Radiation Biophysics - 2nd Edition

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 SECOND EDITION

Edward L. Alpen
Department of Biophysics, Emeritus
University of California
Berkeley, California
and
Department of Radiology, Emeritus
University of California School of Medicine
San Francisco, California

ACADEMIC PRESS

San Diego London Boston New York Sydney Tokyo Toronto

This book is printed on acid-free paper.

Copyright 9 1998, 1990 by ACADEMIC PRESS

All Rights Reserved.

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Library of Congress Card Catalog Number: 97-074421

International Standard Book Number: 0-12-053085-6

PRINTED IN THE UNTIED STATES OF AMERICA

97 98 99 00 01 02 MM 9 8 7 6 5 4 3 2 1
 To my wife, Wynella,
for her unflagging support
and especially for her role
as my proofreader and critic
This Page Intentionally Left Blank
Contents

Preface to the Second Edition xxiii

Preface to the First Edition xxv
Introduction: An Historical Perspective xxvii

Chapter 1
Quantities, Units, and Definitions
Quantities and Units 1
Fundamental Units 2
Derived Units 2
Special Units 2
Radiation Measurement 4
Definitions 4
Directly Ionizing Particles
Indirectly Ionizing Particles 4
Gamma Rays and X-Rays 4
Quantities and Units 5
Exposure 5
Dose (Absorbed Dose) 5
Energy Imparted 5
Equivalent Dose 6
Relative Biological Effectiveness
Particle Fluence 7
Panicle Flux Density 7
Energy Fluence 7
Energy Flux Density 8

~
VU
viii Contents

Kerma 8
Linear Energy Transfer
Charged Particle Equilibrium
Radioactivity Measurements
Decay Constant 9
Activity 10
References 10

Chapter 2
Electromagnetic Radiation: Its Nature
and Properties
Introduction II
Quantum Theory of Electromagnetic Radiation 15
Special Relativity 17
Mass-Energy Equivalence: Einstein's Formulations 18
Relativistic Considerations of Mass and Velocity 19
Atomic Structure 20
Thomson Charge Cloud Model 20
Rutherford's Planetary Model of the Atom 21
Rutherford-Bohr Model of the Atom 21
de Broglie Wave Theory 24
References 25
Suggested Additional Reading 25
Problems 26

Chapter 3
Radioactivity
Introduction 27
Unit of Radioactivity 27
Law of Radioactive Decay 28
Definition of Activity 29
Mean Life 29
Contents ix

Radioactive Decay of Mixtures 30

Chain Decay 30
General Cases for Chain Decay 31
Secular Equilibrium 32
Transient Equilibrium 32
Nonequilibrium 34
Branching Decay Processes 34
Nomenclature of Radioactive Decay 36
Charting Decay Schemes 37
Nuclear Stability 38
Nuclear Mass and Binding Energy 39
Mass Defect 39
Mass Decrement 40
Radioactive Decay by Alpha-Particle Emission 41
Properties of Alpha Decay 42
Negative Electron Emission Decay 42
Positive Electron Emission Decay 43
Annihilation Reaction 44
Decay by Electron Capture 45
K-Capture Fluorescent Radiation 46
Internal Conversion 46
Auger Electron 47
References 48
Suggested Additional Reading 48
Problems 48

Chapter 4
Interaction of Radiation with Matter
Introduction 50
Linear Attenuation Coefficient 51
"Good" and "Bad" Geometry 53
Mass, Electronic, and Atomic Attenuation Coefficients 54
Definition of Cross Section 55
Energy Transfer and Energy Absorption 56
Mechanisms of Energy Transfer from Gamma Rays 58
Photoelectric Scattering Process 58
Energy Transferred-Energy Absorbed Relationships:
Photoelectric Scattering 61
Spatial Distribution of the Emission of Photoelectrons 62
x Contents

Compton Scattering Process: Incoherent Scattering 62

Limits of Energy Transferred 65
Energy Absorbed 66
Summary of the Compton Process 67
Scattering Coefficients 67
Energy of Compton Electrons 70
Electron Binding Energy Effects 70
Pair Production 71
Annihilation Reaction 72
Energy Transferred and Energy Absorbed 73
Summary of the Pair Production Process 73
Triplet Production 74
Variation of Cross Section with Energy 74
Bremsstrahlung--Radiative Energy Loss 74
References 76
Suggested Additional Reading 76
Problems 76

Chapter 5
Energy Transfer Processes
Introduction 78
Importance of the Compton Process in Tissue Systems 79
Total Attenuation Coefficient 80
Total Attenuation Coefficients for Mixtures 83
Interaction of Charged Particles with Matter 83
Final Steps in Energy Absorption 87
Multiple Collision Energy Transfer 87
Photoelectric Process 88
Bremsstrahlung Generation 88
Direct Collisions 89
Life History of a Fast Electron 89
Dose 89
Absorbed Dose and Kerma 90
Neutron Interactions in Tissue 92
Elastic Scatter 93
Inelastic Scatter 94
Nonelastic Scatter 94
Contents xi

Neutron Capture 95
Spallation 95
Kerma and Dose from Neutrons 96
Track Structure and Microdosimetry 96
Rossi Microdosimetric Formulation 97
Linear Energy Transfer 98
Specification of Radiation Quality 98
Local Energy Density (Z) 100
Significance of D(Y) and D(Z) 101
References 102
Suggested Additional Reading 102
Problems 102

Chapter 6
Radiation Chemistry
Introduction 104
Stochastic Nature of Energy Transfer 104
Spurs, Blobs, and Tracks 105
Radiation Chemistry of Water 107
Primary Products of Radiolysis 107
Further Radical Chemistry 108
Recombination 109
Chemical Stage 109
G Value: Expression of Yield in Radiation Chemistry 109
Reactions in the Track: The Role of Scavengers 110
Fricke Dosimeter 111
Interpretation of the Fricke Model 112
Direct and Indirect Action 113
Direct Action 114
Molecular Weight by Direct Action 114
Indirect Action 116
Recombination, Restitution, and Repair 116
Recombination 116
Restitution 117
Macromolecular Target in the Cell 117
Evidence for DNA as the Target Molecule 118
Reactions of the Products of Water Radiolysis 119
Reactions with DNA 120
xii Contents

Chain Scission in DNA 122

Role of DNA Configuration 124
Chromatin Structure 124
DNA Structure and Radiation Damage 124
Repair of DNA 125
Excision Repair 126
Error-Prone Repair 127
Repair of Double-Strand Breaks 127
Repair Fidelity 128
Fidelity of Single-Strand Break Repair 128
Recombinational Repair Fidelity 129
References 129
Suggested Additional Reading 130
Problems 130

Chapter 7
Theories and Models for Cell Survival
Introduction 132
Clonogenic Survival 132
Lea's Target Theory Model 133
Basic Assumptions 134
Biological Survival Curves 135
Exponential Survival Curve 135
Threshold-Type Survival Curve 135
Development of the Target Theory Model 136
Assumptions 137
Derivation 138
General Survival Equation 139
Single-Hit Model 140
Multitarget- Single-Hit Survival 141
Assumptions 141
Properties of the Multitarget-Single-Hit Function 142
Quasi-Threshold Dose 143
Single-Target-Multihit Model 143
Shortcomings of the Multitarget-Single-Hit Model 144
Molecular Models for Cell Death 144
Need for an Alternative Model 144
Role of Enzymatic Repair 145
Contents xiii

Molecular Theory of Radiation Action 146

Assumptions 146
Development of the Molecular Model 147
Two Mechanisms for DNA Damage 147
Derivation of the Molecular Model 148
Linear-Quadratic Formulation 150
Theory of Dual Radiation Action 151
Background 151
Derivation 152
Assumptions 152
Survival Equation 154
Significance of the Dual-Radiation-Action Model 154
Repair-Misrepair Model of Cell Survival 155
Assumptions 155
Formulation of the Repair-Misrepair Model 155
Eurepair and Misrepair 158
Case I: Linear Eurepair-Quadratic Misrepair 158
Comparison with the Conventional Multitarget-
Single-Hit Model 159
Case II: Linear Repair Is Not Always Eurepair 160
Lethal-Potentially Lethal Model 160
Assumptions 160
During Irradiation 162
After Irradiation 162
Survival Equation 163
Low-Dose-Rate Approximation 164
High-Dose-Rate Approximation 164
Linear-Quadratic Approximation 165
Summation 165
References 166
Suggested Additional Reading 167
Problems 167

Chapter 8
Survival Curve and Its Significance
Introduction 169
Technique of the Clonogenic Survival Curve 170
Determination of the Surviving Fraction 171
Feeder Cells 171
xiv Contents

Characteristics of the Mammalian Cell Survival Curve 172

Significance of the Shoulder on the Survival Curve 174
Repair of Sublethal Damage 177
Interpretation of Sublethal Damage 178
High and Low Linear Energy Transfer
and Sublethal Damage 179
Repair of Potentially Lethal Damage 182
Expression of Potentially Lethal Damage Repair 183
Fixation of Potentially Lethal Damage
with Hypertonic Saline 183
Cell Survival and Cell Age 185
Life Cycle of the Cell 185
Cell Age and Radiosensitivity 186
Radiation Induced Cell Progression Delay 189
Mechanisms: Radiation Sensitivity, Progression Delay,
and the Cell Cycle 190
Cell Cycle and Radiation Sensitivity 190
Cell Progression Delay 190
References 191
Suggested Additional Reading 192
Problems 192

Chapter 9
Modification of the Radiation Response
Introduction 194
Role of Water 195
Temperature and Radiation Damage 197
T Less Than 100 K 199
T Greater Than 100 K and Less Than 170 K 199
T Greater Than 170 K and Less Than 420 K 199
Oxygen Effect 200
Oxygen Enhancement Ratio 202
Effect of Oxygen Concentration 203
Time Dependence of the Oxygen Effect 207
Mechanisms of the Oxygen Effect 209
Thiols and Modification of Radiation Response 210
Protection by Exogenous Thiols 212
Synthetic Organic Thiols 213
Contents xv

Nitroaromatic Radiation Sensitizers 215

Mechanism of Action of the Nitroaromatics 217
Sensitization by 5-Halogen-Substituted Pyrimidines 217
References 219
Suggested Additional Reading 220
Problems 220

C h a p t e r 10
Radiation Biology of Normal and
Neoplastic Tissue Systems
Introduction 222
Cell Death in Mammalian Tissues 223
Nature of Cell Populations in Tissue 224
Cell Population Kinetics and Radiation Damage 226
Growth Fraction and Its Significance 226
Cell Kinetics in Normal Tissues and Tumors 228
Models for Cell Survival in Normal Tissues and Tumors 229
Models for Radiobiological Sensitivity of Neoplastic Tissues 230
Hewitt Dilution Assay 230
Lung Colony Assay System 233
Tumor Growth and Tumor "Cure" Models 234
Tumor Volume versus Time 234
TCDso, Tumor Cure 236
Radiobiological Responses of Tumors 236
Hypoxia and Radiosensitivity in Tumor Cells 238
Assay Models for Normal Tissues in Vivo 242
Acute Response of Normal Tissue 242
Hematopoietic System: The Colony Forming Unit 243
Radiation Sensitivity of the Colony Forming Unit 245
Gastrointestinal Crypt Cell Assay 247
Radiation Sensitivity of Gastrointestinal Crypt Cells 248
Spermatogenesis and in Vivo Assays 250
Testis Weight Loss Assay 251
Testis Radiosensitivity-Weight Loss Methods 251
Tubular Regeneration Clonogenic Assay 253
Summary of the Radiosensitivity of the Testis 254
Assays for the Radiosensitivity of Skin 254
Radiation Sensitivity of Skin 255
xvi Contents

Acute Lethal Response in Mammals 255

Principal Organ System Effects 256
Blood Forming Organs 256
Gastrointestinal Tract 259
Lymphoid System 260
Central Nervous System 260
Acute Radiation Syndrome in Man 261
Effects of Dose Rate or Protraction 262
Radiation Effects on the Embryo and Fetus 263
Developmental Sequence and Radiation Effects on Prenatal
Development in the Rodent 264
Preimplantation Period 265
Period of Organogenesis 267
Stage of Fetal Growth 267
Survival of the Murine Conceptus 268
Fertilized Zygote 268
Embryo 268
Interpretation of Animal Findings for Human Subjects 269
Severe Mental Retardation 269
Microcephaly 269
References 270
Suggested Additional Reading 272
Problems 272

C h a p t e r 11
Late Effects of Radiation on Normal Tissues:
Nonstochastic Effects
Introduction 275
Stochastic versus Nonstochastic Effects 277
Radiation Induced Late Pathology in Organ Systems 279
Vascular Endothelium as the Target Tissue 280
Role of Stromal and Parenchymal Damage 282
Functional Subunits 282
Late Effects in Normal Tissue Systems and Organs 283
Gastrointestinal Tract 284
Esophagus 284
Stomach 285
Small and Large Intestine 285
Rectum 286
Contents xvii

Skin 286
Liver 288
Kidneys 288
Lung 289
Central Nervous System 290
Brain 290
Spinal Cord 291
Eye 292
Cataractogenesis 292
Threshold for Cataractogenesis 292
Fractionation and Protraction of Exposure in the Modification of Late
Radiation Injury 293
First Appearance of the Power Law 294
Strandqvist Relationship 294
Nominal Standard Dose (NSD) 295
Summary of the Power Law Relationships 296
Repair and Repopulation after Irradiation 297
Douglas and Fowler F e Formulation 300
Significance of the a / f l Ratio 303
Withers Extension of the F e Model 304
References 305
Suggested Additional Reading 307
Problems 307

Chapter 12
Stochastic Effects Radiation Carcinogenesis
Introduction 308
Historical 308
Stochastic versus Nonstochastic Effects 309
Arguments for and against the Stochastic Model 310
Bases for Our Knowledge of Radiation Carcinogenesis 310
Radiation Carcinogenesis in Experimental Animals 311
Clonal Theory of Carcinogenesis 314
Latency for Tumor Development 316
Initiation-Promotion Hypothesis 317
Oncogenes in Animal and Human Tumors 318
Dose-Rate Effects 319
xviii Contents

Transformed Cell in Vitro 320

In Vitro Cell Transformation Systems 320
Dose-Response Relationship for Transformation 323
Mouse Cell Lines 323
Hamster Embryo Fibroblast System 324
Rodent-Human Cell Differences 324
Effects of Dose Fractionation and Dose Rate 325
High Linear Energy Transfer Radiation Effects 325
Promoters and Cell Transformation 326
In Vivo-in Vitro Assay Systems 327
Role of Viruses in Carcinogenesis 328
Radiation Carcinogenesis in Human Populations 329
Occupational Exposure 329
Medical Exposure 330
Nuclear Weapon Detonation and Accident
Related Exposures 330
Approaches to Risk Estimation 332
Shape of the Dose-Response Relationship 332
Linear Dose-Response Relationship 332
Linear-Quadratic-Dose-Response Relationship 333
Latent Period 334
Absolute Risk versus Relative Risk 335
Absolute Risk 336
Relative Risk 337
Organ-Specific Radiogenic Cancer in Human Beings 338
References 340
Suggested Additional Reading 343

Chapter 13
Stochastic Effects Genetic Effects of
Ionizing Radiation
Introduction 344
Structural Changes in Chromosomes 345
Chromosome Breakage 346
Single-Hit Breakage 348
Inversion of the Fragment or Rejoining at the Wrong End 348
Two-Hit Breakage 349
Multiple Hits in Replicated Chromosomes 350
Breakage Hypothesis and Exchange Hypothesis 352