal across small gaps to other neurons or muscle
cells. A simple sensory-motor neuron circuit is shown in Fig. 13.2. A stimulus
from a muscle produces nerve impulses that travel to the spine. Here the signal
is transmitted to a motor neuron, which in turn sends impulses to control the
muscle. Such simple circuits are often associated with reflex actions. Most
nervous connections are far more complex.
The axon, which is an extension of the neuron cell, conducts the electrical
impulses away from the cell body. Some axons are long indeed—in people,
for example, the axons connecting the spine with the fingers and toes are more
than a meter in length. Some of the axons are covered with a segmented sheath
of fatty material called myelin. The segments are about 2 mm long, separated
by gaps called the Nodes of Ranvier. We will show later that the myelin sheath
increases the speed of pulse propagation along the axon.
182 Chapter 13 Electricity
FIGURE 13.1 A neuron.
FIGURE 13.2 A simple neural circuit.
Section 13.1 The Nervous System 183
Although each axon propagates its own signal independently, many axons
often share a common path within the body. These axons are usually grouped
into nerve bundles.
The ability of the neuron to transmit messages is due to the special electrical
characteristics of the axon. Most of the data about the electrical and
chemical properties of the axon is obtained by inserting small needlelike
probes into the axon. With such probes it is possible to measure currents
flowing in the axon and to sample its chemical composition. Such experiments
are usually difficult as the diameter of most axons is very small. Even
the largest axons in the human nervous system have a diameter of only about
20 μm (20 � 10−4 cm). The squid, however, has a giant axon with a diameter
of about 500 μm (0.5 mm), which is large enough for the convenient insertion
of probes. Much of the information about signal transmission in the nervous
system has been obtained from experiments with the squid axon.
13.1.2 Electrical Potentials in the Axon
In the aqueous environment of the body, salt and various other molecules dissociate
into positive and negative ions. As a result, body fluids are relatively
good conductors of electricity. Still, these fluids are not nearly as conductive
as metals; their resistivity is about 100 million times greater than that of
copper, for example.
The inside of the axon is filled with an ionic fluid that is separated from
the surrounding body fluid by a thin membrane (Fig. 13.3). The axon membrane,
which is only about 50–100 ˚A thick, is a relatively good but not perfect
electrical insulator. Therefore, some current can leak through it.
The electrical resistivities of the internal and the external fluids are about
the same, but their chemical compositions are substantially different. The
FIGURE 13.3 The axon membrane and surroundings.
184 Chapter 13 Electricity
external fluid is similar to sea water. Its ionic solutes are mostly positive
sodium ions and negative chlorine ions. Inside the axon, the positive ions
are mostly potassium ions, and the negative ions are mostly large negatively
charged organic molecules.
Because there is a large concentration of sodium ions outside the axon
and a large concentration of potassium ions inside the axon, we may ask
why the concentrations are not equalized by diffusion. In other words, why
don’t the sodium ions leak into the axon and the potassium ions leak out of it?
The answer lies in the properties of the axon membrane.
In the resting condition, when the axon is not conducting an electrical
pulse, the axon membrane is highly permeable to potassium and only slightly
permeable to sodium ions. The membrane is impermeable to the large organic
ions. Thus, while sodium ions cannot easily leak in, potassium ions can certainly
leak out of the axon. However, as the potassium ions leak out of the
axon, they leave behind the large negative ions, which cannot follow them
through the membrane. As a result, a negative potential is produced inside
the axon with respect to the outside. This negative potential, which has been
measured to be about 70 mV, holds back the outflow of potassium so that in
equilibrium the concentration of ions is as we have stated. Some sodium ions
do in fact leak into the axon, but they are continuously removed by a metabolic
mechanism called the sodium pump. This pumping process, which is not yet
fully understood, transports sodium ions out of the cell and brings in an equal
number of potassium ions.
13.1.3 Action Potential
The description of the axon that we have so far given applies to other types
of cells as well. Most cells contain an excess concentration of potassium ions
and are at a negative potential with respect to their surroundings. The special
property of the neuron is its ability to conduct electrical impulses.
Physiologists have studied the properties of nerve impulses by inserting
a probe into the axon and measuring the changes in the axon voltage with
respect to the surrounding fluid. The nerve impulse is elicited by some stimulus
on the neuron or the axon itself. The stimulus may be an injected chemical,
mechanical pressure, or an applied voltage. In most experiments the stimulus
is an externally applied voltage, as shown in Fig. 13.4.
A nerve impulse is produced only if the stimulus exceeds a certain threshold
value. When this value is exceeded, an impulse is generated at the point
of stimulation and propagates down the axon. Such a propagating impulse is
called an action potential. An action potential as a function of time at one
point on the axon is shown in Fig. 13.5. The scales of time and voltage are
typical of most neurons. The arrival of the impulse is marked by a sudden
Section 13.1 The Nervous System 185
FIGURE 13.4 Measuring the electrical response of the axon.
FIGURE 13.5 The action potential.
rise of the potential inside the axon from its negative resting value to about
+30 mV. The potential then rapidly decreases to about −90 mV and returns
more slowly back to the initial resting state. The whole pulse passes a given
point in a few milliseconds. The speed of pulse propagation depends on the
type of axon. Fast-acting axons propagate the pulse at speeds up to 100 m/sec.
The mechanism for the action potential is discussed in a following section.
Impulses produced by a given neuron are always of the same size and
propagate down the axon without attenuation. The nerve impulses are produced
at a rate proportional to the intensity of the stimulus. There is, however,
186 Chapter 13 Electricity
an upper limit to the frequency of impulses because a new impulse cannot
begin before the previous one is completed.
13.1.4 Axon as an Electric Cable
In the analysis of the electrical properties of the axon, we will use some of the
techniques of electrical engineering. This treatment is more complex than the
methods used in the other sections of the text. The added complexity, however,
is necessary for the quantitative understanding of the nervous system.
Although the axon is often compared to an electrical cable, there are profound
differences between the two. Still, it is possible to gain some insight into
the functioning of the axon by analyzing it as an insulated electric cable submerged
in a conducting fluid. In such an analysis, we must take into account
the resistance of the fluids both inside and outside the axon and the electrical
properties of the axon membrane. Because the membrane is a leaky insulator,
it is characterized by both capacitance and resistance. Thus, we need four
electrical parameters to specify the cable properties of the axon.
The capacitance and the resistance of the axon are distributed continuously
along the length of the cable. It is, therefore, not possible to represent
the whole axon (or any other cable) with only four circuit components. We
must consider the axon as a series of very small electrical-circuit sections
joined together. When a potential difference is set up between the inside and
the outside of the axon, four currents can be identified: the current outside
the axon, the current inside the axon, the current through the resistive component
of the membrane, and the current through the capacitive component of
the membrane (see Fig. 13.6). The electrical circuit representing a small axon
section of length x is shown in Fig. 13.6. In this small section, the resistances
of the outside and of the inside fluids are Ro and Ri, respectively. The
membrane capacitance and resistance are shown as Cm and Rm. The whole
axon is just a long series of these subunits joined together. This is shown in
Fig. 13.7. Sample values of the circuit parameters for both a myelinated and a
nonmyelinated axon of radius 5 � 10−6 m are listed in Table 13.1. (These values
were obtained from [13-4].) Note that the values in Table 13.1 are quoted
for a 1-m length of the axon. The unit mho for the conductivity of the axon
membrane is defined in Appendix B.
An examination of the axon performance shows immediately that the circuit
in Fig. 13.7 does not explain some of the most striking features of the
axon. An electrical signal along such a circuit propagates at nearly the speed
of light (3 � 108 m/sec), whereas a pulse along an axon travels at a speed
that is at most about 100 m/sec. Furthermore, as we will show, the circuit
in Fig. 13.7 dissipates an electrical signal very quickly; yet we know that
action potentials propagate along the axon without any attenuation. Therefore,
Section 13.1 The Nervous System 187
FIGURE 13.6 (a) Currents flowing through a small section of the axon.
(b) Electrical circuit representing a small section of the axon.
FIGURE 13.7 The axon represented as an electrical cable.
TABLE 13.1 Properties of Sample Axons
Property Nonmyelinated axon Myelinated axon
Axon radius 5 � 10−6
m 5 � 10−6
m
Resistance per unit length of fluid
both inside and outside axon (r)
6.37 � 109
/m 6.37 � 109
/m
Conductivity per unit length of axon
membrane (gm)
1.25 � 10−4
mho/m 3 � 10−7
mho/m
Capacitance per unit length of axon (c) 3 � 10−7
F/m 8 � 10−10
F/m
188 Chapter 13 Electricity
we must conclude that an electrical signal along the axon does not propagate
by a simple passive process.
13.1.5 Propagation of the Action Potential
After many years of research the propagation of an impulse along the axon is
now reasonably well understood. (See Fig. 13.8.) When the magnitude of the
FIGURE 13.8 The action potential. (a) The action potential begins with the axon
membrane becoming highly permeable to sodium ions (closed circles) which enter the
axon making it positive. (b) The sodium gates then close and potassium ions (open
circles) leave the axon making the interior negative again.
Section 13.1 The Nervous System 189
voltage across a portion of the membrane is reduced below a threshold value,
the permeability of the axon membrane to sodium ions increases rapidly. As
a result, sodium ions rush into the axon, cancel out the local negative charges,
and, in fact, drive the potential inside the axon to a positive value. This process
produces the initial sharp rise of the action potential pulse. The sharp positive
spike in one portion of the axon increases the permeability to the sodium
immediately ahead of it which in turn produces a spike in that region. In
this way the disturbance is sequentially propagated down the axon, much as a
flame is propagated down a fuse.
The axon, unlike a fuse, renews itself. At the peak of the action potential,
the axon membrane closes its gates to sodium and opens them wide to
potassium ions. The potassium ions now rush out, and, as a result, the axon
potential drops to a negative value slightly below the resting potential. After a
few milliseconds, the axon potential returns to its resting state and that portion
of the axon is ready to receive another pulse.
The number of ions that flow in and out of the axon during the pulse is
so small that the ion densities in the axon are not changed appreciably. The
cumulative effect of many pulses is balanced by metabolic pumps that keep
the ion concentrations at the appropriate levels. Using Eq. B.5 in Appendix B,
we can estimate the number of sodium ions that enter the axon during the
rising phase of the action potential. The initial inrush of sodium ions changes
the amount of electrical charge inside the axon. We can express this change
in charge Q in terms of the change in the voltage V across the membrane
capacitor C, that is,
Q C V (13.1)
In the resting state, the axon voltage is −70 mV. During the pulse, the voltage
changes to about +30 mV, resulting in a net voltage change across the
membrane of 100 mV. Therefore, V to be used in Eq. 13.1 is 100 mV.
The calculations outlined in Exercise 13-1 show that, in the case of the
nonmyelinated axon described in Table 13.1, during each pulse 1.87 � 1011
sodium ions enter per meter of axon length. The same number of potassium
ions leaves during the following part of the action potential. (Measurements
show that actually the ion flow is about three times higher than our simple
estimate.) The exercise also shows that, in the resting state, the number of
sodium ions inside a meter length of the axon is about 7 � 1014 and the number
of potassium ions is 7 � 1015. Thus, the inflow and the outflow of ions
during the action potential is negligibly small compared to the equilibrium
density.
AnothersimplecalculationusingEq.B.6yieldsanestimateoftheminimum
energy required to propagate the impulse along the axon. During the propagation
of one pulse, the whole axon capacitance is successively discharged and
190 Chapter 13 Electricity
then must be recharged again. The energy required to recharge a meter length
of the nonmyelinated axon is
E
1
2
C( V )2 1
2
� 3 � 10−7
� (0.1)2
1.5 � 10−9
J/m (13.2)
where C is the capacitance per meter of the axon. Because the duration of each
pulse is about 10−2 sec, and an axon can propagate at most 100 pulses/sec,
even at peak operation the axon requires only 1.5 � 10−7 W/m to recharge its
capacitance.
13.1.6 An Analysis of the Axon Circuit
The circuit in Fig. 13.7 does not contain the pulse-conducting mechanism of
the axon. It is possible to incorporate this mechanism into the circuit by connecting
small signal generators along the circuit. However, the analysis of
such a complex circuit is outside the scope of this text. Even the circuit in
Fig. 13.7 cannot be fully analyzed without calculus. We will simplify this
circuit by neglecting the capacitance of the axon membrane. The circuit is
then as shown in Fig. 13.9a. This representation is valid when the capacitors
are fully charged so that the capacitive current is zero. With this model, we
will be able to calculate the voltage attenuation along the cable when a steady
voltage is applied at one end. The simplified model, however, cannot make
predictions about the time-dependent behavior of the axon.
The problem then is to calculate the voltage V(x) at point x when a voltage
Va is applied at point x0 (see Fig. 13.9a). The approach is to calculate first
the voltage drop across a small incremental cable section of length x cut by
lines a and b (see [13-5, 13-6]). We assume that the cable is infinite in length
and that the total cable resistance to the right of line b is RT. Thus, the whole
cable to the right of line b is replaced by RT as shown in Fig. 13.9b. Because
the cable is infinite, the resistance to the right of any vertical cut equivalent to
line b is also RT. Specifically, the resistance to the right of line a is RT. We
can, therefore, calculate RT by equating the resistance to the right of line a in
Fig. 13.9b to RT, that is,
RT Ro + Ri +
RTRm
RT + Rm
(13.3)
Measurements show that the resistivities inside and outside the axon are about
the same. Therefore, Ri Ro R and Eq. 13.3 simplifies to
RT 2R +
RTRm
RT + Rm
(13.4)
Section 13.1 The Nervous System 191
FIGURE 13.9 (a) Approximation to the circuit in Fig. 13.7 with the capacitances
neglected. (b) The resistances to the right of line b replaced by the equivalent resistor RT.
The solution of Eq. 13.4 (see Exercise 13-2) yields
RT R + R2
+ 2R Rm
1/2
(13.5)
A simple circuit analysis (see Exercise 13-3) shows that
Vb
Va
1 + (2R)(RT + Rm)
RTRm
Va
1 + β
(13.6)
where β is the quantity in the square brackets.
We can calculate β from the measured parameters shown in Table 13.1.
The resistances R and Rm are the values for a small axon section of length
x. Therefore,
R r x and
1
Rm
gm x or Rm
1
gm x
192 Chapter 13 Electricity
From Eq. 13.5, it can be shown (see Exercise 13-4) that if x is very small
then
RT
2r
gm
1/2
(13.7)
and
β (2rgm)1/2
x
x
λ
(13.8)
where
λ
1
2rgm
1/2
(13.9)
Now returning to Eq. 13.6, since x is vanishingly small, β is also very
small. Therefore, the term 1/(1 + β) is approximately equal to 1 − β (see
Exercise 13-5). Consequently, the voltage Vb at b, a distance x away
from a, is
Vb Va 1 −
x
λ
(13.10)
To obtain the voltage at a distance x away from line a, we divide this distance
into increments of x such that n x x. We can then apply Eq. 13.10
successively down the cable and obtain the voltage at x (see Exercise 13-6) as
V(x) Va 1 −
x
λ
n
(13.11)
It can be shown that, for small x and large n, Eq. 13.11 can be written as
(see Exercise 13-7)
V(x) Vae−x/λ
(13.12)
Equation 13.12 states that if a steady voltage Va is applied across one point
in the axon membrane, the voltage decreases exponentially down the axon.
From Table 13.1, for a nonmyelinated axon λ is about 0.8 mm. Therefore, at
a distance 0.8 mm from the point of application, the voltage decreases to 37%
of its value at the point of application.
Myelinated axons, because of their outer sheath, have a much smaller
membrane conductance than axons without myelin. As a result, the value
of λ is larger. Using the values given in Table 13.1, we can show that λ is
16 mm for a myelinated axon. This result helps to explain the faster pulse
Section 13.1 The Nervous System 193
conduction along myelinated axons. As we mentioned earlier, the myelin
sheath is in 2-mm-long segments. The action potential is generated only at
the nodes between the segments. The pulse propagates through the myelinated
segments as a fast conventional electrical signal. Because λ is 16 mm, the pulse
decreases by only 13% as it traverses one segment, and it is still sufficiently
intense to generate an action potential at the next node.
13.1.7 Synaptic Transmission
So far we have considered the propagation of an electrical impulse down the
axon. Now we shall briefly describe how the pulse is transmitted from the
axon to other neurons or muscle cells.
At the far end, the axon branches into nerve endings which extend to the
cells that are to be activated. Through these nerve endings the axon transmits
signals, usually to a number of cells. In some cases the action potential is
transmitted from the nerve endings to the cells by electrical conduction. In
the vertebrate nervous system, however, the signal is usually transmitted by a
chemicalsubstance. Thenerveendingsareactuallynotincontactwiththecells.
There is a gap, about a nanometer wide (1 nm 10−9 m 10−7 cm) between
the nerve ending and the cell body. These regions of interaction between the
nerve ending and the target cell are called synapses (see Fig. 13.10). When
the impulse reaches the synapse, a chemical substance is released at the nerve
ending which quickly diffuses across the gap and stimulates the adjacent cell.
The chemical is released in bundles of discrete size.
Usually a neuron is in synaptic contact with many sources. Often a number
of synapses must be activated simultaneously to start the action potential in the
target cell. The action potential produced by a neuron is always of the same
FIGURE 13.10 Synapse.
194 Chapter 13 Electricity
magnitude. The neuron operates in an all-or-none mode: It either produces
an action potential of the standard size or does not fire at all. In some cases
the chemicals released at the synapse do not stimulate the cell but inhibit its
response to impulses arriving along a different channel. Presumably, these
types of interactions permit decisions to be made on a cellular level. The
details of these processes are not yet fully understood.
13.1.8 Action Potentials in Muscles
Muscle fibers produce and propagate electrical impulses in much the same
way as neurons. The action potential in the muscle fiber is initiated by the
impulses arriving from motor neurons. This stimulation causes a reduction
of the potential across the fiber membrane which initiates the ionic process
involved in the pulse propagation. The shape of the action potential is the
same as in the neuron except that its duration is usually longer. In skeletal
muscles, the action potential lasts about 20 msec, whereas in heart muscles it
may last a quarter of a second.
After the action potential passes through the muscle fiber, the fiber contracts.
In Chapter 5, we briefly discussed some aspects of muscle contraction.
The details of this process are not yet fully understood.
Within the skeletal muscle fibers, mechanoreceptor organs called muscle
spindles continuously transmit information on the state of muscle contraction.
This information is relayed via neurons for processing and further action. In
this way, the movement of muscles is continuously under control.
It is possible to stimulate muscle fibers by an external application of an
electric current. This effect was first observed in 1780 by Luigi Galvani who
noted that a frog’s leg twitched when an electric current passed through it.
(Galvani’s initial interpretation of this effect was wrong.) External muscle
stimulation is a useful clinical technique for maintaining muscle tone in cases
of temporary muscle paralysis resulting from nerve disorders.
13.1.9 Surface Potentials
The voltages and currents associated with the electrical activities in neurons,
muscle fibers, and other cells extend to regions outside the cells. As an example,
consider the propagation of the action potential along the axon. As the
voltage at one point on the axon drops suddenly, a voltage difference is produced
between this point and the adjacent regions. Consequently, current
flows both inside and outside the axon (see Fig. 13.11). As a result, a voltage
drop develops along the outer surface length of the axon.
Experiments are sometimes performed on a whole nerve consisting of
many axons. As shown in Fig. 13.12, two electrodes are placed along the nerve
Section 13.1 The Nervous System 195
FIGURE 13.11 The action potential produces currents both inside and outside the
axon.
FIGURE 13.12 Surface potential along a nerve bundle.
bundle, and the voltage between them is recorded. This measured voltage is
the sum of the surface potentials produced by the individual axons and yields
some information about the collective behavior of the axons.
Electric fields associated with the activities in cells extend all the way to
the surface of the animal body. Thus, along the surface of the skin, we can
measure electric potentials representing the collective cell activities associated
with certain processes in the body. Based on this effect, clinical techniques
have been developed to obtain, from the skin surface, information about
the activities of the heart (electrocardiography) and the brain (electroencephalography).
The measurement of these surface signals will be discussed
in Chapter 14.
Surface signals are associated with many other activities, such as movement
of the eye, contractions of the gastrointestinal tract, and movement of
muscles. Using a technique called electromyography (EMG), measurement
of action potentials and their speed of propagation along muscles can provide
information about muscle and nerve disorders. (see [13-5]). Surface
196 Chapter 13 Electricity
potentials associated with metabolic activities have also been observed in
plants and bones, as discussed in the following sections.
13.2 Electricity in Plants
The type of propagating electrical impulses we have discussed in connection
with neurons and muscle fibers have also been found in certain plant cells. The
shape of the action potential is the same in both cases, but the duration of the
action potential in plant cells is a thousand times longer, lasting about 10 sec.
The speed of propagation of these plant action potentials is also rather slow,
only a few centimeters per second. In plant cells, as in neurons, the action
potential is elicited by various types of electrical, chemical, or mechanical
stimulation. However, the initial rise in the plant cell potential is produced by
an inflow of calcium ions rather than sodium ions.
The role of action potentials in plants is not yet known. It is possible
that they coordinate the growth and the metabolic processes of the plant and
perhaps control the long-term movements exhibited by some plants.
13.3 Electricity in the Bone
When certain types of crystals are mechanically deformed, the charges in them
are displaced; as a result, they develop voltages along the surface. This phenomenon
is called the piezoelectric effect (Fig. 13.13). Bone is composed of a
crystalline material that exhibits the piezoelectric effect. It has been suggested
FIGURE 13.13 The piezoelectric effect.
Section 13.4 Electric Fish 197
FIGURE 13.14 (a) Charges are produced along the bone as a result of deformation.
(b) Guided by the charge distribution, substance is added and removed from the bone.
(c) Reconstructed bone.
that these piezoelectric voltages play a role in the formation and nourishment
of the bone.
The body has mechanisms for both building and destroying bone. New
bone is formed by cells called osteoblasts and is dissolved by cells called
osteoclasts. It has been known for some time that a living bone will adapt its
structure to a long-term mechanical load. For example, if a compressive force
bends a bone, after a time the bone will assume a new shape. Some portions
of the bone gain substance and others lose it in such a way as to strengthen the
bone in its new position. It has been suggested that the appropriate addition
and removal of bone tissue is guided by the piezoelectric potentials produced
by the deformation (see Fig. 13.14).
Piezoelectricity in the bone may have yet another function. All tissue
including bone has to be nourished by fluids. The nourishing fluids move into
the bone through very narrow canals. Without a pumping mechanism, the
flow of this fluid would be too slow to provide the bone with the necessary
nutrients. It has been suggested that the piezoelectric voltages produced by
forces due to normal body movement act on the ions in the nutrient fluid and
pump it in and out of the bone.
13.4 Electric Fish
Most animals do not possess sensory organs that are specifically designed to
detect external electric fields, but sharks and rays are exceptions. These fish
198 Chapter 13 Electricity
have small organs along their skin which are remarkably sensitive to electric
fields in water. A shark responds to an electric field as small as 1 μV/m, which
is in the range of fields found along the skin of animals. (A flashlight battery
with terminals separated by 1500 km produces a field of this strength.) The
shark uses these electrical organs to locate animals buried in sand and perhaps
even to communicate with other sharks. Sharks are also known to bite boat
propellers, probably in response to the electric field generated in the proximity
of the metal.
An equally remarkable use of electricity is found in the electric eel, which
can generate along its skin electric pulses up to 500 V with currents reaching
80 mA. The eel uses this ability as a weapon. When it comes in contact with
its prey, the high-voltage pulse passes through the victim and stuns it.
The electric organ of the eel consists of specialized muscle fibers that are
connected together electrically. The high voltage is produced by a series interconnection
of many cells, and the large current is obtained by connecting the
series chains in parallel (see Exercise 13-8). A number of other fish possess
similar electric organs.
EXERCISES
13-1. (a) Using Eq. 13.1 and the data in Table 13.1, calculate the number of
ions entering the axon during the action potential, per meter of nonmyelinated
axon length. (The charge on the ion is 1.6 � 10−19 coulomb.)
(b) During the resting state of the axon, typical concentrations of sodium
andpotassiumionsinsidetheaxonare15and150millimole/liter, respectively.
From the data in Table 13.1, calculate the number of ions per
meter length of the axon.
1 mole/liter 6.02 � 1020 particles (ions, atoms, etc.)
cm3
13-2. From Eq. 13.4, obtain a solution for RT. (Remember that RT must be
positive.)
13-3. Verify Eq. 13.6.
13-4. Show that when x is very small, RT is given by Eq. 13.7.
13-5. Show that if β is small, 1/(1 + β) ≈ 1 − β. (Refer to tables of series
expansion.)
13-6. Verify Eq. 13.11.
Chapter 13 Exercises 199
13-7. Using the binomial theorem, show that Eq. 13.11 can be written as
V(x) Va 1 −
n x
λ
+
n(n − 1)
2!
x
λ
2
−
n(n − 1)(n − 2)
3!
x
λ
3
+ � � �
Since x is vanishingly small, n must be very large. Show that the
above equation approaches the expansion for an exponential function.
(Refer to tables of series expansion.)
13-8. (a) From the data provided in the text, estimate the number of cells
that must be connected in series to provide the 500 V observed at the
skin of the electric eel. (b) Estimate the number of chains that must be
connected in parallel to provide the observed currents.
Assume that the size of the cell is 10−5 m, the pulse produced by a
single cell is 0.1 V, and the duration of pulse is 10−2 sec. Use the data
in the text and in Exercise 13-1 to estimate the current flowing into a
single cell during the action potential.
Chapter 14
Electrical Technology
Electrical technology was developed by applying some of the basic principles
of physics to problems in communications and industry. Although this technology
was directed primarily toward industrial and military applications, it
has made great contributions to the life sciences. Electrical technology has
provided tools for the observation of biological phenomena that would have
been otherwise inaccessible. It yielded most of the modern clinical and diagnostic
equipment used in medicine. Even the techniques developed for the
analysis of electrical devices have been useful in the study of living systems.
In this chapter, we will describe some of the applications of electrical
technology in these areas.
14.1 Electrical Technology in Biological Research
Our understanding of the world would be greatly limited if it were based only
on observations made by our unaided senses. Well developed though our
senses are, their responses are limited. We cannot hear sound at frequencies
above 20,000 Hz. We cannot see electromagnetic radiation outside the limited
wavelength region between about 400 nm and 700 nm (1 nm 10−9 m).
Even in this visible range, we cannot detect variations in light intensity that
occur at a rate faster than about 20 Hz. Although many of the vital processes
within our bodies are electrical, our senses cannot detect small electric fields
directly. Electrical technology has provided the means for translating information
from many areas into the domain of our senses.
200
Section 14.1 Electrical Technology in Biological Research 201
FIGURE 14.1 An experiment in biology.
Electrical technology is a vast subject that we cannot possibly cover in
this short chapter. Here we will simply outline the general techniques used in
observing life processes. A description of the various electrical components
is found in [14-3] and [14-4].
A diagram of a typical experimental setup in biology is shown in Fig. 14.1.
The various subunits of the experiment are shown as blocks of specialized
functions. We start with the phenomenon we want to observe but which we
cannot detect with our senses. This may be, for example, a high-frequency
sound emitted by a bat, the electrochemical activity of a cell, the subtle movement
of a muscle, or the light emitted by a fluorescent dye. These phenomena
are first translated into electrical signals which then carry information about
the intensity and time variations of the original event. Specialized devices are
required to perform this task. Some of these devices are commonly found in
our everyday technology; others are rather esoteric. Sound, for example, is
translated into electrical signals by microphones. Light can be translated into
an electrical current by photomultipliers.
The electrical signals generated in this way are usually too weak to drive
the final instrument that displays the signals for our observation, so the power
and amplitude of the signal are increased by a device called an amplifier. The
amplified signal then drives the display unit.
The display unit must be matched to the type of signal that is being
observed. A slowly varying signal can be displayed on a voltmeter, which
has a pointer that moves in accord with the current. Somewhat faster signals
are often displayed on a pen recorder, which draws the shape of the signal on
a chart. Very fast signals are recorded by a device called an oscilloscope. This
instrument is similar to a television set. A beam of electrons generated inside
the device hits a fluorescent screen that emits lights at the point of impact.
202 Chapter 14 Electrical Technology
The motion and intensity of the beam are controlled by electrical signals
applied to the oscilloscope. The resulting picture on the screen displays the
information content of the signal. Speakers can also translate into sound electrical
signals that have a time variation in the audible range of frequencies.
Often the experimental signal is very noisy. In addition to the desired
information, it contains spurious signals due to various sources extraneous to
the main phenomenon. Techniques have been developed for analyzing such
signals and extracting the relevant information from the noise. In modern
experiments, this processing is often performed by a computer.
14.2 Diagnostic Equipment
Most of the diagnostic equipment in medicine utilizes electrical technology
in one form or another. Even the traditional stethoscope is now available
with electronic modifications that increase its sensitivity. We will describe
here only two of the many diagnostic instruments found in a modern clinical
facility: the electrocardiograph and the electroencephalograph.
As a result of the ionic currents associated with electrical activities in the
cells, potential differences are produced along the surface of the body. (See
Chapter 13.) By measuring these potential differences between appropriate
points on the surface of the body, it is possible to obtain information about the
functioning of specific organs. The surface potentials are usually very small
and, therefore, must be amplified before they can be displayed for examination.
14.2.1 The Electrocardiograph
The electrocardiograph (ECG) is an instrument that records surface potentials
associated with the electrical activity of the heart. The surface potentials are
conducted to the instrument by metal contacts called electrodes which are
fixed to various parts of the body. Usually the electrodes are attached to the
four limbs and over the heart. Voltages are measured between two electrodes
at a time. (See Fig. 14.2.)
A typical normal signal recorded between two electrodes is shown in
Fig. 14.3. The main features of this wave form are identified by the letters
P, Q, R, S, and T. The shape of these features varies with the location of
the electrodes. A trained observer can diagnose abnormalities by recognizing
deviations from normal patterns.
The wave shape in Fig. 14.3 is explained in terms of the pumping action
of the heart described in Chapter 8. The rhythmic contraction of the heart
is initiated by the pacemaker, which is a specialized group of muscle cells
located near the top of the right atrium. Immediately after the pacemaker fires,
Section 14.2 Diagnostic Equipment 203
FIGURE 14.2 Electrocardiography.
FIGURE 14.3 An electrocardiogram.
the action potential propagates through the two atria. The P wave is associated
with the electrical activity that results in the contraction of the atria. The QRS
wave is produced by the action potential associated with the contraction of the
ventricles. The T wave is caused by currents that bring about the recovery of
the ventricle for the next cycle.
14.2.2 The Electroencephalograph
The electroencephalograph (EEG) measures potentials along the surface of
the scalp. Here again electrodes are attached to the skin at various positions
204 Chapter 14 Electrical Technology
FIGURE 14.4 EEG potentials between two pairs of electrodes: (a) subject alert,
(b) subject drowsy, (c) light sleep, (d) deep sleep.
along the scalp. The instrument records the voltages between pairs of electrodes.
The EEG signals are much more complex and difficult to interpret
than those produced by the electrocardiograph. The EEG signals are certainly
the result of collective neural activity in the brain. However, so far it has not
been possible to relate unambiguously the EEG potentials to specific brain
functions. Nevertheless, certain types of patterns are known to be related to
specific activities, as illustrated in Fig. 14.4.
Electroencephalographs have been useful in diagnosing various brain disorders.
Epileptic seizures, for example, are characterized by pronounced EEG
abnormalities (see Fig. 14.5). Brain tumors can often be located by a careful
examination of EEG potentials along the whole contour of the scalp.
14.3 Physiological Effects of Electricity
The painful shock produced by electricity is well known to most people.
The shock results from a current passing through the body. An electrical
current has two effects on body tissue. The current stimulates nerves and
muscle fibers, which produces pain and a contraction of muscles, and it also
heats the tissue through dissipation of electrical energy. Both of these effects,
Section 14.3 Physiological Effects of Electricity 205
FIGURE 14.5 Abnormal EEG patterns. Pattern b is typical of petit mal seizures.
if sufficiently intense, can cause severe injury or death. But if the electrical
current is applied in a controlled way, both the heating and the muscle stimulation
can be beneficial. For example, local heating of tissue by high-frequency
electric currents promotes healing in much the same way as by ultrasonic
diathermy.
The amount of current flowing through the body is governed by Ohm’s
law. Thus, it depends on the voltage of the source and the electrical resistance
of the body. The body tissue is a relatively good conductor. Since most of the
electrical resistance is in the skin, the danger of electrical shock increases if
the skin is wet at the point of contact.
Most people begin to feel an electrical current when it reaches a magnitude
of about 500 μA. A 5-mA current causes pain, and currents larger than about
10 mA produce sustained tetanizing contraction of some muscles. This is a
dangerous situation because under these conditions the person cannot release
the conductor that is delivering the current into his or her body.
The brain, the respiratory muscles, and the heart are all very seriously
affected by large electric currents. Currents in the range of a few hundred milliamperes
flowing across the head produce convulsions resembling epilepsy.
Currents in this range are used in electric shock therapy to treat certain mental
disorders.
Currents in the range of a few amperes flowing in the region of the heart
can cause death within a few minutes. In this connection, a large current of
about 10 A is often less dangerous than a 1-A current. When the smaller current
passes through the heart, it may tetanize only part of the heart, thereby
causing a desynchronization of the heart action; this condition is called fibrillation.
The movements of the heart become erratic and ineffective in pumping
blood. Usually fibrillation does not stop when the current source is removed.
A large current tetanizes the whole heart, and when the current is discontinued
the heart may resume its normal rhythmic activity.
206 Chapter 14 Electrical Technology
Fibrillations often occur during a heart attack and during cardiac surgery.
The tetanizing effect of large currents can be used to synchronize the heart.
A clinical device designed for this purpose is called a defibrillator. A capacitor
in this device is charged to about 6000 V and stores about 200 J of energy. Two
electrodes connected to the capacitor through a switch are placed on the chest.
When the switch is closed, the capacitor rapidly discharges through the body.
The current pulse lasts about 5 msec, during which the heart is tetanized (see
Exercise 14-1). After the pulse, the heart may resume its normal beat. Often
the heart must be shocked a few times before it resynchronizes.
Electric current can also be used to stimulate muscles more gently. We
have already mentioned the electric stimulation of paralyzed skeletal muscles
to maintain their tone. Heart muscles can be triggered in a similar way. In
some heart diseases, the pacemaker cells that control the timing of the heartbeat
cease to function properly, and electronic pacemakers have been very
useful. The electronic pacemaker is basically a pulse generator that produces
short periodic pulses that initiate and control the frequency of the heartbeat.
The device can be made small enough for surgical implantation. Unfortunately,
the battery that powers the pacemaker has a finite lifetime and must be
replaced every few years.
14.4 Control Systems
Many of the processes in living systems must be controlled to meet the requirements
of the organism. We have already encountered a few examples of
controlled processes in our earlier discussions. Temperature control in the
body and the growth of bones were two cases where various processes had to
be regulated in order to achieve the desired condition. In this section, we will
describe briefly a useful general method of analyzing such control systems.
Features common to all control systems are shown in Fig. 14.6. Each block
represents an identifiable function within the control system. The control process
consists of:
1. The parameter to be controlled. This may be the temperature of the skin,
the movement of muscles, the rate of heart beat, the size of the bone,
and so on.
2. A means of monitoring the parameter and transmitting information
about its state to some decision-making center. This task is usually performed
by the sensory neurons.
3. Some reference value to which the controlled parameters are required to
comply. The reference value may be in the central nervous system in the
Section 14.4 Control Systems 207
FIGURE 14.6 Control of a biological process.
form of a decision, for example, about the position of the hand. In this
case, the reference value is changeable and is set by the central nervous
system. Many references for body functions are autonomous, however,
not under the cognitive control of the brain.
4. A method for comparing the state of the parameter with the reference
value and for transmitting instructions to bring the two into accord. The
instructions may be transmitted by nerve impulses or in some cases by
chemical messengers called hormones, which diffuse through the body
and control various metabolic functions.
5. A mechanism for translating the messages into actions that alter the state
of the controlled parameter. In the case of the hand position, for example,
this is the contraction of a set of muscle fibers.
We will now illustrate these concepts with a concrete example of the control
of the light intensity reaching the retina of the eye (see Fig. 14.7). Light
enters the eye through the pupil, which is the dark opening in the center of
the iris. (The iris is the colored disk in the eyeball.) The size of the opening
decreases as the light intensity increases. Thus, the iris acts somewhat like the
208 Chapter 14 Electrical Technology
FIGURE 14.7 Control of the light intensity reaching the retina.
automatic diaphragm in a camera. Clearly this action must be governed by a
control system.
Light reaching the retina is converted to neural impulses, which are generated
at a frequency proportional to the light intensity. At some place along
the nervous system of vision, this information is interpreted and compared to
a preset reference value stored probably in the brain. The reference itself can
be altered by hormones and various emotional stimuli. The result of this comparison
is transmitted by means of nerve impulses to the muscles of the iris
which then adjust the size of the opening in response to this signal.
14.5 Feedback
For many years engineers have studied mechanical and electrical systems that
have the general characteristics of control systems in biological organisms.
Voltage regulators, speed controls, and thermostatic heat regulators all have
features in common with biological control systems. Engineers have developed
techniques for analyzing and predicting the behavior of control systems.
These techniques have also been useful in the study of biological systems.
An engineering analysis of such systems is usually done in terms of input
and output. In the light-intensity control example, the input is the light reaching
the retina, and the output is the response of the retina to light. The system
itself is that which yields an output in response to the input. In our case, this is
the retina and the associated nerve circuits. The aim of the iris control system
is to maintain the output as constant as possible.
Section 14.5 Feedback 209
FIGURE 14.8 An amplifier without feedback.
The most significant point to note about control systems such as the one
shown in Fig. 14.7 is that the output affects the input itself. Such systems
are called feedback systems (because information about the output is fed back
to the input). The system is said to have negative feedback if it opposes a
change in the input and positive feedback if it augments a change in the input.
The light control in Fig. 14.7 is a negative feedback because an increase in
the light intensity causes a decrease in the iris opening and a corresponding
reduction of the light reaching the retina. Regulation of body temperature
by sweating or shivering is another example of negative feedback, whereas
sexual arousal is an example of positive feedback. In general, negative feedback
keeps the system response at a relatively constant level. Therefore, most
biological feedback systems are in fact negative.
We will illustrate the method of system analysis with an example from
electrical engineering. We will analyze in these terms a voltage amplifier that
has part of its output fed back to the input. Let us first consider a simple
amplifier without feedback (see Fig. 14.8). The amplifier is an electric device
that increases the input voltage (Vin) by a factor A; that is, the output voltage
Vout is
Vout AVin (14.1)
It is evident from this equation that the amplification A is simply determined
by the ratio of the output and input voltages; that is,
A
Vout
Vin
(14.2)
Now let us introduce feedback (Fig. 14.9). Part of the output (β � Vout) is
added back to the amplifier input so that the voltage at the input terminal of
the amplifier (Vin ) is
Vin Vin + β � Vout (14.3)
Here Vin is the externally applied voltage. The amplification of the total feedback
system is
Afeedback
Vout
Vin
(14.4)
210 Chapter 14 Electrical Technology
FIGURE 14.9 An amplifier with feedback.
Using the fact that Vout AVin , we can show that (see Exercise 14-2)
Afeedback
A
1 − Aβ
(14.5)
Now if β is a negative number, the amplification with feedback is smaller
than the amplification without feedback (i.e., Afeedback is smaller than A).
A negative β implies that the voltage is added out of phase with the external
input voltage. This is negative feedback. With positive β, we have positive
feedback and increased amplification.
This type of analysis has the advantage that we can learn about the system
without a detailed knowledge of the individual system components. We
can vary the frequency, the magnitude, and the duration of the input voltage
and measure the corresponding output voltage. From these measurements,
we can obtain some information about the amplifier and the feedback component
without knowing anything about the transistors, resistors, capacitors, and
other components that make up the device. We could, of course, obtain this
information and much more by a detailed analysis of the device in terms of its
basic components, but this would involve much more work.
In the study of complex biological functions, the systems approach is
often very useful because the details of the various component processes are
unknown. For example, in the iris control system, very little is known about
the processing of the visual signals, the mechanism of comparing these signals
to the reference, or the nature of the reference itself. Yet by shining light at
various intensities, wavelengths, and durations into the eye and by measuring
the corresponding changes in the iris opening, we can obtain significant information
about the system as a whole and even about the various subunits. Here
the techniques developed by the engineers are useful in analyzing the system
(see Exercises 14-3 and 14-4). However, many biological systems are so
complicated with many inputs, outputs, and feedbacks that even the simplified
systems approach cannot yield a tractable formulation.
Section 14.6 Sensory Aids 211
14.6 Sensory Aids
Sight and hearing are the two principal pathways through which our brain
receives information about the external world. The two organs, eyes and ears,
that transmit light and sound information into the brain are often damaged
and their function needs to be supplemented. Eyeglasses came into use in
the 1200s. At first these visual aids provided only simple magnified images
of objects. Gradually a sophisticated technology evolved that produces eyeglasses
to compensate for a wide range of visual problems (see Chapter 15).
Ear horns, in one form or another, have been used to aid hearing for thousands
of years. These devices improve hearing by collecting sound from an
area significantly larger than the pinna (see Chapter 12).
Electrical technology has led to the development of devices that greatly
enhance hearing and in some cases even restore hearing. The restoration of
vision is far more challenging and while several avenues of research are being
pursued the final goal seems far in the future.
14.6.1 Hearing Aids
The basic principle of hearing aids is simple. A microphone converts sound
to an electrical signal. The electrical signal is amplified and converted back
into sound using a speaker-type device. The net result is an amplification of
the sound that enters the ear.
The first hearing aids became commercially available in the 1930s. They
were relatively large cumbersome devices using battery-powered vacuum tube
amplifiers. The batteries had to be replaced daily.
The much smaller transistor amplifiers that became available in the 1950s
made hearing aids truly practicable. Transistorized hearing aids were now
small enough to be placed in the ear. The application of digital computer
technology to hearing aids was another major improvement that allowed individual
tailoring of the device to compensate for the specific hearing deficits
of the user. Using various feedback networks, modern hearing aids automatically
adjust the volume of the sound so that quiet sounds can be heard and
loud sounds not be painfully overwhelming.
14.6.2 Cochlear Implant
A cochlear implant functions differently from a hearing aid. A hearing aid
simply amplifies incoming sound compensating for the diminished functioning
of the ear. A cochlear implant converts sound to electrical signals of
the type produced by the inner ear in response to sound that enters the ear.
212 Chapter 14 Electrical Technology
FIGURE 14.10 Cochlear implant. 1. Sounds are picked up by the microphone.
2. The signal is then “coded” (turned into a special pattern of electrical pulses). 3. These
pulses are sent to the coil and are then transmitted across the skin to the implant. 4. The
implant sends a pattern of electrical pulses to the electrodes in the cochlea. 5. The
auditory nerve picks up these electrical pulses and sends them to the brain. The brain
recognizes these signals as sound.
The electrical signal is wirelessly transmitted to electrodes surgically implanted
in the inner ear. The signals applied to the electrodes stimulate the auditory
nerve to produce the sensation of sound. Thus the cochlear implant actually
mimics the functions of the ear and can restore partial hearing to the deaf.
A sketch of a cochlear implant system is shown in Fig. 14.10. The external
part of the system is small enough to be placed behind the ear. It consists
of a microphone, a signal processor, and a transmitter. The internal part consists
of the receiver and an array of electrodes implanted and wound through
the cochlea.
The microphone converts the sound to an electrical signal. Such electrical
signals as are produced by the microphone could themselves stimulate the
5
4
3
2
1
Chapter 14 Exercises 213
auditory nerve, but the neural signals produced by such stimulation would
not be interpreted by the brain as sound. In the normal ear the fluid filled
cochlea processes the sound signal according to frequency such that the various
frequency components of the incoming sound stimulate nerve endings
along different parts of the basilar membrane (see Chapter 12). This type of a
frequency-selective stimulation of the neural network provided by the cochlea
is essential if the signal is to be interpreted by the brain as sound.
One of the main challenges in the design of cochlear implants was the
development of signal-processing techniques that duplicated the action of a
normal cochlea. Much of the work in this area was done in the 1950s and 60s.
First experiments with human implants began in the mid-1960s and continued
through the 1970s. In 1984, FDA approved implantation into adults and
shortly after, into children.
Usually a person receiving an implant is not immediately able to hear
sounds properly. A period of training and speech therapy are needed before
the full benefits of the device are realized.
EXERCISES
14-1. From the data in the text, compute the capacitance of the capacitor in the
defibrillator and calculate the magnitude of the average current flowing
during the pulse.
14-2. Verify Eq. 14.5.
14-3. Draw a block diagram for the following control systems. (a) Control of
the body temperature in a person. (b) Control of the hand in drawing a
line. (c) Control of the reflex action when the hand draws away from
a painful stimulus. Include here the type of control that the brain may
exercise on this movement. (d) Control of bone growth in response to
pressure.
14-4. For each of the control systems in Exercise 14-3, discuss how the system
could be studied experimentally.
14-5. Discuss the controversy surrounding cochlear implants.
Chapter 15
Optics
Light is the electromagnetic radiation in the wavelength region between about
400 and 700 nm (1 nm 10−9 m). Although light is only a tiny part of the
electromagnetic spectrum, it has been the subject of much research in both
physics and biology. The importance of light is due to its fundamental role
in living systems. Most of the electromagnetic radiation from the sun that
reaches the Earth’s surface is in this region of the spectrum, and life has
evolved to utilize it. In photosynthesis, plants use light to convert carbon
dioxide and water into organic materials, which are the building blocks of
living organisms. Animals have evolved light-sensitive organs which are their
main source of information about the surroundings. Some bacteria and insects
can even produce light through chemical reactions.
Optics, which is the study of light, is one of the oldest branches of physics.
It includes topics such as microscopes, telescopes, vision, color, pigments,
illumination, spectroscopy, and lasers, all of which have applications in the
life sciences. In this chapter, we will discuss four of these topics: vision,
telescopes, microscopes, and optical fibers. Background information needed
to understand this chapter is reviewed in Appendix C.
15.1 Vision
Vision is our most important source of information about the external world.
It has been estimated that about 70% of a person’s sensory input is obtained
through the eye. The three components of vision are the stimulus, which
214
Section 15.3 Structure of the Eye 215
is light; the optical components of the eye, which image the light; and the
nervous system, which processes and interprets the visual images.
15.2 Nature of Light
Experiments performed during the nineteenth century showed conclusively
that light exhibits all the properties of wave motion, which were discussed
in Chapter 12. At the beginning of this century, however, it was shown that
wave concepts alone do not explain completely the properties of light. In
some cases, light and other electromagnetic radiation behave as if composed
of small packets (quanta) of energy. These packets of energy are called photons.
For a given frequency f of the radiation, each photon has a fixed amount
of energy E which is
E hf (15.1)
where h is Planck’s constant, equal to 6.63 � 10−27 erg-sec.
In our discussion of vision, we must be aware of both of these properties
of light. The wave properties explain all phenomena associated with the propagation
of light through bulk matter, and the quantum nature of light must be
invoked to understand the effect of light on the photoreceptors in the retina.
15.3 Structure of the Eye
A diagram of the human eye is given in Fig. 15.1. The eye is roughly a sphere,
approximately 2.4 cm in diameter. All vertebrate eyes are similar in structure
but vary in size. Light enters the eye through the cornea, which is a transparent
section in the outer cover of the eyeball. The light is focused by the lens
system of the eye into an inverted image at the photosensitive retina, which
covers the back surface of the eye. Here the light produces nerve impulses
that convey information to the brain.
The focusing of the light into an image at the retina is produced by the curved
surface of the cornea and by the crystalline lens inside the eye. The focusing
power of the cornea is fixed. The focus of the crystalline lens, however, is
alterable, allowing the eye to view objects over a wide range of distances.
In front of the lens is the iris, which controls the size of the pupil, or
entrance aperture into the eye (see Chapter 14). Depending on the intensity
of the light, the diameter of the aperture ranges from 2 to 8 mm. The cavity of
the eye is filled with two types of fluid, both of which have a refractive index
about the same as water. The front of the eye, between the lens and the cornea,
216 Chapter 15 Optics
FIGURE 15.1 The human eye.
is filled with a watery fluid called the aqueous humor. The space between the
lens and the retina is filled with the gelatinous vitreous humor.
15.4 Accommodation
The focusing of the eye is controlled by the ciliary muscle, which can change
the thickness and curvature of the lens. This process of focusing is called
accommodation. When the ciliary muscle is relaxed, the crystalline lens is
fairly flat, and the focusing power of the eye is at its minimum. Under these
conditions, a parallel beam of light is focused at the retina. Because light
from distant objects is nearly parallel, the relaxed eye is focused to view
distant objects. In this connection, “distant” is about 6 m and beyond (see
Exercise 15-1).
The viewing of closer objects requires greater focusing power. The light
from nearby objects is divergent as it enters the eye; therefore, it must be
focused more strongly to form an image at the retina. There is, however, a
limit to the focusing power of the crystalline lens. With the maximum contraction
of the ciliary muscle, a normal eye of a young adult can focus on
objects about 15 cm from the eye. Closer objects appear blurred. The minimum
distance of sharp focus is called the near point of the eye.
The focusing range of the crystalline lens decreases with age. The near
point for a 10-year-old child is about 7 cm, but by the age of 40 the near point
Section 15.5 Eye and the Camera 217
shifts to about 22 cm. After that the deterioration is rapid. At age 60, the near
point is shifted to about 100 cm. This decrease in the accommodation of the
eye with age is called presbyopia.
15.5 Eye and the Camera
Although the designers of the photographic camera did not consciously imitate
the structure of the eye, many of the features in the two are remarkably similar
(see Fig. 15.2). Both consist of a lens system that focuses a real inverted image
onto a photosensitive surface. In the eye, as in the camera, the diameter of the
light entrance is controlled by a diaphragm that is adjusted in accord with the
available light intensity. In a camera, the image is focused by moving the lens
with respect to the film. In the eye, the distance between the retina and the
lens is fixed; the image is focused by changing the thickness of the lens.
Even the photosensitive surfaces are somewhat similar. Both photographic
film and the retina consist of discrete light-sensitive units, microscopic in size,
which undergo chemical changes when they are illuminated. In fact, under
special circumstances, the retina can be “developed,” like film, to show the
image that was projected on it. This was first demonstrated in the 1870s by
the German physiologist W. Kuhne. He exposed the eye of a living rabbit to
light coming through a barred window. After 3 minutes of exposure to light,
the rabbit was killed and its retina was immersed in an alum solution which
FIGURE 15.2 The eye and the camera.
218 Chapter 15 Optics
fixed the retinal reaction. The barred window was clearly visible on the retina.
A few years later, Kuhne fixed the retina from the head of a guillotined criminal.
He observed an image, but he could not interpret it in terms of anything
that the man had seen before he was beheaded.
The analogy between the eye and the camera, however, is not complete.
As we will describe later, the eye goes far beyond the camera in processing
the images that are projected on the retina.
15.5.1 Aperture and Depth of Field
The iris is the optical aperture of the eye, and its size varies in accordance with
the available light. If there is adequate light, the quality of the image is best
with the smallest possible aperture. This is true for both the eye and the camera.
There are two main reasons for the improved image with reduced aperture.
Imperfections in lenses tend to be most pronounced around the edges. A small
aperture restricts the light path to the center of the lens and eliminates the
distortions and aberrations produced by the periphery.
A smaller aperture also improves the image quality of objects that are not
located at the point on which the eye or the camera is focused. An image
is in sharp focus at the retina (or film) only for objects at a specific distance
from the lens system. Images of objects not at this specific plane are
blurred at the retina (see Fig. 15.3); in other words, a point that is not in exact
FIGURE 15.3 Size of image disk: (a) with large aperture, (b) with small aperture.
Section 15.6 Lens System of the Eye 219
focus appears as a disk on the retina. The amount of blurring depends on the
size of the aperture. As shown in Fig. 15.3, a small aperture reduces the diameter
of the blurred spot and allows the formation of a relatively clear image
from objects that are not on the plane to which the eye is focused. The range
of object distances over which a good image is formed for a given setting of
the focus is called the depth of field. Clearly a small aperture increases the
depth of field. It can be shown that the depth of field is inversely proportional
to the diameter of the aperture (see Exercise 15-2).
15.6 Lens System of the Eye
The focusing of the light into a real inverted image at the retina is produced by
refraction at the cornea and at the crystalline lens (see Fig. 15.4). The focusing
or refractive power of the cornea and the lens can be calculated using Eq. C.9,
(Appendix C). The data required for the calculation are shown in Table 15.1.
TABLE 15.1 Parameters for the Eye
Radius (mm)
Index of
Front Back refraction
Cornea 7.8 7.3 1.38
Lens, min. power 10.00 −6.0
Lens, max. power 6.0 −5.5 1.40
Aqueous and vitreous humor 1.33
FIGURE 15.4 Focusing by the cornea and the crystalline lens (not to scale).
220 Chapter 15 Optics
The largest part of the focusing, about two thirds, occurs at the cornea.
The power of the crystalline lens is small because its index of refraction is
only slightly greater than that of the surrounding fluid. In Exercise 15-3, it is
shown that the refractive power of the cornea is 42 diopters, and the refractive
power of the crystalline lens is variable between about 19 and 24 diopters.
(For a definition of the unit diopter, see Appendix C.)
The refractive power of the cornea is greatly reduced when it is in contact
with water (see Exercise 15-4). Because the crystalline lens in the human
eye cannot compensate for the diminished power of the cornea, the human
eye under water is not able to form a clear image at the retina and vision is
blurred. In fish eyes, which have evolved for seeing under water, the lens is
intended to do most of the focusing. The lens is nearly spherical and has a
much greater focusing power than the lens in the eyes of terrestrial animals
(see Exercise 15-5).
15.7 Reduced Eye
To trace accurately the path of a light ray through the eye, we must calculate
the refraction at four surfaces (two at the cornea and two at the lens). It is
possible to simplify this laborious procedure with a model called the reduced
eye, shown in Fig. 15.5. Here all the refraction is assumed to occur at the front
surface of the cornea, which is constructed to have a diameter of 5 mm. The
eye is assumed to be homogeneous, with an index of refraction of 1.333 (the
same as water). The retina is located 2 cm behind the cornea. The nodal point
n is the center of corneal curvature located 5 mm behind the cornea.
This model represents most closely the relaxed eye which focuses parallel
light at the retina, as can be confirmed using Eq. C.9. For the reduced eye, the
FIGURE 15.5 The reduced eye.
Section 15.7 Reduced Eye 221
second term on the right-hand side of the equation vanishes because the light
is focused within the reduced eye so that nL n2. Equation C.9, therefore,
simplifies to
n1
p
+
nL
q
nL − n1
R
(15.2)
where n1 1, nL 1.333, and R 0.5 cm. Because the incoming light is
parallel, its source is considered to be at infinity (i.e., p ∞). Therefore, the
distance q at which parallel light is focused is given by
1.333
q
1.333 − 1
5
or
q
1.333 � 5
0.333
20 mm
The anterior focal point F for the reduced eye is located 15 mm in front of
the cornea. This is the point at which parallel light originating inside the eye
is focused when it emerges from the eye (see Exercise 15-6).
Although the reduced eye does not contain explicitly the mechanism of
accommodation, we can use the model to determine the size of the image
formed on the retina. The construction of such an image is shown in Fig. 15.6.
Rays from the limiting points of the object A and B are projected through the
nodal point to the retina. The limiting points of the image at the retina are a
and b. This construction assumes that all the rays from points A and B that
enter the eye are focused on the retina at points a and b, respectively. Rays
from all other points on the object are focused correspondingly between these
limits. The triangles AnB and anb are similar; therefore, the relation of object
to image size is given by
Object size
Image size
Distance of object from nodal point
Distance of image from nodal point
(15.3)
or
AB
ab
An
an
222 Chapter 15 Optics
FIGURE 15.6 Determination of the image size on the retina.
Consider as an example the image of a person 180 cm tall standing 2 m
from the eye. The height of the full image at the retina is
Height of image 180 �
1.5
205
1.32 cm
The size of the face in the image is about 1.8 mm, and the nose is about
0.4 mm.
15.8 Retina
The retina consists of photoreceptor cells in contact with a complex network
of neurons and nerve fibers which are connected to the brain via the optic
nerve (see Fig. 15.7). Light absorbed by the photoreceptors produces nerve
impulses that travel along the neural network and then through the optic nerve
into the brain. The photoreceptors are located behind the neural network, so
the light must pass through this cell layer before it reaches the photoreceptors.
There are two types of photoreceptor cells in the retina: cones and rods.
The cones are responsible for sharp color vision in daylight. The rods provide
vision in dim light.
Near the center of the retina is a small depression about 0.3 mm in diameter
which is called the fovea. It consists entirely of cones packed closely together.
Each cone is about 0.002 mm (2 μm) in diameter. Most detailed vision is
obtained on the part of the image that is projected on the fovea. When the eye
scans a scene, it projects the region of greatest interest onto the fovea.
The region around the fovea contains both cones and rods. The structure
of the retina becomes more coarse away from the fovea. The proportion of
cones decreases until, near the edge, the retina is composed entirely of rods.
In the fovea, each cone has its own path to the optic nerve. This allows the
perception of details in the image projected on the fovea. Away from the
Section 15.9 Resolving Power of the Eye 223
FIGURE 15.7 The retina.
fovea, a number of receptors are attached to the same nerve path. Here the
resolution decreases, but the sensitivity to light and movement increases.
With the structure of the retina in mind, let us examine how we view a
scene from a distance of about 2 m. From this distance, at any one instant, we
can see most distinctly an object only about 4 cm in diameter. An object of
this size is projected into an image about the size of the fovea.
Objects about 20 cm in diameter are seen clearly but not with complete
sharpness. The periphery of large objects appears progressively less distinct.
Thus, for example, if we focus on a person’s face 2 m away, we can see clearly
the facial details, but we can pick out most clearly only a subsection about the
size of the mouth. At the same time, we are aware of the person’s arms and
legs, but we cannot detect, for example, details about the person’s shoes.
15.9 Resolving Power of the Eye
So far in our discussion of image formation we have used geometric optics,
which neglects the diffraction of light. Geometric optics assumes that light
from a point source is focused into a point image. This is not the case. When
224 Chapter 15 Optics
FIGURE 15.8 Two points are resolvable if the angle θ is greater than 1.22λ/d.
light passes through an aperture such as the iris, diffraction occurs, and the
wave spreads around the edges of the aperture.1 As a result, light is not
focused into a sharp point but into a diffraction pattern consisting of a disk
surrounded by rings of diminishing intensity.
If light originates from two point sources that are close together, their
image diffraction disks may overlap, making it impossible to distinguish the
two points. An optical system can resolve two points if their corresponding
diffraction patterns are distinguishable. This criterion alone predicts that two
points are resolvable (see Fig. 15.8) if the angular separation between the lines
joining the points to the center of the lens is equal to or greater than a critical
value given by
θ
1.22λ
d
(15.4)
where λ is the wavelength of light and d is the diameter of the aperture. The
angle θ is given in radians (1 rad 57.3◦). With green light (λ 500 nm) and
an iris diameter of 0.5 cm, this angle is 1.22 � 10−4 rad.
Experiments have shown that the eye does not perform this well. Most
people cannot resolve two points with an angular separation of less than
5 � 10−4 rad. Clearly there are other factors that limit the resolution of the
eye. Imperfections in the lens system of the eye certainly impede the resolution.
But perhaps even more important are the limitations imposed by the
structure of the retina.
The cones in the closely packed fovea are about 2 μm diameter. To resolve
two points, the light from each point must be focused on a different cone and
the excited cones must be separated from each other by at least one cone
that is not excited. Thus at the retina, the images of two resolved points are
separated by at least 4 μm. A single unexcited cone between points of excitation
implies an angular resolution of about 3 � 10−4 rad (see Exercise 15-7a).
1
If there are no smaller apertures in the optical path, the lens itself must be considered as the
aperture.
Section 15.10 Threshold of Vision 225
FIGURE 15.9 Resolution of the eye.
Some people with acute vision do resolve points with this separation, but most
people do not. We can explain the limits of resolution demonstrated by most
normal eyes if we assume that, to perceive distinct point images, there must be
three unexcited cones between the areas of excitation. The angular resolution
is then, as observed, 5 � 10−4 rad (see Exercise 15-7b).
Let us now calculate the size of the smallest detail that the unaided eye
can resolve. To observe the smallest detail, the object must be brought to the
closest point on which the eye can focus. Assuming that this distance is 20 cm
from the eye, the angle subtended by two points separated by a distance x is
given by (see Fig. 15.9)
tan−1 θ
2
x/2
20
(15.5)
If θ is very small, as is the case in our problem, the tangent of the angle is
equal to the angle itself and
θ
x
20
Because the smallest resolvable angle is 5 � 10−4 rad, the smallest resolvable
detail x is
x 5 � 10−4
� 20 100 μm 0.1 mm
Using the same criterion, we can show (see Exercise 15-8) that the facial
features such as the whites of the eye are resolvable from as far as 20 m.
15.10 Threshold of Vision
The sensation of vision occurs when light is absorbed by the photosensitive
rods and cones. At low levels of light, the main photoreceptors are the
226 Chapter 15 Optics
rods. Light produces chemical changes in the photoreceptors which reduce
their sensitivity. For maximum sensitivity the eye must be kept in the dark
(dark adapted) for about 30 minutes to restore the composition of the
photoreceptors.
Under optimum conditions, the eye is a very sensitive detector of light.
The human eye, for example, responds to light from a candle as far away
as 20 km. At the threshold of vision, the light intensity is so small that we
must describe it in terms of photons. Experiments indicate that an individual
photoreceptor (rod) is sensitive to 1 quantum of light. This, however, does not
mean that the eye can see a single photon incident on the cornea. At such low
levels of light, the process of vision is statistical.
In fact, measurements show that about 60 quanta must arrive at the cornea
for the eye to perceive a flash. Approximately half the light is absorbed or
reflected by the ocular medium. The 30 or so photons reaching the retina are
spread over an area containing about 500 rods. It is estimated that only 5 of
these photons are actually absorbed by the rods. It seems, therefore, that at
least 5 photoreceptors must be stimulated to perceive light.
The energy in a single photon is very small. For green light at 500 nm, it is
E hf
hc
λ
6.63 � 10−27 � 3 � 1010
5 � 10−5
3.98 � 10−12
erg
This amount of energy, however, is sufficient to initiate a chemical change in
a single molecule which then triggers the sequence of events that leads to the
generation of the nervous impulse.
15.11 Vision and the Nervous System
Vision cannot be explained entirely by the physical optics of the eye. There
are many more photoreceptors in the retina than fibers in the optic nerve. It is,
therefore, evident that the image projected on the retina is not simply transmitted
point by point to the brain. A considerable amount of signal processing
occurs in the neural network of the retina before the signals are transmitted
to the brain. The neural network “decides” which aspects of the image are
most important and stresses the transmission of those features. In a frog, for
example, the neurons in the retina are organized for most active response to
movements of small objects. A fly moving across the frog’s field of vision will
produce an intense neural response, and if the fly is close enough, the frog will
lash out its tongue to capture the fly. On the other hand, a large object, clearly
not food for the frog, moving in the same vision field will not elicit a neural
response. Evidently the optical processing system of the frog enhances its
Section 15.12 Defects in Vision 227
ability to catch small insects while reducing the likelihood of being noticed by
larger, possibly dangerous creatures passing through the neighborhood.
The human eye also possesses important processing mechanisms. It has
been shown that movement of the image is necessary for human vision as well.
In the process of viewing an object, the eye executes small rapid movements,
30 to 70 per second, which alter slightly the position of the image on the retina.
Under experimental conditions, it is possible to counteract the movement of
the eye and stabilize the position of the retinal image. It has been found that,
under these conditions, the image perceived by the person gradually fades.
15.12 Defects in Vision
There are three common defects in vision associated with the focusing system
of the eye: myopia (nearsightedness), hyperopia (farsightedness), and astigmatism.
The first two of these defects are best explained by examining the
imaging of parallel light by the eye.
The relaxed normal eye focuses parallel light onto the retina (Fig. 15.10).
In the myopic eye the lens system focuses the parallel light in front of the
retina (Fig. 15.11a). This misfocusing is usually caused by an elongated eyeball
or an excessive curvature of the cornea. In hyperopia the problem is
reversed (see Fig. 15.12a). Parallel light is focused behind the retina. The
problem here is caused by an eyeball that is shorter than normal or by the
inadequate focusing power of the eye. The hyperopic eye can accommodate
objects at infinity, but its near point is farther away than is normal. Hyperopia
is, thus, similar to presbyopia. These two defects can be summarized as follows:
The myopic eye converges light too much, and the hyperopic eye not
enough.
Astigmatism is a defect caused by a nonspherical cornea. An oval-shaped
cornea, for example, is more sharply curved along one plane than another;
therefore, it cannot form simultaneously sharp images of two perpendicular
lines. One of the lines is always out of focus, resulting in distorted vision.
FIGURE 15.10 The normal eye.
228 Chapter 15 Optics
FIGURE 15.11 (a) Myopia. (b) Its correction.
FIGURE 15.12 (a) Hyperopia. (b) Its correction.
FIGURE 15.13 Cylindrical lens for astigmatism.
All three of these defects can be corrected by lenses placed in front of the
eye. Myopia requires a diverging lens to compensate for the excess refraction
in the eye. Hyperopia is corrected by a converging lens, which adds to the
focusing power of the eye. The uneven corneal curvature in astigmatism is
compensated for by a cylindrical lens (Fig. 15.13), which focuses light along
one axis but not along the other.
Section 15.15 Extension of Vision 229
15.13 Lens for Myopia
Let us assume that the farthest object a certain myopic eye can properly focus
is 2 m from the eye. This is called the far point of the eye. Light from objects
farther away than this is focused in front of the retina (Fig. 15.11a). Here the
purpose of the corrective lens is to make parallel light appear to come from
the far point of the eye (in this case, 2 m). With such a corrective lens, the eye
is able to form images of objects all the way to infinity.
The focal length of the lens is obtained by using Eq. C.6, which is
1
p
+
1
q
1
f
Here p is infinity, as this is the effective distance for sources of parallel light.
The desired location q for the virtual image is −200 cm. The focal length of
the diverging lens (see Eq. C.4) is, therefore,
1
f
1
∞
+
1
−200
or f −200 cm −5 diopters
15.14 Lens for Presbyopia and Hyperopia
In these disorders, the eye cannot focus properly on close objects. The near
point is too far from the eye. The purpose of the lens is to make light from
close objects appear to come from the near point of the unaided eye. Let us
assume that a given hyperopic eye has a near point at 150 cm. The desired
lens is to allow the eye to view objects at 25 cm. The focal length of the lens
is again obtained from Eq. C.6,
1
p
+
1
q
1
f
Here p is the object distance at 25 cm and q is −150 cm, which is the distance
of the virtual image at the near point. The focal length f for the converging
lens is given by
1
f
1
25 cm
−
1
150 cm
or f 30 cm 33.3 diopters
15.15 Extension of Vision
The range of vision of the eye is limited. Details on distant objects cannot be
seen because their images on the retina are too small. The retinal image of a
230 Chapter 15 Optics
20 m-high tree at a distance of 500 m is only 0.6 mm high. The leaves on this
tree cannot be resolved by the unaided eye (see Exercise 15-9). Observation
of small objects is limited by the accommodation power of the eye. We have
already shown that, because the average eye cannot focus light from objects
closer than about 20 cm, its resolution is limited to approximately 100 μm.
Over the past 300 years, two types of optical instruments have been developed
to extend the range of vision: the telescope and the microscope. The
telescope is designed for the observation of distant objects. The microscope
is used to observe small objects that cannot be seen clearly by the naked eye.
Both of these instruments are based on the magnifying properties of lenses.
A third more recent aid to vision is the fiberscope which utilizes total internal
reflection to allow the visualization of objects normally hidden from view.
15.15.1 Telescope
A drawing of a simple telescope is shown in Fig. 15.14. Parallel light from a
distant object enters the first lens, called the objective lens or objective, which
forms a real inverted image of the distant object. Because light from the distant
object is nearly parallel, the image is formed at the focal plane of the objective.
(The drawing shows the light rays from only a single point on the object.) The
second lens, called the eyepiece, magnifies the real image. The telescope is
adjusted so that the real image formed by the objective falls just within the
focal plane of the eyepiece. The eye views the magnified virtual image formed
by the eyepiece. The total magnification—the ratio of image to object size—is
given by
Magnification −
f1
f2
(15.6)
FIGURE 15.14 The telescope.
Section 15.15 Extension of Vision 231
where f1 and f2 are the focal lengths of the objective and the eyepiece
respectively. As can be seen from Eq. 15.6, greatest magnification is obtained
with a long focal-length objective and a short focal-length eyepiece.
15.15.2 Microscope
A simple microscope consists of a single lens that magnifies the object
(Fig. 15.15). Better results can be obtained, however, with a two-lens system
compound microscope, shown in Fig. 15.16. The compound microscope,
like the telescope, consists of an objective lens and an eyepiece, but the
objective of the microscope has a short focal length. It forms a real image I1
of the object; the eye views the final magnified image I2 formed by the
eyepiece.
The microscope is an important tool in the life sciences. Its invention
in the 1600s marked the beginning of the study of life on the cellular level.
The early microscope produced highly distorted images, but years of development
perfected the device nearly to its theoretical optimum. The resolution
of the best modern microscopes is determined by the diffraction properties of
light, which limit the resolution to about half the wavelength of light. In other
FIGURE 15.15 Simple magnifier.
FIGURE 15.16 Schematic diagram of a compound microscope.
232 Chapter 15 Optics
words, with a good modern microscope, we can observe objects as small as
half the wavelength of the illuminating light.
We will not present here the details of microscopy. These can be found
in many basic physics texts (see, for example, [15-1]). We will, however,
describe a special-purpose scanning confocal microscope designed in our laboratory
by Paul Davidovits and M. David Egger.
15.15.3 Confocal Microscopy
With conventional microscopes, it is not possible to observe small objects
embedded in translucent materials. For example, cells located beneath the
surface of tissue, such as buried brain cells in living animals, cannot be satisfactorily
observed with conventional microscopes.
Light can certainly penetrate through tissue. This can be demonstrated
simply by inserting a flashlight into the mouth and observing the light passing
through the cheeks. In principle, therefore, we should be able to form a magnified
image of a cell inside the tissue. This could be done by shining light
into the tissue and collecting the light reflected from the cell. Unfortunately
there is a problem associated with the straightforward use of this technique.
Light is reflected and scattered not only by the cell of interest but also by the
surface of the tissue and by the cells in front and behind the cell being examined.
This spurious light is also intercepted by the microscope and masks the
image of the single cell layer within the tissue (see Fig. 15.17).
FIGURE 15.17 Light scattered and reflected from tissue.
Section 15.15 Extension of Vision 233
Over the years, a number of microscopes have been designed that
attempted to solve this problem. The most successful of these is the confocal
microscope. The principle of confocal microscopy was first described by
Marvin Minsky in 1957. In the 1960s, Davidovits and Egger modified the
Minsky design and built the first successful confocal microscope for observation
of cells within living tissue.
The confocal microscope is designed to accept light only from a thin
slice within the tissue and to reject light reflected and scattered from other
regions. A schematic diagram of the Davidovits-Egger microscope is shown
in Fig. 15.18. Although the device does not resemble a conventional microscope
it certainly does produce magnified images. This microscope requires a
parallel beam of light for illumination of the object. As the source of parallel
FIGURE 15.18 Confocal microscope.
234 Chapter 15 Optics
light we used a laser with a power output that is relatively low so that it does
not damage the tissue under observation. The laser beam is reflected by a
half-silvered mirror into the objective lens, which focuses the beam to a point
inside the tissue. Because the light is parallel, the beam is brought to a point
at the principal focus of the lens. The depth of this point in the tissue can be
changed by altering the distance between the lens and the tissue.
Light is scattered and reflected from all points in the path of the entering
light, and part of this returning light is intercepted by the objective lens. However,
only light originating from the focal point emerges from the lens as a
parallel beam; light from all other points either converges toward or diverges
from the lens axis. The returning light passes through the half-silvered mirror
and is intercepted by the collecting lens. Only the parallel component of the
light is focused into the small exit aperture that is placed at the principal focal
point of the collecting lens. Nonparallel light is defocused at the exit aperture.
A photomultiplier placed behind the exit aperture produces a voltage proportional
to the light intensity transmitted through the exit aperture. This voltage
is then used to control the intensity of an electron beam in the oscilloscope.
So far, we have one spot on the screen of the oscilloscope which glows
with a brightness proportional to the reflectivity of one point inside the tissue.
In order to see a whole cell or region of cells, we must scan the region
point by point. This is done by moving the lens in its own plane so that the
focal point scans an area inside the tissue. The motion of the lens does not
affect the parallelism of the light originating at the focal point of the objective
lens. Therefore, at every instant, the output of the photomultiplier and
the corresponding brightness of the spot on the screen are proportional to the
reflectivity of the point being scanned. While the object is scanned, the electron
beam in the oscilloscope is moved in synchrony with the motion of the
objective lens. Thus, the screen shows a picture of a very thin section within
the tissue.
The magnification of this microscope is simply the ratio of the electron
beam excursion on the oscilloscope face to the excursion of the scanning lens.
For a 0.1-mm excursion of the lens, the electron beam may be adjusted to
move 5 cm. The magnification is then 500. The resolution of the device is
determined by the size of the spot focused by the objective. The diffraction
properties of light limit the minimum spot size to about half the wavelength of
light. The optimum resolution is, therefore, about the same as in conventional
microscopes.
The first biologically significant observations with the confocal microscope
were those of endothelial cells on the inside of the cornea in live frogs.
Such observations cannot be made with conventional microscopes because the
light reflected from the front surface of the cornea masks the weak
reflections from the endothelial cells. The picture of these cells shown in
Section 15.15 Extension of Vision 235
FIGURE 15.19 Corneal endothelial cells in an intact eye of a living bullfrog. Arrows
indicate outlines of the nuclei in two of the cells. Calibration mark, 25 μm.
Fig. 15.19 was obtained by photographing the image on the oscilloscope
screen. The confocal microscope is now a major observational tool in most
biology laboratories. In the more recent versions of the instrument the object
is scanned with moving mirrors and the image is processed by computers.
Image improvement obtained with modern confocal microscopy is illustrated
in Fig. 15.20.
15.15.4 Fiber Optics
Fiber-optic devices are now used in a wide range of medical applications. The
principle of their operation is simple. As discussed in Appendix C in connection
with Snell’s law, light traveling in a material of high index of refraction is
totally reflected back into the material if it strikes the boundary of the material
with lower refractive index at an angle greater than the critical angle θc. In
this way, light can be confined to travel within a glass cylinder as shown in
Fig. 15.21. This phenomenon has been well known since the early days of
optics. However, major breakthroughs in materials technology were necessary
before the phenomenon could be widely utilized.
Optical fiber technology, developed in the 1960s and 1970s made it possible
to manufacture low-loss, thin, highly flexible glass fibers that can carry
light over long distances. A typical optical fiber is about 10 μm in diameter
and is made of high purity silica glass. The fiber is coated with a cladding
to increase light trapping. Such fibers can carry light over tortuously twisting
paths for several kilometers without significant loss.
236 Chapter 15 Optics
FIGURE 15.20 Microscope images of sea urchin embryos obtained with (a) a
conventional microscope showing out-of-focus blur and (b) a modern confocal
microscope. Part (a) from Matsumoto (1993), Meth. Cell Biol. 38, p. 22. Part (b) from
Wright (1989), J. Cell. Sci. 94, 617–624, with permission from the Company of
Biologists Ltd.
Fiberscopes or endoscopes are the simplest of the fiber-optic medical
devices. They are used to visualize and examine internal organs such as the
stomach, heart, and bowels. A fiberscope consists of two bundles of optical
fibers tied into one flexible unit. Each bundle is typically a millimeter in diameter
consisting of about 10,000 fibers. For some applications, the bundles are
thicker, up to about 1.5 cm in diameter. Depending on their use, the bundles
vary in length from 0.3 to 1.2 m.
Chapter 15 Exercises 237
FIGURE 15.21 Light confined to travel inside a glass cylinder by total reflection.
The two bundles as a unit are introduced into the body through orifices,
veins, or arteries and are threaded toward the organ to be examined. Light
from a high intensity source, such as a xenon arc lamp, is focused into one
bundle which carries the light to the organ to be examined. Each of the fibers
in the other bundle collects light reflected from a small region of the organ
and carries it back to the observer. Here the light is focused into an image
which can be viewed by eye or displayed on a cathode ray or some other
type of electronic screen. In the usual arrangement, the illuminating bundle
surrounds the light-collecting bundle. Most endoscopes now utilize attached
miniature video cameras to form images of the internal organs for display on
TV monitors.
The use of fiber-optic devices has been greatly expanded by attaching to
the endoscope remotely controlled miniature instruments to perform surgical
operations without major surgical incisions. More recent applications
of fiber optics include measurement of pressure in arteries, bladder, and
uterus using optical sensors and laser surgery where powerful laser light is
directed through one of the bundles to the tissue which is selectively destroyed
(see Chapter 16).
EXERCISES
15-1. Compute the change in the position of the image formed by a lens with
a focal length of 1.5 cm as the light source is moved from its position at
6 m from the lens to infinity.
15-2. A point source of light that is not exactly in focus produces a disk image
at the retina. Assume that the image is acceptable provided the image
diameter of the defocused point source is less than a. Show that the
depth of field is inversely proportional to the diameter of the aperture.
238 Chapter 15 Optics
15-3. Using data presented in the text, calculate the focusing power of the
cornea and of the crystalline lens.
15-4. Calculate the refractive power of the cornea when it is in contact with
water. The index of refraction for water is 1.33.
15-5. Calculate the focusing power of the lens in the fish eye. Assume that
the lens is spherical with a diameter of 2 mm. (The indices of refraction
are as in Table 15.1.) The index of refraction for water is 1.33.
15-6. Calculate the distance of the point in front of the cornea at which parallel
light originating inside the reduced eye is focused.
15-7. Using the dimensions of the reduced eye (Fig. 15.5), calculate the angular
resolution of the eye (use Fig. 15.6 as an aid) (a) with a single unexcited
cone between points of excitation (b) with four unexcited cones
between areas of excitation.
15-8. Calculate the distance from which a person with good vision can see
the whites of another person’s eyes. Use data in the text and assume the
size of the eye is 1 cm.
15-9. Calculate the size of the retinal image of a 10-cm leaf from a distance
of 500 m.
Chapter 16
Atomic Physics
Modern atomic and nuclear physics is among the most impressive scientific
achievements of this century. There is hardly an area of science or technology
that does not draw on the concepts and techniques developed in this field.
Both the theories and techniques of atomic and nuclear physics have played
an important role in the life sciences. The theories provided a solid foundation
for understanding the structure and interaction of organic molecules, and
the techniques provided many tools for both experimental and clinical work.
Contributions from this field have been so numerous and influential that it is
impossible to do them justice in a single chapter. Of necessity, therefore, our
discussion will be restricted to a survey of the subject. We will present a brief
description of the atom and the nucleus, which will lead into a discussion of
the applications of atomic and nuclear physics to the life sciences.
16.1 The Atom
By 1912, through the work of J. J. Thompson, E. Rutherford, and their
colleagues, a number of important facts had been discovered about atoms
which make up matter. It was found that atoms contain small negatively
charged electrons and relatively heavier positively charged protons. The proton
is about 2000 times heavier than the electron, but the magnitude of the
charge on the two is the same. There are as many positively charged protons
in an atom as negatively charged electrons. The atom as a whole is, therefore,
electrically neutral. The identity of an atom is determined by the number of
239
240 Chapter 16 Atomic Physics
protons it has. For example, hydrogen has 1 proton, carbon has 6 protons,
silver has 47 protons. Through a series of ingenious experiments, Rutherford
showed that most of the atomic mass is concentrated in the nucleus consisting
of protons and that the electrons are somehow situated outside the nucleus.
It was subsequently discovered that the nucleus also contains another particle,
the neutron, which has approximately the same mass as the proton but is
electrically neutral.
Although the nucleus contains most of the atomic mass, it occupies only a
small part of the total atomic volume. The diameter of the whole atom is on
the order of 10−8 cm, but the diameter of the nucleus is only about 10−13 cm.
The configuration of the electrons around the nucleus was not known at that
time.
In 1913, the Danish physicist Niels Bohr proposed a model for the atom
which explained many observations that were puzzling scientists at that time.
When Bohr first became acquainted with atomic physics, the subject was in
a state of confusion. A number of theories had been proposed for the structure
of the atom, but none explained satisfactorily the existing experimental
results. The most surprising observed property of atoms was the light emitted
by them.1 When an element is put into a flame, it emits light at sharply defined
wavelengths, called spectral lines. Each element emits its own characteristic
spectrum of light. This is in contrast to a glowing filament in a light bulb, for
example, which emits light over a continuous range of wavelengths.
Prior to Bohr, scientists could not explain why these colors were emitted
by atoms. Bohr’s model of the atom explained the reason for the sharp spectra.
Bohr started with the model of the atom as proposed by Rutherford. At the
center of the atom is the positive nucleus made up of protons (and neutrons).
The electrons orbit around the nucleus much as the planets orbit around the
sun. They are maintained in orbit by the electrostatic attraction of the nucleus.
And here is the major feature of the Bohr model: So that the model would
explain the emission of spectral lines, Bohr had to postulate that the electrons
are restricted to distinct orbits around the nucleus. In other words, the electrons
can be found only in certain allowed orbits. Bohr was able to calculate
the radii of these allowed orbits and show that the spectral lines are emitted
as a consequence of the orbital restrictions. Bohr’s calculations are found in
most elementary physics texts.
The orbital restrictions are most easily illustrated with the simplest atom,
hydrogen, which has a single-proton nucleus and one electron orbiting around
it (Fig. 16.1). Unless energy is added to the atom, the electron is found in the
allowed orbit closest to the nucleus. If energy is added to the atom, the electron
1
In atomic physics, the word light is not restricted solely to the visible part of the electromagnetic
spectrum. Radiation at shorter wavelength (ultraviolet) and longer wavelength (infrared)
is also often referred to as light.
Section 16.1 The Atom 241
FIGURE 16.1 Bohr model for the hydrogen atom. The electron orbits about the
nucleus and can occupy only discrete orbits with radii 1, 2, 3, and so on.
may “jump” to one of the higher allowed orbits farther away from the nucleus,
but the electron can never occupy the regions between the allowed orbits.
The Bohr model was very successful in explaining many of the experimental
observations for the simple hydrogen atom. But to describe the behavior of
atoms with more than one electron, it was necessary to impose an additional
restriction on the structure of the atom: The number of electrons in a given
orbit cannot be greater than 2n2, where n is the order of the orbit from the
nucleus. Thus, the maximum number of electrons in the first allowed orbit is
2 � (1)2 2; in the second allowed orbit, it is 2 � (2)2 8; in the third orbit,
it is 2 � (3)2 18, and so on.
The atoms are found to be constructed in accordance with these restrictions.
Helium has two electrons, and, therefore, its first orbit is filled. Lithium
has three electrons, two of which fill the first orbit; the third electron, therefore,
must be in the second orbit. This simple sequence is not completely
applicable to the very complex atoms, but basically this is the way the elements
are constructed.
A specific amount of energy is associated with each allowed orbital configuration
of the electron. Therefore, instead of speaking of the electron as
being in a certain orbit, we can refer to it as having a corresponding amount
of energy. Each of these allowed values of energy is called an energy level.
An energy level diagram for an atom is shown in Fig. 16.2. Note that every
element has its own characteristic energy level structure. The electrons in the
atom can occupy only specific energy states; that is, in a given atom the electron
can have an energy E1, E2, E3, and so on, but cannot have an energy
between these two values. This is a direct consequence of the restrictions on
the allowed electron orbital configurations.
The lowest energy level that an electron can occupy is called the ground
state. This state is associated with the orbital configuration closest to the
242 Chapter 16 Atomic Physics
FIGURE 16.2 Energy levels for an atom.
nucleus. The higher allowed energy levels, called excited states, are associated
with larger orbits and different orbital shapes. Normally the electron occupies
the lowest energy level but it can be excited into a higher energy state by
adding energy to the atom.
An atom can be excited from a lower to a higher energy state in a number
of different ways. The two most common methods of excitation are electron
impact and absorption of electromagnetic radiation. Excitation by electron
impact occurs most frequently in a gas discharge. If a current is passed
through a gas of atoms, the colliding electron is slowed down and the electron
in the atom is promoted to a higher energy configuration. When the excited
atoms fall back into the lower energy states, the excess energy is given off as
electromagnetic radiation. Each atom releases its excess energy in a single
photon. Therefore, the energy of the photon is simply the difference between
the energies of the initial state Ei and the final state Ef of the atom. The
frequency f of the emitted radiation is given by
f
Energy of photon
Planck constant
Ei − Ef
h
(16.1)
Transition between each pair of energy levels results in the emission of
light at a specific frequency, called transition or resonance frequency. Therefore,
a group of highly excited atoms of a given element emit light at a number
of well-defined frequencies which constitute the optical spectrum for that
element.
An atom in a given energy level can also be excited to a higher level by
light at a specific frequency. The frequency must be such that each photon
has just the right amount of energy to promote the atom to one of its higher
allowed energy states. Atoms, therefore, absorb light only at the specific
transition frequencies, given by Eq. 16.1. Light at other frequencies is not
Section 16.1 The Atom 243
FIGURE 16.3 The absorption spectrum.
absorbed. If a beam of white light (containing all the frequencies) is passed
through a group of atoms of a given species, the spectrum of the transmitted
light shows gaps corresponding to the absorption of the specific frequencies
by the atoms. This is called the absorption spectrum of the atom. In their
undisturbed state, most of the atoms are in the ground state. The absorption
spectrum, therefore, usually contains only lines associated with transitions
from the ground state to higher allowed states (Fig. 16.3).
Optical spectra are produced by the outer electrons of the atom. The inner
electrons, those closer to the nucleus, are bound more tightly and are consequently
more difficult to excite. However, in a highly energetic collision with
another particle, an inner electron may be excited. When in such an excited
atom an electron returns to the inner orbit, the excess energy is again released
as a quantum of electromagnetic radiation. Because the binding energy here
is about a thousand times greater than for the outer electrons, the frequency of
the emitted radiation is correspondingly higher. Electromagnetic radiation in
this frequency range is called X-rays.
The Bohr model also explained qualitatively the formation of chemical
bonds. The formation of chemical compounds and matter in bulk is due to
the distribution of electrons in the atomic orbits. When an orbit is not filled
244 Chapter 16 Atomic Physics
FIGURE 16.4 A schematic representation for the formation of a hydrogen molecule.
(a) Two separate hydrogen atoms. (b) When the two atoms are close together, the
electrons share each other’s orbit, which results in the binding of the two atoms into a
molecule.
to capacity (which is the case for most atoms), the electrons of one atom can
partially occupy the orbit of another. This sharing of orbits draws the atoms
together and produces bonding between atoms. As an example, we show in
Fig. 16.4 the formation of a hydrogen molecule from two hydrogen atoms. In
the orbit of each of the hydrogen atoms there is room for another electron.
A completely filled orbit is the most stable configuration; therefore, when two
hydrogen atoms are close together, they share each other’s electrons, and, in
this way, the orbit of each atom is completely filled part of the time. This
shared orbit can be pictured as a rubber band pulling the two atoms together.
Therefore, the sharing of the electrons binds the atoms into a molecule. While
the sharing of electrons pulls the atoms together, the coulomb repulsion of the
nuclei tends to keep them apart. The equilibrium separation between atoms in
a molecule is determined by these two counter forces. In a similar way, more
complex molecules, and ultimately bulk matter, are formed.
Atoms with completely filled orbits (these are atoms of the so-called noble
gases—helium, neon, argon, krypton, and xenon) cannot share electrons with
other elements and are, therefore, chemically most inert.
Molecules also have characteristic spectra both in emission and in absorption.
Because molecules are more complicated than atoms, their spectra are
correspondingly more complex. In addition to the electronic configuration,
these spectra also depend on the motion of the nuclei. Still the spectra can be
interpreted and are unique for each type of molecule.
16.2 Spectroscopy
The absorption and emission spectra of atoms and molecules are unique for
each species. They can serve as fingerprints in identifying atoms and molecules
Section 16.2 Spectroscopy 245
in various substances. Spectroscopic techniques were first used in basic experiments
with atoms and molecules, but they were soon adopted in many other
areas, including the life sciences.
In biochemistry, spectroscopy is used to identify the products of complex
chemical reactions. In medicine, spectroscopy is used routinely to determine
the concentration of certain atoms and molecules in the body. From a spectroscopic
analysis of urine, for example, one can determine the level of mercury
in the body. Blood-sugar level is measured by first producing a chemical reaction
in the blood sample which results in a colored product. The concentration
of this colored product, which is proportional to the blood-sugar level, is then
measured by absorption spectroscopy.
The basic principles of spectroscopy are simple. In emission spectroscopy
the sample under investigation is excited by an electric current or a flame. The
emitted light is then examined and identified. In absorption spectroscopy,
the sample is placed in the path of a beam of white light. Examination of
the transmitted light reveals the missing wavelengths which identify the components
in the substance. Both the absorption and the emission spectra can
provide information also about the concentration of the various components
in the substance. In the case of emission, the intensity of the emitted light
in the spectrum is proportional to the number of atoms or molecules of the
given species. In absorption spectroscopy, the amount of absorption can be
related to the concentration. The instrument used to analyze the spectra is
called a spectrometer. This device records the intensity of light as a function
of wavelength.
A spectrometer, in its simplest form, consists of a focusing system, a
prism, and a detector for light (see Fig. 16.5). The focusing system forms
a parallel beam of light which falls on the prism. The prism, which can be
rotated, breaks up the beam into its component wavelengths. At this point, the
fanned-out spectrum can be photographed and identified. Usually, however,
the spectrum is detected a small section at a time. This is accomplished by the
narrow exit slit which intercepts only a portion of the spectrum. As the prism
is rotated, the whole spectrum is swept sequentially past the slit. The position
of the prism is calibrated to correspond with the wavelength impinging on the
slit. The light that passes through the slit is detected by a photodetector which
produces an electrical signal proportional to the light intensity. The intensity
of the signal as a function of wavelength can be displayed on a chart recorder.
Spectrometers used in routine clinical work are automated and can be operated
by relatively unskilled personnel. The identification and interpretation of
the spectra produced by less well-known molecules, however, require considerable
training and skill. In addition to identifying the molecule, such spectra
also yield information about the molecular structure. The use of spectrometers
is further explored in Exercise 16-1.
246 Chapter 16 Atomic Physics
FIGURE 16.5 The measurement of spectra.
16.3 Quantum Mechanics
Although the Bohr model explained many observations, from the very beginning
the theory appeared contrived. Certainly the concept of stable allowed
orbits with a specific number of electrons seemed arbitrary. The model,
however, was a daring step in a new direction that eventually led to the development
of quantum mechanics.
In the quantum mechanical description of the atom, it is not possible to
assign exact orbits or trajectories to electrons. Electrons possess wavelike
properties and behave as clouds of specific shape around the nucleus. The
artificial postulates in Bohr’s theory are a natural consequence of the quantum
mechanical approach to the atom. Furthermore, quantum mechanics explains
many phenomena outside the scope of the Bohr model. The shape of simple
molecules, for example, can be shown to be the direct consequence of the
interaction between the electron configurations in the component atoms.
The concept that particles may exhibit wavelike properties was introduced
in 1924 by Louis de Broglie. This suggestion grew out of an analogy with
light which was then known to have both wave- and particlelike properties. De
Broglie suggested by analogy that particles may exhibit wavelike properties.
He showed that the wavelength λ of the matter waves would be
λ
h
mv
(16.2)
Section 16.4 Electron Microscope 247
FIGURE 16.6 The electron microscope.
where m and v are the mass and velocity of the particle and h is the Planck
constant.
In 1925, de Broglie’s hypothesis was confirmed by experiments which
showed that electrons passing through crystals form wavelike diffraction
patterns with a configuration corresponding to a wavelength given by Eq. 16.2.
16.4 Electron Microscope
In Chapter 15, we pointed out that the size of the smallest object observable
by a microscope is about half the wavelength of the illuminating radiation.
In light microscopes, this limits the resolution to about 200 nm (2000 ˚A).
Because of the wave properties of electrons, it is possible to construct microscopes
with a resolution nearly 1000 times smaller than this value.
It is relatively easy to accelerate electrons in an evacuated chamber to high
velocities so that their wavelength is less than 10−10 m (1 ˚A). Furthermore,
the direction of motion of the electrons can be altered by electric and magnetic
fields. Thus, suitably shaped fields can act as lenses for the electrons.
The short wavelength of electrons coupled with the possibility of focusing
them has led to the development of electron microscopes that can observe
objects 1000 times smaller than are visible with light microscopes. The basic
construction of an electron microscope is shown in Fig. 16.6. The similarities
between the electron and the light microscope are evident: Both have the
same basic configuration of two lenses which produce two-stage magnification.
Electrons are emitted from a heated filament and are then accelerated
and collimated into a beam. The beam passes through the thin sample under
examination which diffracts the electrons in much the same way as light is
diffracted in an optical microscope. But because of their short wavelength,
248 Chapter 16 Atomic Physics
FIGURE 16.7 Electron micrograph of an individual axon in the peripheral nerve of
a mouse. The cross section of the axon at the level of the node of Ranvier is about 2.5 μm
in width. Surrounding the axon is a differentiated region of the myelin sheath.
(Photograph courtesy of Professor Dan Kirschner, Biology Department, Boston
College, and Dr. Bela Kosaras, Primate Center, Southborough, MA.)
the electrons are influenced by much smaller structures within the sample.
The transmitted electrons are focussed into a real image by the objective lens.
This image is then further magnified by the projector lens, which projects
the final image onto film or a fluorescent screen. Although it is possible to
produce electrons with a wavelength much less than 10−10 m (1 ˚A), the
theoretical optimum resolution implied by such short wavelengths has not yet
been realized. At present, the best resolution of electron microscopes is about
5 � 10−10 m (5 ˚A).
Because electrons are scattered by air, the microscope must be contained
in an evacuated chamber. Furthermore, the samples under examination must
be dry and thin. These conditions, of course, present some limitations in the
study of biological materials. The samples have to be specially prepared for
electron microscopic examination. They must be dry, thin, and in some cases
coated. Nevertheless, electron microscopes have yielded beautiful pictures
showing details in cell structure, biological processes, and recently even large
molecules such as DNA in the process of replication (see Fig. 16.7).
Section 16.5 X-rays 249
16.5 X-rays
In 1895, Wilhelm Conrad Roentgen announced his discovery of X-rays. He
had found that when high-energy electrons hit a material such as glass, the
material emitted radiation that penetrated objects which are opaque to light.
He called this radiation X-rays. It was shown subsequently that X-rays are
short-wavelength, electromagnetic radiation emitted by highly excited atoms.
Roentgen showed that X-rays could expose film and produce images of objects
in opaque containers. Such pictures are possible if the container transmits
X-rays more readily than the object inside. A film exposed by the X-rays
shows the shadow cast by the object.
Within three weeks of Roentgen’s announcement, two French physicians,
Oudin and Barth�elemy, obtained X-rays of bones in a hand. Since then,
X-rays have become one of the most important diagnostic tools in medicine.
With current techniques, it is even possible to view internal body organs that
are quite transparent to X-rays. This is done by injecting into the organ a fluid
opaque to X-rays. The walls of the organ then show up clearly by contrast.
X-rays have also provided valuable information about the structure of
biologically important molecules. The technique used here is called crystallography.
The wavelength of X-rays is on the order of 10−10 m, about the
same as the distance between atoms in a molecule or crystal. Therefore, if
a beam of X-rays is passed through a crystal, the transmitted rays produce a
diffraction pattern that contains information about the structure and composition
of the crystal. The diffraction pattern consists of regions of high and low
X-ray intensity which when photographed show spots of varying brightness
(Fig. 16.8).
FIGURE 16.8 Arrangement for detecting diffraction of X-rays by a crystal.
250 Chapter 16 Atomic Physics
Diffraction studies are most successfully done with molecules that can be
formed into a regular periodic crystalline array. Many biological molecules
can in fact be crystallized under the proper conditions. It should be noted,
however, that the diffraction pattern is not a unique, unambiguous picture of
the molecules in the crystal. The pattern is a mapping of the collective effect of
the arrayed molecules on the X-rays that pass through the crystal. The structure
of the individual molecule must be deduced from the indirect evidence
provided by the diffraction pattern.
If the crystal has a simple structure—such as sodium chloride, for
example—the X-ray diffraction pattern is also simple and relatively easy to
interpret. Complicated crystals, however, such as those synthesized from
organic molecules, produce very complex diffraction patterns. But, even in
this case, it is possible to obtain some information about the structure of the
molecules forming the crystal (for details, see [16-1]). To resolve the threedimensional
features of the molecules, diffraction patterns must be formed
from thousands of different angles. The patterns are then analyzed, with the
aid of a computer. These types of studies provided critical information for
the determination of the structure for penicillin, vitamin B12, DNA, and many
other biologically important molecules.
16.6 X-ray Computerized Tomography
The usual X-ray picture does not provide depth information. The image
represents the total attenuation as the X-ray beam passes through the object
in its path. For example, a conventional X-ray of the lung may reveal the
existence of a tumor, but it will not show how deep in the lung the tumor
is located. Several tomographic techniques (CT scans) have been developed
to produce slice-images within the body which provide depth information.
(Tomography is from the Greek word tomos meaning section.) Presently the
most commonly used of these is X-ray computerized tomography (CT scan)
developed in the 1960s. The basic principle of the technique in its simplest
form is illustrated in Fig. 16.9a and b. A thin beam of X-rays passes through
the plane we want to visualize and is detected by a diametrically opposing
detector. For a given angle with respect to the object (in this case the head),
the X-ray source-detector combination is moved laterally scanning the region
of interest as shown by the arrow in Fig. 16.9a. At each position, the detected
signal carries integrated information about X-ray transmission properties of
the full path in this case A−B. The angle is then changed by a small amount
(about 1◦) and the process is repeated full circle around the object. As indicated
in Fig. 16.9b, by rotating the source-detector combination, information
can be obtained about the intersection points of the X-ray beams.
Section 16.6 X-ray Computerized Tomography 251
FIGURE 16.9 (a) Basic principle of computed axial tomography. (b) Rotation of
the source-detector combination provides information about the X-ray transmission
properties of each point within the plane of the object to be studied.
In Fig. 16.9b, we show schematically the scanning beam at two angles with
two lateral positions at each angle. While at each position, the detected signal
carries integrated information about the full path, two paths that intersect contain
common information about the one point of intersection. In the figure,
four such points are shown at the intersection of the beams A−B, A − B ,
C−D, and C −D . The multiple images obtained by translation and rotation
contain information about the X-ray transmission properties of each point
within the plane of the object to be studied. These signals are stored and by a
rather complex computer analysis a point by point image is constructed of the
thin slice scanned within the body.
The visualized slices within the body obtained in this way are typically
about 2 mm thick. In the more recent versions of the instrument, a fan rather
than a beam of X-rays scans the object, and an array of multiple detectors is
used to record the signal. Data acquisition is speeded up in this way yielding
an image in a few seconds.
252 Chapter 16 Atomic Physics
16.7 Lasers
As was pointed out in Section 16.1, when light at the frequency corresponding
to the transition between two energy levels of atoms (or molecules) is passed
through a collection of these atoms, photons are absorbed from the light beam
by atoms in the lower energy level raising them to the higher (excited) level.
Atoms in an excited level can return to the lower state by emitting a photon at
the corresponding resonance frequency (see Eq. 16.1). This type of emission
is called spontaneous emission. However, atoms in an excited state can emit
photons also in another way.
In 1916, Albert Einstein analyzed the interaction of electromagnetic radiation
with matter using quantum mechanics and equilibrium considerations.
His results showed that while light interacting with atoms in a lower energy
state is absorbed, there is a parallel interaction of light with atoms in the
excited energy state. The light at the resonance frequency interacts with the
excited atoms by stimulating them to make a transition back into the lower
energy state. In the process, each stimulated atom emits a photon at the resonance
frequency and in phase with the stimulating light. This type of light
emission is called stimulated emission.
In a collection of atoms or molecules under equilibrium conditions, more
atoms are in a lower energy state than in a higher one. When a beam of light
at resonance frequency passes through a collection of atoms in equilibrium,
more photons are taken out of the beam by absorption than are added to it
by stimulated emission and the light beam is attenuated. However, through a
variety of techniques it is possible to reverse the normal situation and cause
more atoms to occupy a higher than a lower energy state. A collection of
atoms, with more atoms occupying the higher state, is said to have an inverted
population distribution. When light at resonance frequency passes through
atoms with inverted population distribution, more photons are added to the
beam by stimulated emission than are taken out of the beam by absorption.
As a result the intensity of the light beam increases. In other words, the light
is amplified. A medium with an inverted population can be made into a special
type of light source called a laser (light amplification by stimulated emission
of radiation) (see Exercises 16-3 and 16-4).
Light emitted by a laser has some unique properties. It can be formed into
a highly parallel beam that can be subsequently focused into a very small area,
typically on the order of the wavelength of light. In this way a large amount
of energy can be delivered into a small region with high degree of positional
precision. Further, the light emitted by a laser is monochromatic (single color)
with the wavelength determined by the amplifying medium.
The first laser was built in 1960. Since then many different types of laser
have been developed, operating over a wide range of energies and wavelengths
covering the full spectrum from infrared to ultraviolet. Some lasers
Section 16.7 Lasers 253
FIGURE 16.10 Argon-ion laser. From www.nationallaser.com
produce short-duration highly intense light pulses, others operate in a continuous
mode. Lasers are now widely used in science, technology and increasingly
also in medicine. Figure 16.10 shows an Argon-ion laser that emits
green or blue light (depending on settings) and is one of the lasers frequently
used in medical applications.
16.7.1 Lasers Surgery
It was evident soon after the development of the first laser that the device
would be very useful as a surgical tool. Intense laser light focused onto a small
area could burn off and vaporize selected tissue without damage to neighboring
areas. Bleeding and pain during such a procedure would be minimized
because blood vessels are cauterized and nerve endings are sealed. Infections
would likewise be reduced because the cutting tool is not in physical contact
with the tissue.
Before lasers could be successfully used in surgical procedures, a wide
range of studies had to be conducted to understand the effect of intense light
on various types of tissue. Further, technology had to be developed for precise
control of light intensity and duration and for accurate positioning of the focal
point. While the surgical use of lasers is growing in many areas of medicine
and dentistry, the positional accuracy of laser tissue-removal is particularly
important in neurosurgery and eye surgery where a fraction of a millimeter
offset can make the difference between success and failure.
Ophthalmologists were among the first to use lasers for a wide range of
procedures. The repair of retinal detachments and retinal tears is one such
application. As a result of trauma or disease, the retina may detach from the
back of the eye or may develop tears. Left untreated this condition leads to a
loss of vision. Laser procedures have been very successful in arresting the retinal
degeneration and restoring normal vision. Laser light is focused through
254 Chapter 16 Atomic Physics
FIGURE 16.11 Physician performing laser eye surgery. From www.trustyguides.com
the iris onto the boundary of the detached or torn region of the retina. The
tissue is burnt and the subsequent scarring “welds” the retina to the underlying
tissue.
In another ophthalmological application, lasers are used to treat diabetic
retinopathy. Diabetes often causes disorders in blood circulation including
leaks in the retinal blood vessels. Such a condition can cause serious damage
to the retina and the optic nerve. Laser light focused on the damaged blood
vessel seals the leak and halts further retinal deterioration. Unfortunately, the
course of the disease is not halted and new leaks develop that require repeated
treatments. Figure 16.11 shows a typical eye surgery setup.
A relatively recent but now widely used application of lasers in ophthalmology
is the LASIK technique (Laser-Assisted in Situ Keratomileusis). This
is a laser surgical procedure that reshapes the cornea with the aim of correcting
focusing problems associated with myopia, hyperopia and astigmatism. In
this procedure, the computer that controls the laser is first programmed for the
amount and location of the corneal tissue to be removed. Then using a cutting
instrument called a microkeratome a flap is cut in the front part of the cornea
Chapter 16 Exercises 255
and the flap is folded back. The mid-part of the cornea is reshaped by the
computer controlled laser pulses that deliver the correct amount of energy to
evaporate the corneal tissue at the set locations. As a result of this procedure
the need for eyeglasses is often eliminated.
EXERCISES
16-1. Explain the operation of a spectrometer and describe two possible uses
for this device.
16-2. Describe the process of X-ray computerized tomography. What information
does this process provide that ordinary X-ray images do not?
16-3. Describe the operation of a helium–neon laser. Include a description of
the method for obtaining the inverted population distribution.
16-4. Two laser commonly used in laser surgery are the CO2 laser and
the argon-ion laser. Describe the method for obtaining the inverted
population distribution in these two lasers.
Chapter 17
Nuclear Physics
17.1 The Nucleus
Although all the atoms of a given element have the same number of protons in
their nucleus, the number of neutrons may vary. Atoms with the same number
of protons but different number of neutrons are called isotopes. All the nuclei
of the oxygen atom, for example, contain 8 protons but the number of neutrons
in the nucleus may be 8, 9, or 10. These are the isotopes of oxygen. They are
designated as 16
8 O, 17
8 O, and 18
8 O. This is a general type of nuclear symbolism
in which the subscript to the chemical symbol of the element is the number of
protons in the nucleus and the superscript is the sum of the number of protons
and neutrons. The number of neutrons often determines the stability of the
nucleus.
The nuclei of most naturally occurring atoms are stable. They do not
change when left alone. There are, however, many unstable nuclei which
undergo transformations accompanied by the emission of energetic radiation.
It has been found that the emanations from these radioactive nuclei fall into
three categories: (1) alpha (α) particles, which are high-speed helium nuclei
consisting of two protons and two neutrons; (2) beta (β) particles, which are
very high-speed electrons; and (3) gamma (γ) rays, which are highly energetic
photons.
The radioactive nucleus of a given element does not emit all three radiations
simultaneously. Some nuclei emit alpha particles, others emit beta particles,
and the emission of gamma rays may accompany either event.
256
Section 17.2 Magnetic Resonance Imaging 257
Radioactivity is associated with the transmutation of the nucleus from one
element to another. Thus, for example, when radium emits an alpha particle,
the nucleus is transformed into the element radon. The details of the process
are discussed in most physics texts (see [16-2]).
The decay or transmutation of a given radioactive nucleus is a random
event. Some nuclei decay sooner; others decay later. If, however, we deal
with a large number of radioactive nuclei, it is possible, by using the laws of
probability, to predict accurately the decay rate for the aggregate. This decay
rate is characterized by the half-life, which is the time interval for half the
original nuclei to undergo transmutation.
There is a great variation in the half-life of radioactive elements. Some
decay very quickly and have a half-life of only a few microseconds or less.
Others decay slowly with a half-life of many thousands of years. Only the
very long-lived radioactive elements occur naturally in the Earth’s crust. One
of these, for example, is the uranium isotope 238
92U, which has a half-life of
4.51 � 109 years. The short-lived radioactive isotopes can be produced in
accelerators by bombarding certain stable elements with high-energy particles.
Naturally occurring phosphorus, for example, has 15 protons and 16 neutrons
in its nucleus (31
15P). The radioactive phosphorus isotope 32
15P with 17 neutrons
can be produced by bombarding sulfur with neutrons. The reaction is
32
16S + neutron → 32
15P + proton
This radioactive phosphorus has a half-life of 14.3 days. Radioactive isotopes
of other elements can be produced in a similar way. Many of these isotopes
have been very useful in biological and clinical work.
17.2 Magnetic Resonance Imaging
Images of the shapes of internal organs obtained with computerized X-ray
tomography are excellent. However, X-rays, do not provide information about
the internal structure of tissue. CT scans may therefore, miss changes in
tissue structure and pathological alteration inside internal organs. Magnetic
resonance imaging (MRI), introduced in the early 1980s, is the most recent
addition to medical imaging techniques. This technique utilizes the magnetic
propertiesofthenucleustoprovideimagesofinternalbodyorganswithdetailed
information about soft-tissue structure.
The imaging techniques we have discussed so far (X-ray and ultrasound)
are in principle relatively simple. They utilize reflected or transmitted energy
to visualize internal structures. Magnetic resonance imaging is more complex.
It utilizes the principles of nuclear magnetic resonance (NMR) developed in
258 Chapter 17 Nuclear Physics
the 1940s. A detailed description of MRI is beyond the scope of this text, but
the principles are relatively simple to explain. A discussion of MRI begins
with an introduction to the principles of nuclear magnetic resonance.
17.2.1 Nuclear Magnetic Resonance
Protons and neutrons which are the constituents of atomic nuclei possess the
quantum mechanical property of spin which has magnitude and direction. We
can imagine these particles as if they were small spinning tops. As a result
of spin, the nuclear particles act as small bar magnets. Inside the nucleus,
these small magnets associated with the nucleons (protons and neutrons) line
up so as to cancel each other’s magnetic fields. However, if the number of
nucleons is odd, the cancellation is not complete, and the nucleus possesses
a net magnetic moment. Therefore, nuclei with an odd number of nucleons
behave as tiny magnets. Hydrogen, which has a nucleus consisting of a single
proton, does, of course, have a nuclear magnetic moment. The human body is
made of mostly water and other hydrogen-containing molecules. Therefore,
MRI images of structures within the body can be most effectively produced
using the magnetic properties of the hydrogen nucleus. Our discussion will be
restricted to the nuclear magnetic properties of hydrogen.
Normally, the little nuclear magnets in bulk material are randomized in
space as is shown in Fig. 17.1a, and the material does not possess a net magnetic
moment (M 0). The nuclear magnets are represented as small arrows.
However, the situation is altered in the presence of an external magnetic field.
When an external magnetic field is applied to a material possessing nuclear
magnetic moments, the tiny nuclear magnets line up either parallel or antiparallel
with the magnetic field as shown in Fig. 17.1b. The direction of the external
magnetic field is usually designated as the z-axis. As shown in the figure,
the x-y plane is orthogonal to the z-axis. Because the nuclear magnets parallel
to the field (+z) have a somewhat lower energy than those that are antiparallel
(−z), more of the nuclei are in the parallel state than in the antiparallel state. In
an external magnetic field, the assembly of parallel/antiparallel nuclear spins
as a whole has a net magnetic moment M that behaves as a magnet pointing
in the direction of the magnetic field.
The energy spacing Em between the parallel and antiparallel alignments
is
Em
γhB
2π
(17.1)
Here B is the externally applied magnetic field, h is the Planck constant as
defined earlier, and γ is called gyromagnetic ratio which is a property of a
given nucleus. Typically the strength of magnetic fields used in MRI is about
Section 17.2 Magnetic Resonance Imaging 259
FIGURE 17.1 (a) In the absence of an external magnetic field, nuclear spins are
randomized. (b) When an external magnetic field is applied to a material possessing
nuclear magnetic moments, the tiny nuclear magnets line up either parallel or antiparallel
with the magnetic field. The parallel configuration is at a lower energy.
2 tesla (T). (By comparison, the strength of the magnetic field of the Earth is
on the order of 10−4 T.)
The discrete energy spacing Em, between the two states shown in
Fig. 17.1b, makes this a resonant system. The frequency corresponding to
the energy difference between the two states is called the Larmor frequency
and in accord with Eq. 16.1 is given by
fL
Em
h
γB
2π
(17.2)
The gyromagnetic ratio γ for a proton is 2.68 � 108 T−1 sec−1. Magnetic
fields used in MRI are typically in the range 1 to 4 T. The corresponding
Larmor frequencies are about 43 to 170 MHz. These frequencies are in the
260 Chapter 17 Nuclear Physics
radio frequency (RF) range, which are much lower than X-rays and do not
disrupt living tissue.
If by some means the magnetic moment is displaced from the field, as
shown in the Fig. 17.2, it will precess (rotate) around the field as a spinning
top precesses in the gravitational field of the Earth. The frequency of precession
is the Larmor frequency given by Eq. 17.2. The displacement of the
magnetic moment is due to a reversal of alignment for some of the individual
nuclear magnetic moments from parallel to antiparallel alignment as shown
in Fig. 17.2. A displacement of 90◦ corresponds to equalizing the population
of the spin up and spin down states. To reverse the alignment of antiparallel
spins requires energy which must be supplied by an external source.
The energy required to displace the magnetic moment from the direction
of the external field is supplied by a short radio frequency driving pulse at
the Larmor frequency which is the natural (resonant) frequency of precession.
(This is analogous to setting a pendulum swinging by applying to it a force
FIGURE 17.2 A short radio frequency driving pulse at the Larmor frequency
displaces the magnetic moment from the external magnetic field by an angle determined
by the strength and duration of the driving pulse.
Section 17.2 Magnetic Resonance Imaging 261
at the frequency of the pendulum resonance.) The driving pulse is applied by
a coil surrounding the sample as shown in Fig. 17.2. At the end of the spinflipping
driving pulse, the magnetic moment is displaced from the external
magnetic field by an angle determined by the strength and duration of the
driving pulse.
The displaced magnetic moment produced by the radio frequency driving
pulse, precesses around the external magnetic field and itself generates a radio
frequency signal at the Larmor frequency of rotation. This emitted NMR signal
can be detected by a separate coil or by the driving coil itself. The detected
NMR signal decreases exponentially with time due to two distinct processes,
(1) the return of the nuclear spin orientations to the equilibrium distribution
and (2) variations in the local magnetic field.
Process 1: As was stated earlier, in the presence of an external field more of
the nuclei are lined up parallel to the field than antiparallel. The radio
frequency pulse flips some of the parallel spins into the antiparallel configuration.
As soon as the driving pulse is over, the nuclear spins and the
associated magnetic moment begin to return back to the original equilibrium
alignment. The equilibration is brought about by the exchange of
energy between the nuclear spins and the surrounding molecules. With
the return of the magnetic moment to the original alignment with the
external magnetic field, the precession angle decreases, as does the associated
NMR signal. The decay of the NMR signal is exponential with
time constant T1, called the spin lattice relaxation time.
Process 2: The local magnetic field throughout the object under examination
is not perfectly uniform. Variations in the magnetic field are produced
by the magnetic properties of molecules adjacent to the nuclear spins.
Such variations in the local magnetic field cause the Larmor frequency
of the individual nuclear magnetic moments to differ slightly from each
other. As a result, the precessions of the nuclei get out of phase with
each other, and the total NMR signal decreases. This dephasing is
likewise exponential, with a time constant T2, called the spin-spin relaxation
time.
The driving pulse and the emitted NMR signal are shown schematically
in Fig. 17.3. The NMR signal detected after the driving pulse contains information
about the material being studied. For a given initial driving pulse, the
magnitude of the emitted NMR signal is a function of the number of hydrogen
nuclei in the material. Bone, for example, which contains relatively few water
or other hydrogen-containing molecules, produces a relatively low NMR signal.
The post-pulse radiation emitted by fatty tissue is much higher.
262 Chapter 17 Nuclear Physics
FIGURE 17.3 The driving pulse and the emitted NMR signal.
The time constants T1 and T2 characterizing the rate of decay of the emitted
NMR signal provide information about the nature of the material within which
the precessing nuclei are located. The spinning top provides a useful analogy.
A well-designed top in vacuum will spin for a long time. In air, the duration
of the spin will be somewhat shorter because collisions with air molecules
will dissipate its rotational energy. In water, where the frictional losses are
yet greater, the top will spin hardly at all. The decay rate of the spinning
top provides information about the nature of the medium surrounding the top.
Similarly, the characteristic time constants T1 and T2 provide information about
the matter surrounding the precessing nuclei (see [16-4]). For example, with
an external magnetic field of 1 T, for fat T1 240 msec and T2 80 msec; for
heart tissue T1 570 msec and T2 57 msec (see [16-4]). Malignant tissue
is often characterized by higher values of T1.
The NMR principles described have been used since the 1940s to identify
molecules in various physics, chemistry, and biological applications. In
this application the detected NMR signal is derived from the entire volume
exposed to the magnetic field. The technique as discussed so far cannot provide
information about the location of the signal within the volume studied.
17.2.2 Imaging with NMR
In order to obtain a three-dimensional image using nuclear magnetic resonance,
we must isolate and identify the location of signals from small sections of the
Section 17.2 Magnetic Resonance Imaging 263
body and then build the image from these individual signals. In CT scans, such
tomographic spatial images are obtained by extracting the information from
intersection points of narrowly focused X-ray beams. This cannot be done
with NMR because the wavelengths of the radio frequency driving signals are
long, in the range of meters, which cannot be collimated into the narrow beams
required to examine small regions of interest.
In the 1970s, several new techniques were developed to utilize NMR
signals for the construction of two-dimensional tomographic images similar
to those provided by CT scans. One of the first of these was described by
P. C. Lauterbur in 1973. He demonstrated the principle using two tubes of
water, A and B as shown in Fig. 17.4. In a uniform magnetic field (B0) the
Larmor frequency of the two tubes is the same. Therefore, the post-pulse
NMR signals from tubes A and B cannot be distinguished. The NMR signals
from the two tubes can be made distinguishable by superimposing on the uniform
field B0 a magnetic field gradient B(x) as shown in Fig. 17.4b. The total
magnetic field now changes with position along the x-axis, and the associated
Larmor frequencies at the location of tubes A and B are now different.
As is evident, each point (actually small region x) on the x-axis is now
characterized by its unique Larmor frequency. Therefore, the NMR signal
FIGURE 17.4 (a) In a uniform magnetic field (B0) the Larmor frequency of two
locations in space A and B is the same. (b) When a magnetic field gradient is
superimposed on the uniform field, the Larmor frequencies at the locations A and B
are different.
264 Chapter 17 Nuclear Physics
observed after excitation with a pulse of a given frequency can be uniquely
associated with a specific region in the x-space. A field gradient in one direction
yields projection of the object onto that axis. To obtain a tomographic
image in the x-y plane, a field gradient in both the x- and y-directions must
be introduced (see Fig. 17.5). A magnetic field gradient is also applied in the
z-direction to select within the body the slice to be examined. A very large
number of NMR signals have to be collected and synthesized to construct an
MRI image. For this purpose, the intensity as well as the time constants T1
and T2 of the NMR signal are needed. The process is more complex than for
a CT scan and requires highly sophisticated computer programs.
A sketch of a whole-body MRI apparatus is shown in Fig. 17.5. Most such
devices use liquid helium cooled superconducting magnets to produce the high
FIGURE 17.5 Sketch of a whole body MRI imaging system.
Section 17.2 Magnetic Resonance Imaging 265
FIGURE 17.6 MRI image of brain. From V. Kuperman, “Magnetic Resonance
Imaging,” 2000, Academic Press.
magnetic fields required for the production of high resolution images. An MRI
image of the brain is shown in Fig. 17.6.
The MRI technique yields detailed visualization of soft tissue structures
with a resolution of about 0.5 mm. Such visualizations have been particularly
useful in neurology. All parts of the brain structure including arteries as thin as
a hair can be clearly seen deep inside the brain. However, conventional MRI
does not provide information about the functions performed by the brain. Such
a display in vivo of neural activity in the brain as it performs various tasks and
functions can be obtained with a modified MRI technique called functional
magnetic resonance imaging fMRI.
17.2.3 Functional Magnetic Resonance Imaging (fMRI)
Prior to the development of modern imaging techniques, information about
specific localized functions of the brain was obtained primarily from studies
(usually post mortem) of brain tumors and injuries. For example, in 1861
through a post mortem examination of an aphasic patient, a French physician,
Paul Pierre Broca, determined that the patient had a lesion in the left cerebral
hemisphere and confirmed earlier studies suggesting that this part of the
brain controlled speech production. The development of fMRI has made it
possible to observe non-invasively a wide range neural functions of interest in
psychology and clinical medicine.
When a specific region of the brain is activated, the energy requirement of
that region rises. Oxygenated blood flow to that part of the brain increases to
ensure an adequate supply of oxygen required to meet the increased energy
requirements. The fMRI technique makes use of the fact that oxygenated
hemoglobindoesnothaveamagneticmomentwhiledeoxygenatedhemoglobin
266 Chapter 17 Nuclear Physics
does. In the presence of deoxygenated hemoglobin with its magnetic moment,
thedephasingofthehydrogenNMRsignalismorerapidandthesignalintensity
is weaker than with the hemoglobin oxygenated. Therefore, regions of greater
brain activity, infused with more oxygenated blood, will produce a more
intense T2-weighted NMR signals. In this way regions of increased brain
activity can be clearly identified.
FMRI has been applied to identify regions of brain activity in the performance
of a wide range of cognitive, motor and sensory activities. Most of the
fMRI applications to date have been research-related. For example, in a 2003
study Eisenberg and colleagues set up a situation where the subjects experienced
social exclusion. Their fMRI studies showed that in such a situation the
same part of the brain is activated as is during physical pain.
The clinical applications of fMRI are still in their infancy, but there is little
doubt that within a few years fMRI will have a major role in medicine. Research
is being conducted to use fMRI as an early diagnostic tool to detect neurological
diseases such as Alzheimer’s, Parkinson’s and Huntington’s diseases.
Emerging results indicate that fMRI can provide important information to make
neurosurgical procedures such as removal of tumors more accurate. Pain management
and more accurate functional testing of psychoactive drugs are some
of the other areas where fMRI is likely to become a useful tool.
17.3 Radiation Therapy
The photons of X-rays and gamma-rays and the particles emitted by radioactive
nuclei all have energies far greater than the energies that bind electrons to
atoms and molecules. As a result, when such radiation penetrates into biological
materials, it can rip off electrons from the biological molecules and produce
substantial alterations in their structure. The ionized molecule may break
up, or it may chemically combine with another molecule to form an undesirable
complex. If the damaged molecule is an important component of a cell,
the whole cell may die. Water molecules in the tissue are also broken up by
the radiation into reactive fragments (H + OH). These fragments combine with
the biological molecules and alter them in a detrimental way. In addition, radiation
passing through tissue may simply give up its energy and heat the tissue
to a dangerously high temperature. A large dose of radiation may damage so
many cells that the whole organism dies. Smaller but still dangerous doses may
produce irreversible changes such as mutations, sterility, and cancer.
In controlled doses, however, radiation can be used therapeutically. In the
treatment of certain types of cancer, an ampul containing radioactive material
such as radium or cobalt 60 is implanted near the cancerous growth. By
careful placement of the radioactive material and by controlling the dose, the
hope is to destroy the cancer without greatly damaging the healthy tissue.
Section 17.4 Food Preservation by Radiation 267
Unfortunately some damage to healthy tissue is unavoidable. As a result, this
treatment is often accompanied by the symptoms of radiation sickness (diarrhea,
nausea, loss of hair, loss of appetite, and so on). If long-lived isotopes
are used in the therapy, the material must be removed after a prescribed period.
Short-lived isotopes, such as gold 198 with a half-life of about 3 days, decay
quickly enough so that they do not need to be removed after treatment.
Certain elements introduced into the body by injection or by mouth tend to
concentrate in specific organs. This phenomenon is used to advantage in radiation
therapy. The radioactive isotope phosphorus 32 (half-life, 14.3 days)
mentioned earlier accumulates in the bone marrow. Iodine 131 (half-life,
8 days) accumulates in the thyroid and is given for the treatment of hyper-
thyroidism.
An externally applied beam of gamma rays or X-rays can also be used to
destroy cancerous tumors. The advantage here is that the treatment is administered
without surgery. The effect of radiation on the healthy tissue can be
reduced by frequently altering the direction of the beam passing through the
body. The tumor is always in the path of the beam, but the dosage received by
a given section of healthy tissue is reduced.
17.4 Food Preservation by Radiation
Without some attempt at preservation, all foods decay rather quickly. Within
days and often within hours, many foods spoil to a point where they become
inedible. The decay is usually caused by microorganisms and enzymes that
decompose the organic molecules of the food.
Over the years, a number of techniques have been developed to retard
spoilage. Keeping the food in a cold environment reduces the rate of activity
for both the enzymes and the microorganisms. Dehydration of food achieves
the same goal. Heating the food for a certain period of time destroys many
microorganisms and again retards decay. This is the principle of pasteurization.
These methods of retarding spoilage are all at least 100 years old. There
is now a new technique of preserving food by irradiation.
High-energy radiation passing through the food destroys microorganisms
that cause decay. Radiation is also effective in destroying small insects that
attack stored foods. This is especially important for wheat and other grains
which at present are often fumigated before shipping or storage. Chemical
fumigation kills the insects but not their eggs. When the eggs hatch, the new
insects may destroy a considerable fraction of the grain. Radiation kills both
the insects and the eggs.
Gamma rays are used most frequently in food preservation. They have a
great penetrating power and are produced by relatively inexpensive isotopes
268 Chapter 17 Nuclear Physics
such as cobalt 60 and cesium 137. High-speed electrons produced by
accelerators have also been used to sterilize food. Electrons do not have the
penetrating power of gamma rays, but they can be aimed better and can be
turned off when not in use.
In the United States and in many other countries, there are now a number
of facilities for irradiating food. In the usual arrangement, the food on a
conveyor passes by the radioactive source, where it receives a controlled dose
of radiation. The source must be carefully shielded to protect the operator.
This problem is relatively simple to solve, and at present the technical problems
seem to be well in hand. One plant for irradiating food, in Gloucester,
Massachusetts, initially built by the Atomic Energy Commission, has been
operating successfully since 1964. It can process 1000 lb of fish per hour.
There is no doubt that irradiation retards spoilage of food. Irradiated strawberries,
for example, remain fresh for about 15 days after they have been
picked whereas strawberries that have not been treated begin to decay after
about 10 days. Irradiated unfrozen fish also lasts a week or two longer. Tests
have shown that the taste, nutritional value, and appearance of the food remain
acceptable. The important question is the safety of the procedure. Irradiation
at the levels used in the treatment does not make the food radioactive. There
is, however, the possibility that the changes induced by radiation may make
the food harmful. Over the past three decades, there have been many test programs
both with animals and with human volunteers to ascertain the safety of
food irradiation. At this point, the technique has been judged safe and is in
commercial use (see Exercise 10-3).
17.5 Isotopic Tracers
Most elements have isotopes differing from each other by the number of
neutrons in their nuclei. The isotopes of a given element are chemically
identical—that is, they participate in the same chemical reactions—but they
can be distinguished from each other because their nuclei are different. One
difference is, of course, in their mass. This property alone can be used to separate
one isotope from another. A mass spectrometer is one of the devices that
can perform this task. Another way to distinguish isotopes is by their radioactivity.
Many elements have isotopes that are radioactive. These isotopes are
easily identified by their activity. In either case, isotopes can be used to trace
the various steps in chemical reactions and in metabolic processes. Tracer
techniques have been useful also in the clinical diagnoses of certain disorders.
Basically the technique consists of introducing a rare isotope into the process
and then following the course of the isotope with appropriate detection
techniques. We will illustrate this technique with a few examples. Nitrogen
Section 17.6 Laws of Physics and Life 269
is one of the atoms in the amino acids that compose the protein molecules.
In nature, nitrogen is composed primarily of the isotope 14N. Only 0.36% of
natural nitrogen is in the form of the nonradioactive isotope 15N. Ordinarily
the amino acids reflect the natural composition of nitrogen.
It is possible to synthesize amino acids in a laboratory. If the synthesis is
done with pure 15N, the amino acids are distinctly marked. The amino acid
glycine produced in this way is introduced into the body of a subject where
it is incorporated into the hemoglobin of the blood. Periodic sampling of the
blood measures the number of blood cells containing the originally introduced
glycine. Such experiments have shown that the average lifetime of a red blood
cell is about four months.
Radioactive isotopes can be traced more easily and in smaller quantities
than the isotopes that are not radioactive. Therefore, in reactions with
elements that have radioactive isotopes, radioactive tracer techniques are preferred.
Since the 1950s, when radioactive isotopes first became widely available,
hundreds of important experiments have been conducted in this field.
An example of this technique is the use of radioactive phosphorus in the
study of nucleic acids. The element phosphorus is an important component
of the nucleic acids DNA and RNA. Naturally occurring phosphorus is all in
the form 31P, and, of course, this is the isotope normally found in the nucleic
acids. However, as discussed earlier, by bombarding sulphur 32 with neutrons,
it is possible to produce the radioactive phosphorus 32P which has a half-life
of 14.3 days. If the 32P isotope is introduced into the cell, the nucleic acids
synthesized in the cell incorporate this isotope into their structure. The nucleic
acids are then removed from the cell and their radioactivity is measured. From
these measurements it is possible to calculate the rate at which nucleic acids
are manufactured by the cell. These measurements, among others, provided
evidence for the roles of DNA and RNA in cell functions.
Radioactive tracers have been useful also in clinical measurements. In
one technique, the radioactive isotope of chromium is used to detect internal
hemorrhage. This isotope is taken up by the blood cells, which then become
radioactive. The radioactivity is, of course, kept well below the danger level.
If the circulation is normal, the radioactivity is distributed uniformly throughout
the body. A pronounced increase in radioactivity in some region indicates
a hemorrhage at that point.
17.6 Laws of Physics and Life
We have discussed in this book many phenomena in the life sciences that
are clearly explained by the theories of physics. Now we come to the most
fundamental question: Can physics explain life itself? In other words, if we
270 Chapter 17 Nuclear Physics
put together the necessary combination of atoms, at each step following the
known laws of physics, do we inevitably end up with a living organism, or
must we invoke some new principles outside the realm of current physics in
order to explain the occurrence of life? This is a very old question which still
cannot be answered with certainty. But it can be clarified.
Quantum mechanics, which is the fundamental theory of modern atomic
physics, has been very successful in describing the properties of atoms and
the interaction of atoms with each other. Starting with a single proton and
one electron, the theory shows that their interaction leads to the hydrogen
atom with its unique configuration and properties. The quantum mechanical
calculations for larger atoms are more complicated. In fact, so far a complete
calculation has been performed only for the hydrogen atom. The properties
of heavier atoms must be computed using various approximation techniques.
Yet there is little doubt that quantum mechanics describes all the properties of
atoms from the lightest to the heaviest. The experimental evidence gathered
over the past 100 years fully confirms this view.
The interactions between atoms, which result in the formation of molecules,
are likewise in the domain of quantum mechanics. Here again exact solutions
of the quantum mechanical equations have been obtained only for the simplest
molecule, H2. Still it is evident that all the rules for both organic and
inorganic chemistry follow from the principles of quantum mechanics. Even
though our present numerical techniques cannot cope with the enormous calculations
required to predict the exact configuration of a complex molecule, the
concepts developed in physics and chemistry are applicable. The strengths of
the interatomic bonds and the orientations of the atoms within the molecules
are all in accord with the theory. This is true even for the largest organic
molecules such as the proteins and DNA.
Past this point, however, we encounter a new level of organization: the cell.
The organic molecules, which are in themselves highly complex, combine to
form cells, which in turn are combined to form larger living organisms, which
possess all the amazing properties of life. These organisms take nourishment
from the environment, grow, reproduce, and at some level begin to govern
their own actions. Here it is no longer obvious that the theories governing the
interaction of atoms lead directly to these functions that characterize life. We
are now in the realm of speculations.
The phenomena associated with life show such remarkable organization
and planning that we may be tempted to suggest that perhaps some new undiscovered
law governs the behavior of organic molecules that come together to
form life. Yet there is no evidence for any special laws operating within living
systems. So far, on all levels of examination, the observed phenomena associated
with life obey the well-known laws of physics. This does not mean that
the existence of life follows from the basic principles of physics, but it may. In
fact the large organic molecules inside cells are sufficiently complex to contain
Chapter 17 Exercises 271
within their structures the information necessary to guide in a predetermined
way the activities associated with life. Some of these codes contained in the
specific groupings of atoms within the molecules have now been unraveled.
Because of these specific structures, a given molecule always participates in
a well-defined activity within the cell. It is very likely that all the complex
functions of cells and of cell aggregates are simply the collective result of
the enormously large number of predetermined but basically well-understood
chemical reactions.
This still leaves the most important question unanswered: What are the
forces and the principles that initially cause the atoms to assemble into coded
molecules which then ultimately lead to life. The answer here is probably
again within the scope of our existing theories of matter.
In 1951, S. L. Miller simulated in his laboratory the type of conditions that
may have existed perhaps 3.5 billion years ago in the atmosphere of the primordial
Earth. He circulated a mixture of water, methane, ammonia, and hydrogen
through an electric discharge. The discharge simulated the energy sources that
were then available from the sun, lightning, and radioactivity. After about one
week Miller found that the chemical activities in the mixture produced organic
molecules including some of the simple amino acids, which are the building
blocks of proteins. Since then, hundreds of other organic molecules have been
synthesized under similar conditions. Many of them resemble the components
of the important large molecules found in cells. It is thus plausible that in the
primordial oceans, rich in organic molecules produced by the prevailing chemical
reactions, life began. A number of smaller organic molecules combined
accidentally to form a large self-replicating molecule such as DNA. These, in
turn, combined into organized aggregates and finally into living cells.
Although the probability for the spontaneous occurrence of such events is
small, the time span of evolution is probably long enough to make this scenario
plausible. If that is indeed the case, the current laws of physics can explain all
of life. At the present state of knowledge about life processes, the completeness
of the descriptions provided by physics cannot be proved. The principles of
physics have certainly explained many phenomena, but mysteries remain. At
present, however, there seems to be no need to invoke any new laws.
EXERCISES
17-1. Describe the basic principles of magnetic resonance imaging.
17-2. What is your (considered) opinion of food preservation by radiation?
17-3. Through a literature search describe the most recent use of fMRI.
17-4. Discuss some of the most notable attributes of living systems that distinguish
them from inanimate ones.
Appendix A
Basic Concepts in Mechanics
In this section, we will define some of the fundamental concepts in mechanics.
We assume that the reader is familiar with these concepts and that here a
simple summary will be sufficient. A detailed discussion can be found in
basic physics texts, some of which are listed in the Bibliography.
A.1 Speed and Velocity
Velocity is defined as the rate of change of position with respect to time. Both
magnitude and direction are necessary to specify velocity. Velocity is, therefore,
a vector quantity. The magnitude of the velocity is called speed. In the
special case when the velocity of an object is constant, the distance s traversed
in time t is given by
s vt (A.1)
In this case, velocity can be expressed as
v
s
t
(A.2)
If the velocity changes along the path, the expression s/t yields the average
velocity.
272
Section A.2 Acceleration 273
A.2 Acceleration
If the velocity of an object along its path changes from point to point, its
motion is said to be accelerated (or decelerated). Acceleration is defined
as the rate of change in velocity with respect to time. In the special case of
uniform acceleration, the final velocity v of an object that has been accelerated
for a time t is
v v0 + at (A.3)
Here v0 is the initial velocity of the object, and a is the acceleration.1 Acceleration
can, therefore, be expressed as
a
v − v0
t
(A.4)
In the case of uniform acceleration, a number of useful relations can be
simply derived. The average velocity during the interval t is
vav
v + v0
2
(A.5)
The distance traversed during this time is
s vavt (A.6)
Using Eqs. A.4 and A.5, we obtain
s v0t +
at2
2
(A.7)
By substituting t (v − v0)/a (from Eq. A.4) into Eq. A.7, we obtain
v2
v2
0 + 2as (A.8)
1
Both velocity and acceleration may vary along the path. In general, velocity is defined as the
time derivative of the distance along the path of the object; that is,
v lim
t → 0
s
t
ds
dt
Acceleration is defined as the time derivative of the velocity along the path; that is,
a
dv
dt
d
dt
ds
dt
d2
s
dt2
274 Appendix A Basic Concepts in Mechanics
A.3 Force
Force is a push or a pull exerted on a body which tends to change the state of
motion of the body.
A.4 Pressure
Pressure is the force applied to a unit area.
A.5 Mass
We have stated that a force applied to a body tends to change its state of
motion. All bodies have the property of resisting change in their motion. Mass
is a quantitative measure of inertia or the resistance to a change in motion.
A.6 Weight
Every mass exerts an attractive force on every other mass; this attraction is
called the gravitational force. The weight of a body is the force exerted on the
body by the mass of the Earth. The weight of a body is directly proportional
to its mass. Weight being a force is a vector, and it points vertically down in
the direction of a suspended plumb line.
Mass and weight are related but distinct properties of an object. If a body
were isolated from all other bodies, it would have no weight, but it would still
have mass.
A.7 Linear Momentum
Linear momentum of a body is the product of its mass and velocity; that is,
Linear momentum mv (A.9)
A.8 Newton’s Laws of Motion
The foundations of mechanics are Newton’s three laws of motion. The laws
are based on observation, and they cannot be derived from more basic principles.
These laws can be stated as follows:
First Law: A body remains at rest or in a state of uniform motion in a straight
line unless it is acted on by an applied force.
Section A.9 Conservation of Linear Momentum 275
Second Law: The time rate of change of the linear momentum of a body is
equal to the force F applied to it.
Except at very high velocities, where relativistic effects must be considered,
the second law can be expressed mathematically in terms of the
mass m and acceleration a of the object as2
F ma (A.10)
This is one of the most commonly used equations in mechanics. It
shows that if the applied force and the mass of the object are known, the
acceleration can be calculated. When the acceleration is known, the
velocity of the object and the distance traveled can be computed from
the previously given equations.
The Earth’s gravitational force, like all other forces, causes an acceleration.
By observing the motion of freely falling bodies, this acceleration
has been measured. Near the surface of the Earth, it is approximately
9.8 m/sec2. Because gravitational acceleration is frequently used
in computations, it has been given a special symbol g. Therefore, the
gravitational force on an object with mass m is
Fgravity mg (A.11)
This is, of course, also the weight of the object.
Third Law: For every action, there is an equal and opposite reaction. This
law implies that when two bodies A and B interact so that A exerts a
force on B, a force of the same magnitude but opposite in direction is
exerted by B on A. A number of illustrations of the third law are given
in the text.
A.9 Conservation of Linear Momentum
It follows from Newton’s laws that the total linear momentum of a system of
objects remains unchanged unless acted on by an outside force.
2
The second law can be expressed mathematically in terms of the time derivative of
momentum: that is,
Force t → 0
mv(t + t) − mv(t)
t
d
dt
(mv) m
dv
dt
ma
276 Appendix A Basic Concepts in Mechanics
FIGURE A.1 The radian.
A.10 Radian
In the analysis of rotational motion, it is convenient to measure angles in a
unit called a radian. With reference to Fig. A.1, the angle in radian units is
defined as
θ
s
r
(A.12)
where s is the length of the circular arc and r is the radius of rotation. In a full
circle, the arc length is the circumference 2πr. Therefore in radian units the
angle in a full circle is
θ
2πr
r
2π rad
Hence,
1 rad
360◦
2π
57.3◦
A.11 Angular Velocity
The angular velocity ω is the angular displacement per unit time; that is, if a
body rotates through an angle θ (in radians) in a time t, the angular velocity is
ω
θ
t
(rad/sec) (A.13)
Section A.14 Equations for Angular Momentum 277
A.12 Angular Acceleration
Angular acceleration α is the time rate of change of angular velocity. If the
initial angular velocity is ω0 and the final angular velocity after a time t is ωf,
the angular acceleration is3
α
ωf − ω0
t
(A.14)
A.13 Relations between Angular and Linear Motion
As an object rotates about an axis, each point in the object travels along the
circumference of a circle; therefore, each point is also in linear motion. The
linear distance s traversed in angular motion is
s rθ
The linear velocity v of a point that is rotating at an angular velocity ω a
distance r from the center of rotation is
v rω (A.15)
The direction of the vector v is at all points tangential to the path s. The linear
acceleration along the path s is
a rα (A.16)
A.14 Equations for Angular Momentum
The equations for angular motion are analogous to the equations for translational
motion. For a body moving with a constant angular acceleration α and
initial angular velocity ω0, the relationships are shown in Table A.1.
3
Both angular velocity and angular acceleration may vary along the path. In general, the
instantaneous angular velocity and acceleration are defined as
ω
dθ
dt
; α
dω
dt
d2
θ
dt2
278 Appendix A Basic Concepts in Mechanics
TABLE A.1 Equations for
Rotational Motion (angular
acceleration, α constant)
ω ω0 + αt
θ ω0t + 1
2
αt2
ω2
ω2
0 + 2αθ
ωav
(ω0 + ω)
2
A.15 Centripetal Acceleration
As an object rotates uniformly around an axis, the magnitude of the linear
velocity remains constant, but the direction of the linear velocity is continuously
changing. The change in velocity always points toward the center
of rotation. Therefore, a rotating body is accelerated toward the center of
rotation. This acceleration is called centripetal (center-seeking) acceleration.
The magnitude of the centripetal acceleration is given by
ac
v2
r
ω2
r (A.17)
where r is the radius of rotation and v is the speed tangential to the path of
rotation. Because the body is accelerated toward its center of rotation, we
conclude from Newton’s second law that a force pointing toward the center of
rotation must act on the body. This force, called the centripetal force Fc, is
given by
Fc mac
mv2
r
mω2
r (A.18)
where m is the mass of the rotating body.
For a body to move along a curved path, a centripetal force must be applied
to it. In the absence of such a force, the body moves in a straight line, as
required by Newton’s first law. Consider, for example, an object twirled at
the end of a rope. The centripetal force is applied by the rope on the object.
From Newton’s third law, an equal but opposite reaction force is applied on
the rope by the object. The reaction to the centripetal force is called the centrifugal
force. This force is in the direction away from the center of rotation.
The centripetal force, which is required to keep the body in rotation, always
acts perpendicular to the direction of motion and, therefore, does no work
Section A.17 Torque 279
TABLE A.2 Moments of Inertia of Some Simple Bodies
Body Location of axis Moment of inertia
A thin rod of length l Through the center ml2
/12
A thin rod of length l Through one end ml2
/3
Sphere of radius r Along a diameter 2mr2
/5
Cylinder of radius r Along axis of symmetry mr2
/2
(see Eq. A.28). In the absence of friction, energy is not required to keep a
body rotating at a constant angular velocity.
A.16 Moment of Inertia
The moment of inertia in angular motion is analogous to mass in translational
motion. The moment of inertia I of an element of mass m located a distance r
from the center of rotation is
I mr2
(A.19)
In general, when an object is in angular motion, the mass elements in the
body are located at different distances from the center of rotation. The total
moment of inertia is the sum of the moments of inertia of the mass elements
in the body.
Unlike mass, which is a constant for a given body, the moment of inertia
depends on the location of the center of rotation. In general, the moment of
inertia is calculated by using integral calculus. The moments of inertia for a
few objects useful for our calculations are shown in Table A.2.
A.17 Torque
Torque is defined as the tendency of a force to produce rotation about an axis.
Torque, which is usually designated by the letter L, is given by the product of
the perpendicular force and the distance d from the point of application to the
axis of rotation; that is (see Fig. A.2),
L F cos θ � d (A.20)
The distance d is called the lever arm or moment arm.
280 Appendix A Basic Concepts in Mechanics
FIGURE A.2 Torque produced by a force.
A.18 Newton’s Laws of Angular Motion
The laws governing angular motion are analogous to the laws of translational
motion. Torque is analogous to force, and the moment of inertia is analogous
to mass.
First Law: A body in rotation will continue its rotation with a constant angular
velocity unless acted upon by an external torque.
Second Law: The mathematical expression of the second law in angular
motion is analogous to Eq. A.10. It states that the torque is equal to the
product of the moment of inertia and the angular acceleration; that is,
L Iα (A.21)
Third Law: For every torque, there is an equal and opposite reaction torque.
A.19 Angular Momentum
Angular momentum is defined as
Angular momentum Iω (A.22)
From Newton’s laws, it can be shown that angular momentum of a body is
conserved if there is no unbalanced external torque acting on the body.
Section A.20 Addition of Forces and Torques 281
FIGURE A.3 The resolution of a force into its vertical and horizontal components.
A.20 Addition of Forces and Torques
Any number of forces and torques can be applied simultaneously to a given
object. Because forces and torques are vectors, characterized by both a magnitude
and a direction, their net effect on a body is obtained by vectorial addition.
When it is required to obtain the total force acting on a body, it is often convenient
to break up each force into mutually perpendicular components. This
is illustrated for the two-dimensional case in Fig. A.3. Here we have chosen
the horizontal x- and the vertical y-directions as the mutually perpendicular
axes. In a more general three-dimensional case, a third axis is required for the
analysis.
The two perpendicular components of the force F are
Fx F cos θ
Fy F sin θ
(A.23)
The magnitude of the force F is given by
F F2
x + F2
y (A.24)
When adding a number of forces (F1, F2, F3, . . .) the mutually perpendicular
components of the total force FT are obtained by adding the corresponding
282 Appendix A Basic Concepts in Mechanics
components of each force; that is,
(FT)x (F1)x + (F2)x + (F3)x + � � �
(FT)y (F1)y + (F2)y + (F3)y + � � �
(A.25)
The magnitude of the total force is
FT (FT)2
x + (FT)2
y (A.26)
Thetorqueproducedbyaforceactstoproducearotationineitheraclockwise
or a counterclockwise direction. If we designate one direction of rotation as
positive and the other as negative, the total torque acting on a body is obtained
by the addition of the individual torques each with the appropriate sign.
A.21 Static Equilibrium
A body is in static equilibrium if both its linear and angular acceleration are
zero. To satisfy this condition, the sum of the forces F acting on the body,
as well as the sum of the torques L produced by these forces must be zero;
that is,
P
F 0 and
P
L 0 (A.27)
A.22 Work
In our everyday language, the word work denotes any types of effort whether
physicalormental. Inphysics,amorerigorousdefinitionisrequired. Herework
is defined as the product of force and the distance through which the force acts.
Only the force parallel to the direction of motion does work on the object. This
is illustrated in Fig. A.4. A force F applied at an angle θ pulls the object along
the surface through a distance D. The work done by the force is
Work F cos θ � D (A.28)
A.23 Energy
Energy is an important concept. We find reference to energy in connection
with widely different phenomena. We speak of atomic energy, heat energy,
potential energy, solar energy, chemical energy, kinetic energy; we even speak
Section A.24 Forms of Energy 283
FIGURE A.4 Work done by a force.
of people as being full of energy. The common factor that ties together these
manifestations is the possibility of obtaining work from these sources. The
connection between energy and work is simple: Energy is required to do
work. Energy is measured in the same units as work; in fact, there is a oneto-one
correspondence between them. It takes 2 J of energy to do 2 J of work.
In all physical processes, energy is conserved. Through work, one form of
energy can be converted into another, but the total amount of energy remains
unchanged.
A.24 Forms of Energy
A.24.1 Kinetic Energy
Objects in motion can do work by virtue of their motion. For example, when
a moving object hits a stationary object, the stationary object is accelerated.
This implies that the moving object applied a force on the stationary object
and performed work on it. The kinetic energy (KE) of a body with mass m
moving with a velocity v is
KE
1
2
mv2
(A.29)
In rotational motion, the kinetic energy is
KE
1
2
Iω2
(A.30)
A.24.2 Potential Energy
Potential energy of a body is the ability of the body to do work because of its
position or configuration. A body of weight W raised to a height H with respect
284 Appendix A Basic Concepts in Mechanics
to a surface has a potential energy (PE)
PE WH (A.31)
This is the amount of work that had to be performed to raise the body to height
H. The same amount of energy can be retrieved by lowering the body back
to the surface.
A stretched or compressed spring possesses potential energy. The force
required to stretch or compress a spring is directly proportional to the length
of the stretch or compression (s); that is,
F ks (A.32)
Here k is the spring constant. The potential energy stored in the stretched or
compressed spring is
PE
1
2
ks2
(A.33)
A.24.3 Heat
Heat is a form of energy, and as such it can be converted to work and other
forms of energy. Heat, however, is not equal in rank with other forms of
energy. While work and other forms of energy can be completely converted
to heat, heat energy can only be converted partially to other forms of energy.
This property of heat has far-reaching consequences which are discussed in
Chapter 10.
Heat is measured in calorie units. One calorie (cal) is the amount of heat
required to raise the temperature of 1 g of water by 1 C◦. The heat energy
required to raise the temperature of a unit mass of a substance by 1 degree is
called the specific heat. One calorie is equal to 4.184 J.
A heat unit frequently used in chemistry and in food technology is the
kilocalorie or Cal which is equal to 1000 cal.
A.25 Power
The amount of work done—or energy expended—per unit time is called
power. The algebraic expression for power is
P
E
t
(A.34)
where E is the energy expended in a time interval t.
Section A.26 Units and Conversions 285
A.26 Units and Conversions
In our calculations we will mostly use SI units in which the basic units for
length, mass, and time are meter, kilogram, and second. However, other units
are also encountered in the text. Units and conversion factors for the most
commonly encountered quantities are listed here with their abbreviations.
A.26.1 Length
SI unit: meter (m)
Conversions: 1 m 100 cm (centimeter) 1000 mm (millimeter)
1000 m 1 km
1 m 3.28 feet 39.37 in
1 km 0.621 mile
1 in 2.54 cm
In addition, the micron and the angstrom are used frequently in physics and
biology.
1 micron (μm) 10−6
m 10−4
cm
1 angstrom ( ˚A)∗
10−8
cm
A.26.2 Mass
SI unit: kilogram (kg)
Conversions: 1 kg 1000 g
The weight of a 1-kg mass is 9.8 newton (N).
A.26.3 Force
SI Unit: kg m s−2, name of unit: newton (N)
Conversions: 1 N 105 dynes (dyn) 0.225 lbs
A.26.4 Pressure
SI unit: kg m−1 s−2, name of unit: pascal (Pa)
Conversions: 1 Pa 10−1 dynes/cm2 9.87 � 10−6 atmosphere (atm)
1.45 � 10−4 lb/in2
1 atm 1.01 � 105 Pa 760 mmHg (torr)
286 Appendix A Basic Concepts in Mechanics
A.26.5 Energy
SI unit: kg m−2 s−2, name of unit: joule (J)
Conversion: 1 J 1 N-m 107 ergs 0.239 cal 0.738 ft-lb
A.26.6 Power
SI unit: J s−1, name of unit: watt (W)
Conversion: 1 W 107 ergs/sec 1.34 � 10−3 horsepower (hp)
Appendix B
Review of Electricity
B.1 Electric Charge
Matter is composed of atoms. An atom consists of a nucleus surrounded by
electrons. The nucleus itself is composed of protons and neutrons. Electric
charge is a property of protons and electrons. There are two types of electric
charge: positive and negative. The proton is positively charged, and the electron
isnegativelycharged. Allelectricalphenomenaareduetotheseelectriccharges.
Charges exert forces on each other. Unlike charges attract and like charges
repel each other. The electrons are held around the nucleus by the electrical
attraction of the protons. Although the proton is about 2000 times heavier than
the electron, the magnitude of the charge on the two is the same. There are as
many positively charged protons in an atom as negatively charged electrons.
The atom as a whole is, therefore, electrically neutral. The identity of an atom
is determined by the number of protons in the nucleus. Thus, for example,
hydrogen has 1 proton; nitrogen has 7 protons; and gold has 79 protons.
Itispossibletoremoveelectronsfromanatom, makingitpositivelycharged.
Such an atom with missing electrons is called a positive ion. It is also possible
to add an electron to an atom which makes it a negative ion.
Electric charge is measured in coulombs (C). The magnitude of the charge
on the proton and the electron is 1.60 � 10−19 C. The force F between two
charged bodies is proportional to the product of their charges Q1 and Q2 and
is inversely proportional to the square of the distance R between them; that is,
F
KQ1Q2
R2
(B.1)
287
288 Appendix B Review of Electricity
This equation is known as Coulomb’s law. If R is measured in meters, the
constant K is 9 � 109, and F is obtained in newtons.
B.2 Electric Field
An electric charge exerts a force on another electric charge; a mass exerts a
force on another mass; and a magnet exerts a force on another magnet. All
these forces have an important common characteristic: Exertion of the force
does not require physical contact between the interacting bodies. The forces
act at a distance. The concept of lines of force or field lines is useful in visualizing
these forces which act at a distance.
Any object that exerts a force on another object without contact can be
thought of as having lines of force emanating from it. The complete line configuration
is called a force field. The lines point in the direction of the force,
and their density at any point in space is proportional to the magnitude of the
force at that point.
The lines of force emanate from an electric charge uniformly in all directions.
By convention, the lines point in the direction of the force that the source
charge exerts on a positive charge. Thus, the lines of force point away from a
positive source charge and into a negative source charge (see Fig. B.1). The
number of lines emanating from the charge is proportional to the magnitude
of the electric charge. If the size of the source charge is doubled, the number
of force lines is also doubled.
Lines of force need not be straight lines; as we mentioned, they point in
the direction in which the force is exerted. As an example, we can consider the
FIGURE B.1 Two-dimensional representation of the electric field produced by a
positive point charge (a) and a negative point charge (b).
Section B.4 Electric Current 289
FIGURE B.2 Lines of force produced by a positive and a negative charge separated
by a distance d.
net field due to two charges separated by a distance d. To determine this field
we must compute the direction and size of the net force on a positive charge
at all points in space. This is done by adding vectorially the force lines due
to each charge. The force field due to a positive and negative charge of equal
magnitude separated by a distance d from each other is shown in Fig. B.2.
Here the lines of force are curved. This is, of course, the direction of the net
force on a positive charge in the region surrounding the two fixed charges.
The field shown in Fig. B.2 is called a dipole field, and it is similar to the field
produced by a bar magnet.
B.3 Potential Difference or Voltage
The electric field is measured in units of volt per meter (or volt per centimeter).
The product of the electric field and the distance over which the field extends
is an important parameter which is called potential difference or voltage. The
voltage (V ) between two points is a measure of energy transfer as the charge
moves between the two points. Potential difference is measured in volts. If
there is a potential difference between two points, a force is exerted on a charge
placed in the region between these points. If the charge is positive, the force
tends to move it away from the positive point and toward the negative point.
B.4 Electric Current
An electric current is produced by a motion of charges. The magnitude of the
current depends on the amount of charge flowing past a given point in a given
period of time. Current is measured in amperes (A). One ampere is 1 coulomb
(C) of charge flowing past a point in 1 second (sec).
290 Appendix B Review of Electricity
B.5 Electric Circuits
The amount of current flowing between two points in a material is proportional
to the potential difference between the two points and to the electrical
properties of the material. The electrical properties are usually represented by
three parameters: resistance, capacitance, and inductance. Resistance measures
the opposition to current flow. This parameter depends on the property
of the material called resistivity, and it is analogous to friction in mechanical
motion. Capacitance measures the ability of the material to store electric
charges. Inductance measures the opposition in the material to changes in
current flow. All materials exhibit to some extent all three of these properties;
often, however, one of these properties is predominant. It is possible to
manufacture components with specific values of resistance, capacitance, or
inductance. These are called, respectively, resistors, capacitors, and inductors.
The schematic symbols for these three electrical components are shown in
Fig. B.3. Electrical components can be connected together to form an electric
circuit. Currents can be controlled by the appropriate choice of components
and interconnections in the circuit. An example of an electric circuit is shown
in Fig. B.4. Various techniques have been developed to analyze such circuits
and to calculate voltages and currents at all the points in the circuit.
B.5.1 Resistor
The resistor is a circuit component that opposes current flow. Resistance (R)
is measured in units of ohm ( ). The relation between current (I ) and
FIGURE B.3 Circuit components.
FIGURE B.4 Example of an electric circuit.
Section B.5 Electric Circuits 291
voltage (V ) is given by Ohm’s law, which is
V IR (B.2)
Materials that present a very small resistance to current flow are called conductors.
Materials with a very large resistance are called insulators. A flow
of current through a resistor is always accompanied by power dissipation as
electrical energy is converted to heat. The power (P) dissipated in a resistor is
given by
P I2
R (B.3)
The inverse of resistance is called conductance, which is usually designated by
the symbol G. Conductance is measured in units of mho, also called Siemens.
The relationship between conductance and resistance is
G
1
R
(B.4)
B.5.2 Capacitor
The capacitor is a circuit element that stores electric charges. In its simplest
form it consists of two conducting plates separated by an insulator (see
Fig. B.5). Capacitance (C) is measured in farads. The relation between the
stored charge (Q), and the voltage across the capacitor is given by
Q CV (B.5)
In a charged capacitor, positive charges are on one side of the plate, and
negative charges are on the other. The amount of energy (E) stored in such a
configuration is given by
E
1
2
CV2
(B.6)
FIGURE B.5 A simple capacitor.
292 Appendix B Review of Electricity
B.5.3 Inductor
The inductor is a device that opposes a change in the current flowing through
it. Inductance is measured in units called henry.
B.6 Voltage and Current Sources
Voltages and currents can be produced by various batteries and generators.
Batteries are based on chemical reactions that result in a separation of positive
and negative charges within a material. Generators produce a voltage by the
motion of conductors in magnetic fields. The circuit symbols for these sources
are shown in Fig. B.6.
B.7 Electricity and Magnetism
Electricity and magnetism are related phenomena. A changing electric field
always produces a magnetic field, and a changing magnetic field always produces
an electric field. All electromagnetic phenomena can be traced to this
basic interrelationship. A few of the consequences of this interaction follow:
1. An electric current always produces a magnetic field at a direction
perpendicular to the current flow.
2. A current is induced in a conductor that moves perpendicular to a
magnetic field.
3. An oscillating electric charge emits electromagnetic waves at the
frequency of oscillation. This radiation propagates away from the source
at the speed of light. Radio waves, light, and X-rays are examples of
electromagnetic radiation.
FIGURE B.6 Circuit symbols for a battery and a generator.
Appendix C
Review of Optics
C.1 Geometric Optics
The characteristics of optical components, such as mirrors and lenses, can be
completely derived from the wave properties of light. Such detailed calculations,
however, are usually rather complex because one has to keep track
of the wave front along every point on the optical component. It is possible
to simplify the problem if the optical components are much larger than the
wavelength of light. The simplification entails neglecting some of the wave
properties of light and considering light as a ray traveling perpendicular to the
wave front (Fig. C.1). In a homogeneous medium, the ray of light travels in a
straight line; it alters direction only at the interface between two media. This
simplified approach is called geometric optics.
The speed of light depends on the medium in which it propagates. In
vacuum, light travels at a speed of 3 � 108 m/sec. In a material medium, the
speed of light is always less. The speed of light in a material is characterized
by the index of refraction (n) defined as
n
c
v
(C.1)
where c is the speed of light in vacuum and v is the speed in the material.
When light enters from one medium into another, its direction of propagation
is changed (see Fig. C.2). This phenomenon is called refraction. The relationship
between the angle of incidence (θ1) and the angle of refraction (θ2)
293
294 Appendix C Review of Optics
FIGURE C.1 Light rays perpendicular to the wave front.
is given by
sin θ1
sin θ2
n2
n1
(C.2)
The relationship in Eq. C.2 is called Snell’s law. As shown in Fig. C.2, some
of the light is also reflected. The angle of reflection is always equal to the
angle of incidence.
In Fig. C.2a, the angle of incidence θ1 for the entering light is shown to
be greater than the angle of refraction θ2. This implies that n2 is greater than
n1 as would be the case for light entering from air into glass, for example
(see Eq. C.2). If, on the other hand, the light originates in the medium of
higher refractive index, as shown in Fig. C.2b, then the angle of incidence
θ1 is smaller than the angle of refraction θ2. At a specific value of angle θ1
called the critical angle (designated by the symbol θc), the light emerges tangent
to the surface, that is, θ2 90◦. At this point, sin θ2 1 and, therefore,
sin θ1 sin θc n2/n1. Beyond this angle, that is for θ1 > θc, light originating
in the medium of higher refractive index does not emerge from the
medium. At the interface, all the light is reflected back into the medium. This
phenomenon is called total internal reflection. For glass, n2 is typically 1.5,
and the critical angle at the glass-air interface is sin θc 1/1.5 or θc 42◦.
Transparent materials such as glass can be shaped into lenses to alter the
direction of light in a specific way. Lenses fall into two general categories:
converging lenses and diverging lenses. A converging lens alters the direction
of light so that the rays are brought together. A diverging lens has the opposite
effect; it spreads the light rays apart.
Using geometric optics, we can calculate the size and shape of images
formed by optical components, but we cannot predict the inevitable blurring
of images which occurs as a result of the wave nature of light.
Section C.2 Converging Lenses 295
FIGURE C.2 (Top) Reflection and refraction of light. (Bottom) Total internal
reflection.
C.2 Converging Lenses
A simple converging lens is shown in Fig. C.3. This type of a lens is called a
convex lens.
Parallel rays of light passing through a convex lens converge at a point
called the principal focus of the lens. The distance of this point from the lens
is called the focal length f. Conversely, light from a point source at the focal
point emerges from the lens as a parallel beam. The focal length of the lens is
296 Appendix C Review of Optics
FIGURE C.3 The convex lens illuminated (a) by parallel light, (b) by point source at
the focus.
determined by the index of refraction of the lens material and the curvature of
the lens surfaces. We adopt the following convention in discussing lenses.
1. Light travels from left to right.
2. The radius of curvature is positive if the curved surface encountered by
the light ray is convex; it is negative if the surface is concave.
It can be shown that for a thin lens the focal length is given by
1
f
(n − 1)
1
R1
−
1
R2
(C.3)
where R1 and R2 are the curvatures of the first and second surfaces, respectively
(Fig. C.4). In Fig. C.4, R2 is a negative number.
Focal length is a measure of the converging power of the lens. The shorter
the focal length, the more powerful the lens. The focusing power of a lens is
Section C.2 Converging Lenses 297
FIGURE C.4 Radius of curvature defined for a lens.
often expressed in diopters defined as
Focusing power
1
f (meters)
(diopters) (C.4)
If two thin lenses with focal lengths f1 and f2, respectively, are placed close
together, the focal length fT of the combination is
1
fT
1
f1
+
1
f2
(C.5)
Light from a point source located beyond the focal length of the lens is
converged to a point image on the other side of the lens (Fig. C.5a). This type
of an image is called a real image because it can be seen on a screen placed at
the point of convergence.
If the distance between the source of light and the lens is less than the focal
length, the rays do not converge. They appear to emanate from a point on the
source side of the lens. This apparent point of convergence is called a virtual
image (Fig. C.5b).
For a thin lens, the relationship between the source and the image distances
from the lens is given by
1
p
+
1
q
1
f
(C.6)
Here p and q, respectively, are the source and the image distances from the
lens. By convention, q in this equation is taken as positive if the image is
formed on the side of the lens opposite to the source and negative if the image
is formed on the source side.
Light rays from a source very far from the lens are nearly parallel; therefore,
by definition we would expect them to be focused at the principal focal
point of the lens. This is confirmed by Eq. C.6, which shows that as p becomes
very large (approaches infinity), q is equal to f.
298 Appendix C Review of Optics
FIGURE C.5 Image formation by a convex lens: (a) real image, (b) virtual image.
If the source is displaced a distance x from the axis, the image is formed
at a distance y from the axis such that
y
x
q
p
(C.7)
This is illustrated for a real image in Fig. C.6. The relationship between p and
q is still given by Eq. C.6.
C.3 Images of Extended Objects
So far we have discussed only the formation of images from point sources.
The treatment, however, is easily applied to objects of finite size.
When an object is illuminated, light rays emanate from every point on the
object (Fig. C.7a). Each point on the object plane a distance p from the lens
Section C.3 Images of Extended Objects 299
FIGURE C.6 Image formation off axis.
FIGURE C.7 Image of an object: (a) real, (b) virtual.
is imaged at the corresponding point on the image plane a distance q from the
lens. The relationship between the object and the image distances is given by
Eq. C.6. As shown in Fig. C.7, real images are inverted and virtual images are
upright. The ratio of image to object height is given by
Image height
Object height
−
q
p
(C.8)
300 Appendix C Review of Optics
FIGURE C.8 A diverging lens.
C.4 Diverging Lenses
An example of a diverging lens is the concave lens shown in Fig. C.8. Parallel
light diverges after passing through a concave lens. The apparent source of
origin for the diverging rays is the focal point of the concave lens. All the
equations we have presented for the converging lens apply in this case also,
provided the sign conventions are obeyed. From Eq. C.3, it follows that the
focal length for a diverging lens is always negative and the lens produces only
virtual images (Fig. C.8).
C.5 Lens Immersed in a Material Medium
The lens equations that we have presented so far apply in the case when the
lens is surrounded by air that has a refraction index of approximately 1. Let
us now consider the more general situation shown in Fig. C.9, which we will
need in our discussion of the eye. The lens here is embedded in a medium that
has a different index of refraction (n1 and n2) on each side of the lens. It can
be shown (see [15-3]) that under these conditions the relationship between the
object and the image distances is
n1
p
+
n2
q
nL − n1
R1
−
nL − n2
R2
(C.9)
Section C.5 Lens Immersed in a Material Medium 301
FIGURE C.9 Lens immersed in a material medium.
Here nL is the refraction index of the lens material. The effective focal
length in this case is
1
f
n2 − n1
R1
−
nL − n2
R2
(C.10)
Note that in air n1 n2 1 and Eq. C.10 reduces to Eq. C.3.
The lens equations we have presented in this appendix assume that the
lenses are thin. This is not a fully valid assumption for the lenses in the eye.
Nevertheless these equations are adequate for our purposes.
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McGraw-Hill Book Co., 1969.
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Answers to Numerical
Exercises
Chapter 1
1-1(b). F 254 N (57.8 lb)
1-3. θ 72.6◦
1-4. Maximum weight 335 N (75 lb)
1-5(a). Fm 2253 N (508 lb), Fr 2386 N (536 lb)
1-6. Fm 720 N, Fr 590 N
1-7(a). Fm 2160 N, Fr 1900 N
1-8. Fm 103 N, Fr 84 N
1-10. x 19.6 cm, v of tendon 4 cm/sec, v of weight 38 cm/sec
1-11. Fm 0.47 W, Fr 1.28 W
1-12(a). Fm 2000 N, Fr 2200 N; (b). Fm 3220 N, Fr 3490 N
1-13. FA 2.5 W, FT 3.5 W
Chapter 2
2-1(a). Distance 354 m; (b). Independ of mass
2-2(a). μ 0.067
2-3(a). μ 1.95; (b). with μ 1.0, θ 39.4◦, with μ 0.01, θ 0.6◦
Chapter 3
3-1. P 4120 watt
3-2. H 126 cm
310
Answers to Numerical Exercises 311
3-3. Fr 1.16 W, θ 65.8◦
3-4. T 0.534 sec
3-5(a). R 13.5 m; (b). H 3.39 m; (c). 4.08 sec
3-6. v 8.6 m/sec
3-7. r 1.13 m
3-8(a). v 8.3 m/sec; (b) 16.6 cm/sec
3-9. Energy expended/sec 1350 J/sec
3-10. P 371 watt
Chapter 4
4-2. F 10.1 N
4-3. ω 1.25 rad/sec; linear velocity 6.25 m/sec
4-4. ω 1.25 rad/sec 33.9 rpm
4-5. v 62.8 m/sec
4-6. Speed 1.13 m/sec 4.07 km/h 2.53 mph
4-7. T 1.6 sec
4-8. E 1.64 mv2
4-9. Fall time 1 sec
Chapter 5
5-1. v 2.39 m/sec (5.3 mph)
5-2. v 8 m/sec; with 1 cm2area v 2 m/sec
5-3. h 5.1 m
5-4. t 3 � 10−2 sec
5-5. v 17 m/sec (37 mph)
5-6. Force/cm2 4.6 � 106 dyn/cm2, yes
5-7. v 0.7 m/sec, no
Chapter 6
6-1. F 2 W
6-2. 0.052 mm
6-3. h 18.4 cm
6-4. 10.3 cm
312 Answers to Numerical Exercises
Chapter 7
7-2. P 7.8 W
7-3. v [gV(ρw − ρ)/Aρw]1/2; P 1/2[W{(ρw/ρ) −1}3/2]/(Aρw)1/2
7-5. P 1.51 � 107dyn/cm2 15 atm
7-6. Volume of swim bladder 3.8%
7-7. ρ2 ρ1(W1/W1 − W2)
7-8. p 1.46 � 105 dyn/cm2
7-11. Perimeter 9.42 km
7-12. Speed 29 cm/sec
Chapter 8
8-1. P 3.19 � 10−2 torr
8-2. P 4.8 torr
8-3. h 129 cm
8-4(a). p 61 torr; (b). p 200 torr
8-5(b). R1/R2 0.56
8-6. v 26.5 cm/sec
8-7. N 7.5 � 104
8-8. p 79 torr
8-9. P 10.1 W
8-10(a). P 0.25 W; (b). P 4.5 W
Chapter 9
9-2. V 29.3
9-3(a). t 10−2 sec; (b). t 10−5 sec
9-5. N 1.08 � 1020 molecules/sec
9-6. No. breaths/min . 10.4
9-7(a). Rate 1.71 liter/hr-cm2; (b). diameter 0.5 cm
9-8. P 2.87 atm
Chapter 11
11-2. t 373 hours
11-3. v 4.05 m3
11-4. t 105 days
11-5. Weight loss 0.892 kg
11-6. H 18.7 Cal/h
Answers to Numerical Exercises 313
11-8(b). Change 22%; (c). Kr 6.0 Cal/m2-h-C◦
11-9. Heat removed 8.07 Cal/h
11-10. Heat loss 660 Cal/m2-h
11-11. H 14.4 Cal/h
Chapter 12
12-1. R 31.6 km
12-2. 1.75 times
12-3. p 2.9 � 10−4 dyn/cm2
12-6. D 11.5 m
12-8. Min. size 1.7 � 10−2 cm
Chapter 13
13-1(a). No. of ions 1.88 � 1011; (b). no. of Na+ ions 7.09 � 1014/m;
No. of K+ ions 7.09 � 1015/m
13-8(a). no of cells in series 5000; (b). no of cells in parallel 2.7 � 109
Chapter 14
14-1. i 13.3 amp
Chapter 15
15-1. Change in position 0.004 cm
15-3. For cornea 41.9 diopters; for lens, min power 18.7 diopters, max
power 24.4 diopters
15-4. 1/f −0.39 diopters
15-5. Focusing power �70 diopters
15-6. p 1.5 cm
15-7(a). Resolution 2.67 � 10−4 rad; (b). Resolution 6.67 � 10−4 rad
15-8. D 20 m
15-9. H 3 � 10−4 cm
Index
Absorption
electromagnetic radiation, 123, 242
spectroscopy, 245
spectrum, 243
Acceleration
angular, 277
centripetal, 278–279
defined, 273
equations of translational motion for,
30–32
gravitational, 5, 33
of jumper, 31–34
Accommodation of eye, 216–217, 230
Achilles tendon, 19
Actin, 95, 96
Action potential, 184–186
in muscles, 194
in plants, 196
measurement, 195
propagation, 188–190
Adhesion, 90–91
strength of, 93
Adrenaline, 155
Aging, and vision, 216–217
Air
inspired vs. expired, 130–131
motion through, 40–42
pressure in ear, 170
velocity and Kc , 151–152
viscosity, 104
Airbag, 68–69
Alcohol, caloric value, 147
Aluminum
specific heat, 119
thermal conductivity, 122
Alveoli, 129
size, 132
Amino acid, 271
Amplification, in human ear, 175
Amplifier, 201
transistor, 211
Angstrom, 285
Angular acceleration, 277
Angular momentum
defined, 280
equations of, 277–278
Angular motion
forces on a curved path, 45–48
Newton’s laws, 280
pendulum
physical, 51–52
simple, 48–50
running, 53–56
vs. linear motion, 277
walking, 50–53
Angular velocity, 276
maximum, 54, 78–79
314
Index 315
Animal
energetics, 136
motion, 1
sounds produced by, 176
Anvil (middle ear), 169
Aorta, 106
blood pressure drop, 107–108
turbulent flow, 110–111
Aperture of eye, 218–219
Apocrine sweat gland, 155
Aqueous humor, 215–216
Archimedes’ principle
defined, 87
fish buoyancy and, 88–89
power required to stay afloat and,
87–88
Arteriole, 106–107, 109
Arterisclerosis, and blood flow, 111–112
Artery, 105
elasticity, 112
narrowing, 111, 112
natural frequency, 112
plaque deposit, 111–112
pressure drop, 107–108
pulmonary, 105
Astigmatism, 227
lens for, 228, 229
Atom
absorption spectrum, 243
energy state, 241–242
excitation of, 242–243
interactions between, 270
nucleus, 240, 256–257
structure, 239–240
Atomic physics, 239
Axon, 181
action potential, 184–186
action potential propagation,
188–190
capacitance and resistance of, 186
circuit, analysis of, 190–193
diameter of, 183
electrical potentials, 183–184, 185
electrical properties, 186–187
length of, 181
membrane
as leaky insulator, 186
permeability, 184
myelin, 181
myelinated, 192–193
myelinated vs. nonmyelinated, 187
nodes of Ranvier, 181
propagation vs. speed of light, 186
resistance of, 186
sodium pump, 184
voltage, 184
vs. electric cable, 186–188
Back
backaches, 18
lever representation, 17–19
Bacteria, thermophilic, 145
Ballistocardiograph, 115
Barth�elemy, Toussaint, 249
Basal metabolic rate, 147
Basilar membrane, 170
Bats
chirping, 175–176
echo location, 175
Battery, 292
Bernoulli’s equation, 101–102, 103
stenosis and, 111
Biceps, 7
movement of, 11–15
Biological control system, 208
features, 206–207
feedback, 208–210
in iris, 210
Biomechanics, 2
Blood
adrenaline in, 155
cells, radioactive, 269
circulation, 105–107
flow
arterisclerosis, 111–112
control, 109
energetics, 110
laminar, 103, 104, 110, 111
rate, 112–113
to brain, 109
turbulence, 110–111
velocity, 110, 178
kinetic energy, 110
pressure
arterial, 107–109
at capillaries, 107
measurement, 113–114
systolic and diastolic, 107
venal, 109
sugar level, 245
venal, 136
viscosity, 104
316 Index
Bohr model of atom, 240–241, 247
formation of chemical bonds,
243–244
hydrogen, 241
Bohr, Niels, 240
Boltzmann constant, 117–118
Bone
density
cuttlefish, 88–89
electricity and, 196–197
fracture
energy involved, 64–66
force needed to cause, 67–68
neck, 69–70
NMR signal, 261
osteoblasts and osteoclasts, 197
Boyle’s law, 119
Brain
activity identification, 265–266
arteries, 109
blood flow to, 109
diagnosing disorders, 204
ischemic stroke, 112
nerve centers in, 150
nerve impulses, 162
role in hearing, 175
signal processing, 226
Breathing
cold-blooded animals, 132
heat loss by, 155–156, 157
surfactants and, 132
Broad jump
running position, 39–40
standing position, 37–39
Broca, Paul Pierre, 265
Broken heart syndrome, 109
Bruit, 111
Buoyancy, of fish, 88–89
Calorie, 119
intake, 148
Capacitor, 291
Capillary action, 91, 92, 93
Cardiomyopathy, stress, 109
Cardiovascular disease
arterisclerosis, 111–112
stress cardiomyopathy, 109
Catfish spine fin, 27–29
Cell, 270–271
Center of gravity, 2
human body, 3–4
Center of mass motion
in running, 57–58
in walking, 56–57
Centrifugal force, 45–46, 47
defined, 278
Centripetal acceleration, 277–278
Centripetal force, 46, 47, 277–278
defined, 277
137
Cesium, 268
Chatecholamine, 109
Chemical bond, formation of, 243–244
Chemical energy, 139
Chemical fumigation, 267
Chlorine ion, and membrane potential, 184
Chromium isotope, in medicine, 269
Circulatory system, 105–107
body heat transfer and, 151
mechanism of energy losses, 107
turbulent flow, 111–112
60
Cobalt, 268
Cochlea, 170–171
implants, 211–213
Coefficient
convection, 122, 152
diffusion, 127
friction, 24, 25, 46, 71
kinetic, 25, 71
static, 25
thermal conductivity, 121
Collision
automobile, 69–70
duration of, 66–67
force of, 67–68
protective device, 68–69
Compression, 61–62
Computerized tomography, 250–251, 257
Conductance (G), 291
Conduction, thermal, 120–121
in human body, 150, 151
Conductor, 291
Cones and rods, 222, 223, 224, 225–226
Confocal microscopy, 232–235
Conservation
energy, 135–136
Bernoulli’s equation and, 101–102
human physiology and, 136
linear momentum, 275
Constructive interference, 166
Control system, 206–208
feedback, 208–210
Convection, 121–122
in human body, 151–153
Index 317
Converging lens, 294, 295–298
Cooling mechanism, 136, 141, 150,
155–156, 158
Cork, thermal conductivity of, 122
Cornea, 215
receive oxygen by diffusion, 133
refractive power, 220
Coulomb, 287
Coulomb’s law, 287–288
Critical angle, 294
Critical flow velocity, 104
Cromer, A. H., 43
Crystallography, 249–250
CT scan, 250–251, 257
Cut-off blood pressure measurement,
113–114
Cuttlefish, bone density, 88–89
Davidovits, Paul, 232, 233
da Vinci, Leonardo, 1, 7–8
De Broglie, Louis, 246, 247
Defibrillator, 206
Dehydration, 155
Density
constant, 83
of water, and floating, 87–88
porous bones and swim bladders,
88–89
Depth of field, 219
Destructive interference, 166
Diabetic retinopathy, laser treatment, 254
Diagnostic equipment
computerized tomography, 250–251,
257
electrocardiograph, 195, 202–203
electroencephalograph, 195,
203–204
electromyograph (EMG), 195
magnetic resonance imaging,
257–258
stethoscope, 111, 113, 177, 202
X-rays, 249–250
Diastolic pressure, 107
Diathermy, 178
Diffraction, 168
in eye, 224
studies with molecules, 250
Diffusion, 123–125
coefficient, 127
contact lens and, 133
in respiratory system, 129–132
mean free path, 124
molecular transport through,
126–127
random walk, 124–125
through biological membrane,
128–129
Diopter, 219
Dipole field, 289
Diverging lens, 294, 300
DNA, 143–144, 248, 250, 269, 270, 271
Doppler effect, 178
Doughnut, energy content, 43
Dyne, 285
Ear, 168
amplification in, 175
canal, 169
horns, 211
inner, 170–171
balance maintenance, 21
middle, 169–170
outer, 168–169
performance, 171–172
sound detection capability, 172–173
sound intensity, 173–175
threshold of hearing and pain, 173, 174
Eardrum, 162, 168, 169–170, 175
Earth, forces on, 33
Eccrine sweat gland, 155
ECG, See Electrocardiography
Echoes, bats and, 175
EEG, See Electroencephalography
Eel, electric, 198
Egger, M. David, 232, 233
Einstein, Albert, 252
Elasticity, 61
artery, 112
insect wings, 79–80
spring, 62–64
Elbow, movement of, 11–15
Electrical technology, in biological
research, 200–202
Electric charge, 287–288
Electric circuit, 290–292
Electric current, 289
effect on brain, 205
sources, 292
Electric eel, 198
Electric field, 288–289, 292
in water, 198
Electric fish, 197–198
318 Index
Electricity
as a natural phenomena, 180
in bone, 196–197
fish and, 197–198
in plants, 196
magnetism and, 292
nervous system and, 180–196
physiological effects, 204–206
piezoelectricity, 196–197
Electric shock, 204–205
Electrocardiography (ECG), 195, 202–203
Electrode, 202
Electroencephalography (EEG), 195,
203–204
Electromagnetic radiation, 214
energy and, 122–123
excitation of atom and, 242–243
Electromyography (EMG), 195
Electron, 239, 240–241
binding energy, 243
diffraction patterns, 247
electric charge of, 287
energy level, 241–242
excitation, methods of, 242
excited state, 242
ground state, 241–242
high-speed (Beta particles), 256
and food preservation, 268
impact, 242
inner, 243
in oscilloscope, 201
orbital restrictions, 240–241
orbit around nucleus, 240
outer, 243
radiation and, 122–123
shared, 244
wavelength, 247
wavelike properties, 246
Electron microscope, 247–248
EMG, See Electromyography
Emission
spectroscopy, 245
spontaneous, 252
stimulated, 252
Emissivity, 123
of skin, 153
Endoscope, 236–237
Energy, 282–283
chemical, 139
in food, 141
conservation, 135–136
consumption in physical activity,
42–43
load carrying, 58–59
running, 54–56
electromagnetic, 122–123
forms, 283
from food, 147–149
internal, 117, 139
involved in bone fracture, 64–66
kinetic, 283
insect wing in flight, 78–79
of particles in gas, 117
level, 241–242
mechanical, in ultrasonic wave, 178
requirements, 146–149
during pregnancy, 149
unit and conversion, 286
Entropy, 142
Epilepsy, 205
Equilibrium
human body considerations, 3–4
stability and, 2–3
static, 2–3, 282
Eustachian tube, 170
Evaporation, skin temperature control by,
155–156
Excited state, 242
Exercise, osteoarthritis and, 71
Eye
aging and, 216–217
aperture and depth of field, 218–219
eyeglasses, 211
focusing, 216–217, 230
laser treatment, 253–255
lens system, 219–220
light intensity reaching retina,
207–208, 209
near point, 216–217
parameters, 220
reduced, 220–222
resolution of, 223–225
structure, 215–216
vs. camera, 217–218
Eyepiece, 230, 231
Falling
fracture due to, 67–68
from great height, 70
on snow, 70
Farads, 291
Fasting, world record, 149
Index 319
Feedback system, 208–209
negative feedback, 209
positive feedback, 209
Fetus
energy required, 149
heart, examination, 178
Fiber optics, 235
fiberscopes, 236–237
Fibrillation, 205–206
Field line, 288
Fish
buoyancy, 88–89
catfish spine fin, 27–29
electric, 197–198
electronic lures, 176
eye, lens focusing power, 219–220
Flight
insect, 73–80
hovering, 73–75
Fluid
Archimedes’ principle, 87–89
blood, See Blood
body, 183
defined, 82
force and pressure, 82–86
friction and, 103–104
motion of, 101
Bernoulli’s equation, 101–102
laminar, 103, 104
Poiseuille’s law, 103–104, 107–108
turbulent flow, 104–105
viscous friction, 24, 103, 107
surface tension, 89–96
Flux, 126–127, 128
solar, 154
Focal length of lens, 295–297
Focus, principal, of the lens, 295
Food
composition and energy content, 148
energy from, 141, 147–149
oxidation, 147
preservation by fumigation, 267
preservation by radiation, 267–268
requirements for humans, 147–148
Force
addition of torques and, 281–282
adhesive vs. cohesive, 90
centrifugal, 45–46, 47, 278
centripetal, 46, 47, 277–278
contraction of muscle, 96
defined, 274
field, 288
fluid, 82–86
impulsive, 66–67
bone fracture and, 67–68
lines of, 288–289
on a curved path, 45–48
on the foot, 47
pressure in a fluid and, 82–83
static, 1–2
stopping, 69
unit and conversion, 285
Fourier, J. B. J., 171
Fovea, 222
Fracture
due to a fall, 67–68
energy involved, 64–66
neck bone, 69–70
Frequency
larmor, 259–261
natural, of healthy artery, 112
pendulum swings, 48–49
resonant, 167
sound, 163, 164
Friction, 23–24
at hip joint, 26–27
catfish spine fin and, 27–29
coefficient, 24, 25, 46, 71
fluid, in air, 40
standing at an incline, 25–26
viscous, 24, 103, 107
Frog
alveolal radii, 132
diffusion transfer of oxygen,
131–132
neurons in retina, 226–227
Fulcrum, 9–10
Fumigation, chemical, 267
Functional magnetic resonance imaging
(fMRI), 265–266
Fur, 157
Galvani, Luigi, 194
Gamma ray, use in food preservation,
267–268
Gas
behavior, 139
behavior of matter as a function of
temperature in, 117–119
diffusion, 125
greenhouse, 159
noble, 244
pressure, 118
Generator, 292
320 Index
Geometric optics, 293–295
Gland
apocrine, 155
eccrine, 155
Glass
lens, 294
radiation and, 123, 249
silica, 235
Glycerine, viscosity of, 104
Gravitational force, 274
Greenhouse effect, 159
Greenhouse gas, 159
Ground state, 241–242
Gyromagnetic ratio, 258, 259
Hales, Stephen, 113
Hammer (middle ear), 169
Hearing, 168
aids, 211
ear horns, 211
brain’s role in, 175
cochlear implants, 211–213
in bats, 175–176
sound frequency and pitch, 172–173
threshold of, 173, 174
transistorized aids for, 211
Heart
aorta, 106
atrium and ventricle, 105–106
capillaries, 107
desynchronization of heart action,
205–206
fetus, examination, 178
power produced by, 112–113
stress, 109
Heat, 284, See also Thermodynamics
cold and, 156–157
defined, 116
latent, 120
life and, 145–146
loss by breathing, 155–156, 157
radiative by sun, 153–154
specific, 119, 284
therapeutic effects, 161
transfer of
conduction, 120–121, 150, 151
convection, 121–122, 151–153
diffusion, 123–133
in human body, 149–157
radiation, 122–123, 139, 153–154
unit of, 119, 284
vs. other energy forms, 138–140
Helicotrema, 170
Henry, 292
Hertz, 163
Hertz, Heinrich, 163
High jump, 36–37
Hip joint
friction at, 26–27
movement of, 15–17
walking on injured, 17
Hooke, Robert, 62, 63
Hooke’s law, 62, 79
Hormone, 109, 207
Hovering flight, 73–75
power required, 76–79
Human body, See also Specific parts,
organs and systems
adaptation for heat vs. cold, 156
critical temperature, 156
energy requirements, 146–148
food requirements, 147–148
metabolic rate, 146
motion, 1–2
oxygen requirements, 130–131
posture, 19–21
radiative heating, 154
resistance to cold, 156–157
senses, limitations of, 200
sound production, 176
specific heat, 119
stability of, 3–4
under action of external force, 4–7
sweat production, 155–156, 209
temperature
regulation, 149–151
regulation by convection, 151–153
regulation by evaporation, 155–156
regulation by radiation, 153
Hydrogen
Bohr model for atom of, 241
formation of molecule of, 244
nuclear magnetic properties of, 258
Hydrostatic skeleton, 84–86
Hyperopia, 227
lens for, 228, 229
Ice, specific heat of, 119
Image
of extended objects, 298–300
on retina, 217–218
size, 221–222, 223, 229–230
Index 321
real, 297
size of aperture and, 218–219
virtual, 297
Imaging
computerized tomography, 250–251, 257
magnetic resonance imaging (MRI),
257–258
ultrasound, 177–178
with NMR, 262–265
X-ray, 243, 249–250
Impulsive force, 66–67
fracture and, 67–68
Inductor, 292
Inertia, moment of, 279
Infant respiratory distress syndrome, 132
Inner ear, 170–171
balance maintenance, 21
basilar membrane, 170
cochlea, 170–171
implants, 211–213
helicotrema, 170
Insect
flight, 73
hovering, 73–75, 76–79
locomotion on water, 93–95, 99
Microvelia, 99
wing
elasticity, 79–80
kinetic energy when in flight,
78–79
muscles, 75–76
Insulation, fur and feather, 122, 157
Insulator, 291
Intensity
of light
control, in reaching retina,
207–208, 209
of sound, 163
and loudness, 173–175
Interference, 166–167
Internal energy, 117, 139
Internal reflection, total, 294, 295
Interneuron, 181
131
Iodine, 267
Ion
membrane potential and, 183–184
negative, 287
positive, 287
Iris, 215
control system, 210
defined, 207
optical aperture, 218–219
Irradiation, food, 267–268
Ischemic stroke, 112
Isotope, 256
oxygen, 256
radioactive, 257
tracers, 268–269
Joint
hip
friction at, 26–27
movement, 15–17
walking on injured, 17
knee problems, 71
osteoarthritis, 70–71
Jump
broad
from running position, 39–40
from standing position, 37–39
high, 36–37
vertical
effect of gravity on, 35
height of, 32–35
Kilocalorie, 284
Kinesiology, 2
Kinetic energy
defined, 283
insect wing in flight, 78–79
of particles in gas, 117
Kinetic friction, 23–24
coefficient, 25, 71
Kinetic theory of matter, 116–119
Knee joint, problems, 71
Kuhne, W., 217, 218
Laminar flow, 103, 104, 110, 111
Larmor frequency, 259–261
Laser, 252–253
surgery, 253
ophthalmological applications,
253–255
LASIK (Laser-assisted in Situ Keratomileusis),
254–255
Latent heat, 120
Lauterbur, P. C., 263
Lavoisier, Laurent, 135, 136
Lens, 215
astigmatism, 228, 229
contact lens and diffusion, 133
converging, 294, 295–298
diverging, 294, 300
eyepiece, 230, 231
322 Index
Lens (cont.)
immersed in a material medium,
300–301
myopia, 228, 229
objective, 230, 231
of eye, 219–220
focusing power, 216–217
presbyopia and hyperopia, 228, 229
Lever, 9–11
arm, 279
elbow movement, 11–15
hip movement, 15–17
spine movement, 17–19
standing on tip-toe on one foot,
19, 20
Light, 162, 214
emitted by laser, 252
fiber-optic devices and, 237
intensity at retina, 207–208, 209
penetration through tissue, 232
properties, 215
speed, 293
vision and, 214–215
Limping, 17, 18
Linear momentum, 274
conservation, 275
Linear motion, 277
Lines of force, 288
Lipoprotein, solubility, 98
Lithium, 241
Load carrying, energy consumption, 58–59
Logarithmic sound intensity, 174
Long jump, See Broad jump
Loudness, 173–175
Lubrication, 25
effect on human hip joint, 27
Lumbar vertebra, fifth, 17–19
Lung
gas exchange in, 129–130
water vapor and, 155
X-ray, 250
Magnetic moment, 258, 259
Magnetic resonance imaging (MRI), 257–
258, 262–265
functional, 265–266
Magnetism, electricity and, 292
Marangoni propulsion, 99
Mass, 274
unit and conversion, 285
Matter, kinetic theory of, 116–119
Maximum angular velocity, 54, 78–79
Mayer, Robert, 135–136
Mean free path, 124
Medfly (Mediterranean fly), control of,
177
Membrane
axon
as leaky insulator, 186
capacitance and resistance, 186
permeability, 184
basilar, 170
biological, diffusion through,
128–129
oval window in ear, 169
tympanic, 162, 168, 169–170, 175
Membrane protein, solubility, 98
Mercury, viscosity of, 104
Metabolic rate, 145–146
defined, 146
for selected activities, 146
Metabolism, 157
Mho, 291
Micron, 285
Microscope, 231
compound, 231
confocal, 232–235
electron, 247–248
resolution, 231–232
Microvelia, 99
Middle ear, 169–170
Eustachian tube, 170
hammer, anvil, stirrup, 169
ossicles, 169, 170, 175
Miller, S. L., 271
Minsky, Marvin, 233
Moisture tension in soil, 92–93
Molecule
characteristic spectra, 244
diffraction studies with, 250
formation of hydrogen, 244
organic, 270–271
transport through diffusion, 126–127
Moment arm, 279
Moment of inertia, 279
Momentum
angular, 280
equations of, 277–278
linear, 274
conservation, 275
Motion, 1–2
angular
Newton’s laws, 280
vs. linear, 277
Index 323
Newton’s laws, 274–275
rotational, 30, 31
equations for, 278
thermal, 117, 124, 140
through air, 40–42
translational, 30–32
Motor neuron, 181
MRI, See Magnetic resonance imaging
Muscle
action potentials in, 194
biceps, 7, 11–15
contraction, 8, 95–96
efficiency, 42–43
fibers, 194
insect wings, 75–76
myofibrils, 95
skeletal, 7–9, 95–96
spindle, 194
stimulation by electric current, 206
triceps, 7, 11, 12
Musculoskeletal system,
interconnectedness, 21
Myelin, 181
Myelinated axon, 192–193
vs. nonmyelinated, 187
Myofibrils, 95
Myopia, 227
lens for, 228, 229
Myosin, 95, 96
Near point of the eye, 216–217
Neck bone, fracture, 69–70
Negative feedback, 209–210
Negative ion, 287
Nervous system
action potential, 184–186
action potential, propagation, 188–190
electrical phenomena and, 180–181
electrical potentials in axon, 183–184,
185
signal propagation, 181
surface potentials, 194–196
synaptic transmission, 193–194
vision and, 226–227
Neuron, 180, 181–183
axons and dendrites, 181, 183, See also
Axon
classes, 181
Neutron, 287
Newton, 5
Newton, Isaac, 1
Newton’s
laws of angular motion, 280
laws of motion
first, 274
second, 275
third, 275
14
Nitrogen, 269
NMR, See Nuclear magnetic resonance
Noble gas, 244
Nodes of Ranvier, 181
Noise
bruit, 111
laminar flow, 113
Nuclear magnetic resonance (NMR), 257–
262
imaging with, 262–265
Nuclear spin, 261
Nucleus, 240, 256–257
transmutation, 257
Objective lens, 230, 231
Ohm, 290
Ohm’s law, 205, 291
Optical spectra, 243
Optics, 214
fiber, 235–237
geometric, 293–295
vision and, 214–215
Oscilloscope, 201–202
Osmosis, 129
Ossicles, 169, 170, 175
Osteoarthritis, 70–71
exercise and, 71
Osteoblast, 197
Osteoclast, 197
Oudin, Paul, 249
Outer ear, 168–169
ear canal, 169
pinna, 168
tympanic membrane, 162, 168, 169–
170, 175
Oxidation of food, 147
Oxygen
consumption, calories produced by, 147
diffusion through skin, 129
small animals, 131–132
isotopes of, 256
oxidation of food, 147
requirement for humans, 130–131
Pacemaker, 202
electronic, 206
324 Index
Particle, wavelike properties, 246–247
Pascal (Pa), 83
Pascal’s principle, 83–84
Pastuerization, 267
Pendulum
physical, 51–52, 54–56
simple, 48–50
Period, of pendulum motion, 48–49
Phosphorus, radioactive, 257, 267, 269
Photodetector, 245
Photon, 215
Photoreceptor, 222, 225–226
Photosynthesis, 214
Physics and life, 269–271
Piezoelectric effect, 196–197
Pinna, 168
Pitch of sound, 172–173
Planck’s constant, 215, 246–247, 258
Plant
action potential in, 196
electricity in, 196
soil water and, 92–93
Plaque, arterial, 111–112
Poise, 103, 104
Poiseuille, L. M., 101
Poiseuille’s law, 103–104
estimation of blood pressure drop and,
107–108
Positive feedback, 209–210
Positive ion, 287
Posture, 19–21
Potassium ion, axon potential and, 184
Potential
axon, 184
difference, 289
energy, 283–284
Power, 284
defined, 78
generated by limbs, 88
produced by, 112–113
required to hover, 76–79
required to stay afloat, 87–88
unit and conversion, 286
Precession, 260
Pregnancy, energy requirements, 149
Presbyopia, 217
lens for, 229
Pressure
Bernoulli’s equation and, 101
blood
arterial, 107–109
at capillaries, 107
measurement, 113–114
systolic and diastolic, 107
venal, 109
defined, 274
fluid, 82–84
gas, 118
in porous bones, 89
inside worm, 85
on eardrum, 170
Poiseuille’s equation and, 103–104
sound, 164–165, 175
unit and conversion, 285
Principia Mathematica, 1
Projectile, range of, 37
Protein
caloric value, 147
consumption during fasting, 149
resilin, 79–80
solubility of membrane protein and
lipoprotein, 98
specific heat, 119
Proton, 239–240, 287
Pulmonary artery, 105
Pupil, 215
defined, 207
Pure tone, 163–164
P wave, 203
Quality
image, 218
sound, 171
Quantum mechanics, 246–247, 270
Radian, 276
Radiation, 139
electromagnetic, 122–123, 214
food preservation by, 267–268
human body, 153
solar, 153–154
and soil, 159
therapy, 266–267
thermal, 122
Radioactive
isotopes, 257
tracers, 269
Radioactivity, 256–257
Random thermal motion, 124, 140
Random walk, 124–125
Real image, 297
Reduced eye, 220–222
Reflection, 165–166
total internal, 294, 295
Index 325
Refraction, 165–166
defined, 293
index of, 220
refractive power of cornea, 219–220
Resilin, 79–80
Resistance of air, 40–42
Resistivity, 290
Resistor, 290–291
Resolution
eye, 223–225
microscope, 231–232
Resonant frequency, 167
Respiratory system
diffusion process, 129–132
surfactants and breathing, 132
Retina, 215, 222–223
cones and rods, 222, 223, 224,
225–226
degeneration arrest, 253–254
image size on, 221–222, 223,
229–230
light intensity, control of, 207–208, 209
photographic film and, 217–218
Reynold’s number, 104
Righting reflex, 21
Rods and cones, 222, 223, 224, 225–226
Roentgen, Wilhelm Conrad, 249
Rolling friction, 24
Root (plant), and pressure, 92
Rotational motion, 30, 31
equations for, 278
Running
broad jump, 39–40
center of mass motion in, 57–58
energy expended in, 54–56
metabolic rate, 43
on a curved track, 47–48
speed, 53–54
Rupture strength, 63
Rutherford, E., 239, 240
Sensitivity
of ear, 169, 172, 174–175
logarithmic, 174
mechanical reasons for, 175
of eye, 226
Sensory aid, 211
Sensory neuron, 181
Shannon, Claude, 143
Shark, and electric field, 198
Shock, electric, 204–205
stimulation of muscle with, 206
Siemen, 291
Silver, thermal conductivity of, 122
Simple harmonic motion, 48
walking in terms of, 50–51
Sinusoidal sound wave, 163–164, 171, 172
Skeletal muscle, 7–9
contraction, 95–96
Skin
convection and, 151–153
emissivity of, 153
evaporative cooling, 156–157
frostbite, 157
oxygen diffusion through, 129
radiative heating of, 153–154
temperature, 150–151
control, 151
Snell’s law, 235
defined, 293–294
Sodium
ions, 184, 189
pump, 184
Soil
loam vs. clay, 93
moisture tension, 92–93
specific heat, 119
temperature, 158–159
water, 92–93
Solar radiation, 153–154
soil and, 159
Somatosensory system, balance
maintenance, 21
Sound, 162
acoustic traps, 176–177
bell in a jar, 163
clinical uses, 177
frequency, 163, 164, 172–173
intensity, 163
and loudness, 173–175
logarithmic, 174
perception of, 175
pitch, 172–173
produced by animals, 176
properties, 162–165
pure tone, 163–164
speed, 164
wave, 162
wavelength (λ), 164
Specific heat, 119, 284
Spectral line, 240
Spectrometer, 245
Spectroscopy, 244–245
absorption, 245
326 Index
Spectroscopy (cont.)
emission, 245
Spectrum, absorption, 243
Speed
defined, 272
light, 293
running, 53–54
sound, 164
walking, 52–53
Spindle, 194
Spontaneous emission, 252
Spring, 62–64
Squid, axon of, 183
Stability
equilibrium and, 2–3
human body, 4–7
Standing
at an incline, 25–26
broad jump, 37–39
tip-toe on one foot, 19, 20
Standing wave, 166–167
Static equilibrium, 2–3
defined, 282
Static force, 1–2
Static friction, 23–24
coefficient, 25
Stefan-Boltzmann constant, 123
Stenosis, 111, 112
Stethoscope, 111, 113, 177
electronic, 202
Stimulated emission, 252
Stirrup (middle ear), 169
Strength of material, 61
bone, 64–68
Stress
defined, 62
stress cardiomyopathy, 109
Stretching
longitudinal, 61–62
spring, 62–64
Stroke, ischemic, 112
32
Sulphur, 269
Surface potential, 194–196
recording of, 202–203
Surface tension, 89–91
insect locomotion on water and,
93–95, 99
muscle contraction and, 95–96
soil water, 92–93
spherical liquid drops, 91–92
Surfactants, 97–98
breathing and, 132
secreted by insects, 99
Sweating
as negative feedback, 209
cooling mechanisms, 155–156
dehydration, 155
rate, 155
Synapse, 193
synaptic transmission, 193–194
Synovial fluid, 25, 27
Systems approach, 209–210
Systolic pressure, 107
Telescope, 230–231
Temperature, 117–118
body, regulation of, 149–151
critical, 156
defined, 117
skin, 150–151
Terminal velocity, 41–42
Thermal conductivity, 120–121
in human body, 150, 151
Thermal motion, 117
random, 124, 140
Thermal radiation, 122
emitted by soil, 158–159
Thermal velocity, 118
Thermodynamics, See also Heat
defined, 135
first law, 135–136
of living systems, 140–142
second law, 137–138
information and, 143–144
Thermophilic bacteria, 145
Thompson, J. J., 239
Threshold
of hearing, 173, 174
of pain, 173, 174
of vision, 225–226
Tissue
light penetration, 232
thermal conductivity, 122, 150
Tomography, computerized, 250–251, 257
Torque, 279–280
addition of force and, 281–282
Torr, 83
Torricelli, Evangelista, 83
Total internal reflection, 294, 295
Tracer, isotopic, 268–269
Transistor amplifier, 211
Translational motion, 30
energy consumption, 42–43
for constant acceleration, 30–32
high jump, 36–37
Index 327
long jump
standing, 37–39
running, 39–40
projectile range, 37
through air, 40–42
vertical jump, 32–35
Transmutation of nucleus, 257
Transport, of molecules, 126–127
Triceps, 7
movement of, 11, 12
Turbulent fluid flow, 104–105
blood, 110–111
T wave, 203
Tympanic membrane, 162, 168, 169–170,
175
Ultrasonic
diathermy, 178
flow meter, 178
waves, 177–178
Ultrasound imaging, 177–178
Unit
calorie, 119
coulomb, 287
diopter, 219
dyne, 285
farads, 291
henry, 292
hertz, 163
kilocalorie, 284
mho, 291
newton, 5
of energy, 286
of force, 285
of length, 285
of mass, 285
of power, 286
of pressure, 285
pascal (Pa), 83
poise, 103, 104
radian, 276
siemen, 291
torr, 83
Uranium, isotopes of, 257
Vein, 105
blood pressure in, 109
pulmonary, 105
Velocity
angular, 276
maximum, 54, 78–79
critical flow, 104
defined, 272
terminal, 41–42
thermal, 118
Venule, 107
Vertical jump
effect of gravity on, 35
height of, 32–35
Vespertilionidae bat, echo location,
175–176
Vestibular system, balance maintenance,
21
Virtual image, 297
Viscosity, and Poiseuille’s law, 103–104
Viscous friction, 24, 103, 107
Vision, 214–215
astigmatic, 227, 228, 229
hyperopic, 227, 228, 229
image quality, 218–219
myopic, 227, 228, 229
nervous system and, 226–227
presbyopic, 217, 229
range, 229–230
threshold of, 225–226
Vitreous humor, 216
Vocal cord, 176
Voltage, 289
and current sources, 292
Walking, 50
center of mass motion in, 56–57
on injured hip, 17
simple harmonic motion, 50–51
speed, 52–53
Water
content of food, 148
density of, and floating, 87–88
elimination from body, 148
index of refraction, 220
insect locomotion on, 93–95
latent heat of vaporization, 155
mean free path of molecules in, 124
osmosis, 129
sea, 89
soil, 92–93
sound and, 166
specific heat, 119
speed of sound in, 164
surface tension, 89
viscosity, 104
Wave, See also Sound
defined, 162
328 Index
Wave (cont.)
diffraction, 168
fundamental and harmonic, 171, 172
interference, 166–167
P, 203
reflection and refraction, 165–166
standing, 166–167
T, 203
ultrasonic, 177–178
wavelength, 164
Weight, 274
loss, 155
Whiplash injury, 69–70
Work
chemical energy and, 42
defined, 43, 282
heat converted into, 139–140
implication of second law of thermodynamics,
138
muscular movement, 42
Worm
hydrostatic forces in moving, 84–86
movement of, 84
X-ray, 243, 249–250
computerized tomography, 250–251,
257
of lungs, 250
Young’s modulus, 62
of resilin, 79–80
rupture strength for materials and, 65
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