Ebook Radiobiology for the radiologist (8/E): Part 1

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Library of Congress Cataloging-in-Publication Data
Names: Hall, Eric J., author. | Giaccia, Amato J., author.
Title: Radiobiology for the radiologist / Eric J. Hall, Amato
J. Giaccia.
Description: Eighth edition. | Philadelphia : Wolters Kluwer,
[2019] | Includes bibliographical references and index.
Identifiers: LCCN 2017057791 | ISBN 9781496335418

Subjects: | MESH: Radiation Effects | Radiobiology | Radiotherapy
Classification: LCC R895 | NLM WN 600 | DDC 616.07/57—dc23 LC record
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3


Preface to the First Edition
This book, like so many before it, grew out of a set of lecture notes. The lectures
were given during the autumn months of 1969, 1970, and 1971 at the ColumbiaPresbyterian Medical Center, New York City. The audience consisted primarily
of radiology residents from Columbia, affiliated schools and hospitals, and
various other institutions in and around the city.
To plan a course in radiobiology involves a choice between, on the one hand,
dealing at length and in depth with those few areas of the subject in which one
has personal expertise as an experimenter or, on the other hand, surveying the
whole field of interest to the radiologist, necessarily in less depth. The former
course is very much simpler for the lecturer and in many ways more satisfying; it
is, however, of very little use to the aspiring radiologist who, if this course is
followed, learns too much about too little and fails to get an overall picture of
radiobiology. Consequently, I opted in the original lectures, and now in this
book, to cover the whole field of radiobiology as it pertains to radiology. I have
endeavored to avoid becoming evangelical over those areas of the subject which
interest me, those to which I have devoted a great deal of my life. At the same
time I have attempted to cover, with as much enthusiasm as I could muster and
from as much knowledge as I could glean, those areas in which I had no
particular expertise or personal experience.
This book, then, was conceived and written for the radiologist—specifically,
the radiologist who, motivated ideally by an inquiring mind or more realistically
by the need to pass an examination, elects to study the biological foundations of
radiology. It may incidentally serve also as a text for graduate students in the life
sciences or even as a review of radiobiology for active researchers whose
viewpoint has been restricted to their own area of interest. If the book serves

these functions, too, the author is doubly happy, but first and foremost, it is
intended as a didactic text for the student of radiology.
Radiology is not a homogenous discipline. The diagnostician and therapist
have divergent interests; indeed, it sometimes seems that they come together
only when history and convenience dictate that they share a common course in
physics or radiobiology. The bulk of this book will be of concern, and hopefully
of interest, to all radiologists. The diagnostic radiologist is commended
particularly to Chapters 11, 12, and 13 concerning radiation accidents, late
effects, and the irradiation of the embryo and fetus. A few chapters, particularly
4


Chapters 8, 9, 15, and 16, are so specifically oriented towards radiotherapy that
the diagnostician may omit them without loss of continuity.
A word concerning reference material is in order. The ideas contained in this
book represent, in the author’s estimate, the consensus of opinion as expressed in
the scientific literature. For ease of reading, the text has not been broken up with
a large number of direct references. Instead, a selection of general references has
been included at the end of each chapter for the reader who wishes to pursue the
subject further.
I wish to record the lasting debt that I owe my former colleagues at Oxford
and my present colleagues at Columbia, for it is in the daily cut and thrust of
debate and discussion that ideas are formulated and views tested.
Finally, I would like to thank the young men and women who have regularly
attended my classes. Their inquiring minds have forced me to study hard and
reflect carefully before facing them in a lecture room. As each group of students
has grown in maturity and understanding, I have experienced a teacher’s
satisfaction and joy in the belief that their growth was due in some small
measure to my efforts.

E. J. H.
New York
July 1972

5


Preface
The eighth edition is a significant revision of this textbook and includes new
chapters that were not included in the seventh edition. We have retained the
same format as the seventh edition, which divided the book into two parts.
Section I contains 16 chapters and represents both a general introduction to
radiation biology and a complete self-contained course in the subject, suitable for
residents in diagnostic radiology and nuclear medicine. It follows the format of
the syllabus in radiation biology prepared by the Radiological Society of North
America (RSNA). Section II consists of 12 chapters of more in-depth material
designed primarily for residents in radiation oncology.
Dickens’s famous beginning to a Tale of Two Cities, “It was the best of
times, it was the worst of times, it was the age of wisdom, it was the age of
foolishness, it was the epoch of belief, it was the epoch of incredulity . . . ,” very
much applies to the current world order. Although medical science and
technology have made great advances in alleviating disease and suffering,
irrational and unpredictable events occur quite frequently, instilling fear and
apprehension about potential nuclear terrorism. The eighth edition contains a
new chapter (Chapter 9) on “Medical Countermeasures to Radiation Exposure”
that summarizes the current therapies available to prevent or mitigate radiation
damage to normal tissues. This chapter nicely complements Chapter 14 on
“Radiologic Terrorism.”
Due to the strong request for including more information on molecular
techniques, we have included a new Chapter 17 on “Molecular Techniques in

Radiology.” The techniques described in this chapter should be useful to both the
novice as well as the skilled practitioner in molecular biology.
In this edition, we have eliminated the chapter on “Molecular Imaging.” The
basis for this decision was that the subject matter covered in this chapter does not
involve any radiobiologic principles, and in any case, there are several textbooks
devoted solely to the subject of molecular imaging. For these reasons, we have
decided to remove this chapter from the eighth edition. Overall, we believe that
this new edition represents a well-balanced compilation of both traditional and
molecular radiation biology principles.
The ideas contained in this book represent, we believe, the consensus of
opinion as expressed in the scientific literature. We have followed the precedent
of previous editions, in that, the pages of text are unencumbered with flyspeck6


like numerals referring to footnotes or original publications, which are often too
detailed to be of much interest to the general reader. On the other hand, there is
an extensive and comprehensive bibliography at the end of each chapter for
those readers who wish to pursue the subject further.
We commend this new edition to residents in radiology, nuclear medicine,
and radiation oncology, for whom it was conceived and written. If it serves also
as a text for graduate students in the life sciences or even as a review of basic
science for active researchers or senior radiation oncologists, the authors will be
doubly happy.

Eric J. Hall
Columbia University, New York
Amato J. Giaccia
Stanford University, California
October 2017

7


Acknowledgments
We would like to thank the many friends and colleagues who generously and
willingly gave permission for diagrams and illustrations from their published
work to be reproduced in this book.
Although the ultimate responsibility for the content of this book must be
ours, we acknowledge with gratitude the help of several friends who read
chapters relating to their own areas of expertise and made invaluable suggestions
and additions. With each successive edition, this list grows longer and now
includes Drs. Ged Adams, Philip Alderson, Sally Amundson, Joel Bedford,
Roger Berry, Max Boone, Victor Bond, David Brenner, J. Martin Brown, Ed
Bump, Denise Chan, Julie Choi, James Cox, Nicholas Denko, Bill Dewey, Mark
Dewhirst, Frank Ellis, Peter Esser, Stan Field, Greg Freyer, Charles Geard,
Eugene Gerner, Julian Gibbs, George Hahn, Simon Hall, Ester Hammond, Tom
Hei, Robert Kallman, Richard Kolesnick, Norman Kleiman, Gerhard Kraft,
Adam Krieg, Edward LaGory, Dennis Leeper, Howard Lieberman, Philip Lorio,
Edmund Malaise, Gillies McKenna, Mortimer Mendelsohn, George Merriam,
Noelle Metting, Jim Mitchell, Thomas L. Morgan, Anthony Nias, Ray Oliver,
Stanley Order, Tej Pandita, Marianne Powell, Simon Powell, Julian Preston,
Elaine Ron, Harald Rossi, Robert Rugh, Chang Song, Fiona Stewart, Herman
Suit, Robert Sutherland, Roy Tishler, Len Tolmach, Liz Travis, Lou Wagner,
John Ward, Barry Winston, Rod Withers, and Basil Worgul. The principal credit
for this book must go to the successive classes of residents in radiology,
radiation oncology, and nuclear medicine that we have taught over the years at
Columbia and Stanford, as well as at American Society for Radiation Oncology
(ASTRO) and RSNA refresher courses. Their perceptive minds and searching
questions have kept us on our toes. Their impatience to learn what was needed of
radiobiology and to get on with being doctors has continually prompted us to

summarize and get to the point.
We are deeply indebted to the U.S. Department of Energy, the National
Cancer Institute, and the National Aeronautical and Space Administration, which
have generously supported our work and, indeed, much of the research
performed by numerous investigators that is described in this book.
We owe an enormous debt of gratitude to Ms. Sharon Clarke, who not only
typed and formatted the chapter revisions but also played a major role in editing
and proofreading. Our publisher, Lauren Pecarich, guided our efforts at every
8


stage.
Finally, we thank our wives, Bernice Hall and Jeanne Giaccia, who have
been most patient and have given us every encouragement with this work.

9


Contents
Preface to the First Edition
Preface
Acknowledgments

SECTION I For Students of Diagnostic Radiology, Nuclear
Medicine, and Radiation Oncology
1 Physics and Chemistry of Radiation Absorption
TYPES OF IONIZING RADIATIONS
Electromagnetic Radiations
Particulate Radiations
ABSORPTION OF X-RAYS

DIRECT AND INDIRECT ACTION OF RADIATION
ABSORPTION OF NEUTRONS
ABSORPTION OF PROTONS AND HEAVIER IONS SUCH AS
CARBON
SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
2 Molecular Mechanisms of DNA and Chromosome Damage and Repair
GENERAL OVERVIEW OF DNA STRAND BREAKS
MEASURING DNA STRAND BREAKS
DNA REPAIR PATHWAYS
Base Excision Repair
Nucleotide Excision Repair
DNA Double-Strand Break Repair
Nonhomologous End-Joining
Homologous Recombination Repair
Crosslink Repair
10


Mismatch Repair
RELATIONSHIP BETWEEN DNA DAMAGE AND CHROMOSOME
ABERRATIONS
CHROMOSOMES AND CELL DIVISION
THE ROLE OF TELOMERES
RADIATION-INDUCED CHROMOSOME ABERRATIONS
EXAMPLES OF RADIATION-INDUCED ABERRATIONS
CHROMOSOME ABERRATIONS IN HUMAN LYMPHOCYTES
SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
3 Cell Survival Curves

REPRODUCTIVE INTEGRITY
THE IN VITRO SURVIVAL CURVE
THE SHAPE OF THE SURVIVAL CURVE
MECHANISMS OF CELL KILLING
DNA as the Target
The Bystander Effect
Apoptotic and Mitotic Death
Autophagic Cell Death
Senescence
SURVIVAL CURVES FOR VARIOUS MAMMALIAN CELLS IN
CULTURE
SURVIVAL CURVE SHAPE AND MECHANISMS OF CELL DEATH
ONCOGENES AND RADIORESISTANCE
GENETIC CONTROL OF RADIOSENSITIVITY
INTRINSIC RADIOSENSITIVITY AND CANCER STEM CELLS
EFFECTIVE
REGIMEN

SURVIVAL

CURVE

FOR

CALCULATIONS OF TUMOR CELL KILL
11

A

MULTIFRACTION

Problem 1
Answer
Problem 2
Answer
Problem 3
Answer
Problem 4
Answer
THE RADIOSENSITIVITY OF MAMMALIAN CELLS COMPARED
WITH MICROORGANISMS
SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
4 Radiosensitivity and Cell Age in the Mitotic Cycle
THE CELL CYCLE
SYNCHRONOUSLY DIVIDING CELL CULTURES
THE EFFECT OF X-RAYS ON SYNCHRONOUSLY DIVIDING CELL
CULTURES
MOLECULAR CHECKPOINT GENES
THE EFFECT OF OXYGEN AT VARIOUS PHASES OF THE CELL
CYCLE
THE AGE-RESPONSE FUNCTION FOR A TISSUE IN VIVO
VARIATION OF SENSITIVITY WITH CELL AGE FOR HIGH–LINEAR
ENERGY TRANSFER RADIATIONS
MECHANISMS FOR THE AGE-RESPONSE FUNCTION
THE POSSIBLE IMPLICATIONS
FUNCTION IN RADIOTHERAPY

OF

SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
5 Fractionated Radiation and the Dose-Rate Effect

12

THE

AGE-RESPONSE


OPERATIONAL CLASSIFICATIONS OF RADIATION DAMAGE
Potentially Lethal Damage Repair
Sublethal Damage Repair
MECHANISM OF SUBLETHAL DAMAGE REPAIR
REPAIR AND RADIATION QUALITY
THE DOSE-RATE EFFECT
EXAMPLES OF THE DOSE-RATE EFFECT IN VITRO AND IN VIVO
THE INVERSE DOSE-RATE EFFECT
THE DOSE-RATE EFFECT SUMMARIZED
BRACHYTHERAPY OR ENDOCURIETHERAPY
Intracavitary Brachytherapy
Permanent Interstitial Implants
SUMMARY OF PERTINENT CONCLUSIONS
Potentially Lethal Damage Repair
Sublethal Damage Repair
Dose-Rate Effect
Brachytherapy
BIBLIOGRAPHY

6 Oxygen Effect and Reoxygenation
THE NATURE OF THE OXYGEN EFFECT
THE TIME AT WHICH OXYGEN ACTS AND THE MECHANISM OF
THE OXYGEN EFFECT
THE CONCENTRATION OF OXYGEN REQUIRED
CHRONIC AND ACUTE HYPOXIA
Chronic Hypoxia
Acute Hypoxia
THE FIRST EXPERIMENTAL DEMONSTRATION OF HYPOXIC
CELLS IN A TUMOR

13


PROPORTION OF HYPOXIC CELLS IN VARIOUS ANIMAL TUMORS
EVIDENCE FOR HYPOXIA IN HUMAN TUMORS
TECHNIQUES TO MEASURE TUMOR OXYGENATION
Oxygen Probe Measurements
Markers of Hypoxia
REOXYGENATION
TIME SEQUENCE OF REOXYGENATION
MECHANISM OF REOXYGENATION
THE IMPORTANCE OF REOXYGENATION IN RADIOTHERAPY
HYPOXIA AND CHEMORESISTANCE
HYPOXIA AND TUMOR PROGRESSION
SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
7 Linear Energy Transfer and Relative Biologic Effectiveness
THE DEPOSITION OF RADIANT ENERGY
LINEAR ENERGY TRANSFER

RELATIVE BIOLOGIC EFFECTIVENESS
RELATIVE BIOLOGIC EFFECTIVENESS AND FRACTIONATED
DOSES
RELATIVE BIOLOGIC EFFECTIVENESS FOR DIFFERENT CELLS
AND TISSUES
RELATIVE BIOLOGIC EFFECTIVENESS AS A FUNCTION OF
LINEAR ENERGY TRANSFER
THE OPTIMAL LINEAR ENERGY TRANSFER
FACTORS
THAT
EFFECTIVENESS

DETERMINE

RELATIVE

BIOLOGIC

THE OXYGEN EFFECT AND LINEAR ENERGY TRANSFER
RADIATION WEIGHTING FACTOR
SUMMARY OF PERTINENT CONCLUSIONS
14


BIBLIOGRAPHY
8 Acute Radiation Syndrome
ACUTE RADIATION SYNDROME
EARLY LETHAL EFFECTS
THE PRODROMAL RADIATION SYNDROME
THE CEREBROVASCULAR SYNDROME

THE GASTROINTESTINAL SYNDROME
THE HEMATOPOIETIC SYNDROME
THE FIRST AND MOST RECENT
HEMATOPOIETIC SYNDROME

DEATHS

FROM

THE

PULMONARY SYNDROME
CUTANEOUS RADIATION INJURY
SYMPTOMS ASSOCIATED
SYNDROME

WITH

THE

ACUTE

RADIATION

TREATMENT OF RADIATION ACCIDENT VICTIMS EXPOSED TO
DOSES CLOSE TO THE LD50/60
TRIAGE
SURVIVORS OF SERIOUS RADIATION ACCIDENTS IN THE
UNITED STATES
RADIATION EMERGENCY ASSISTANCE CENTER

SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
9 Medical Countermeasures to Radiation Exposure
INTRODUCTION AND DEFINITIONS
THE DISCOVERY OF RADIOPROTECTORS
MECHANISM OF ACTION
DEVELOPMENT OF MORE EFFECTIVE COMPOUNDS
AMIFOSTINE
(WR-2721)
RADIOTHERAPY

AS

15

A

RADIOPROTECTOR

IN


AMIFOSTINE AS A PROTECTOR AGAINST RADIATION-INDUCED
CANCER
A NEW FAMILY OF AMINOTHIOL RADIOPROTECTORS
RADIATION MITIGATORS
RADIONUCLIDE ELIMINATORS
DIETARY
SUPPLEMENTS
RADIATION

AS

COUNTERMEASURES

TO

SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
10 Radiation Carcinogenesis
TISSUE
REACTIONS
STOCHASTIC EFFECTS

(DETERMINISTIC

EFFECTS)

AND

ESTIMATES

AND

CARCINOGENESIS: THE HUMAN EXPERIENCE
THE LATENT PERIOD
ASSESSING THE RISK
COMMITTEES CONCERNED
RADIATION PROTECTION

WITH

RISK

RADIATION-INDUCED CANCER IN HUMAN POPULATIONS
Leukemia
Thyroid Cancer
Breast Cancer
Lung Cancer
Bone Cancer
Skin Cancer
QUANTITATIVE RISK ESTIMATES FOR RADIATION-INDUCED
CANCER
DOSE AND DOSE-RATE EFFECTIVENESS FACTOR
SUMMARY OF RISK ESTIMATES
SECOND MALIGNANCIES IN RADIOTHERAPY PATIENTS

16


Second Cancers after Radiotherapy for Prostate Cancer
Radiation Therapy for Carcinoma of the Cervix
Second Cancers among Long-Term Survivors from Hodgkin Disease
DOSE–RESPONSE
RELATIONSHIP
CARCINOGENESIS AT HIGH DOSES

FOR

RADIATION

CANCER RISKS IN NUCLEAR INDUSTRY WORKERS
EXTRAPOLATING CANCER RISKS FROM HIGH TO LOW DOSES
MORTALITY PATTERNS IN RADIOLOGISTS
CHILDHOOD CANCER AFTER RADIATION EXPOSURE IN UTERO
NONNEOPLASTIC DISEASE AND RADIATION
SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
11 Heritable Effects of Radiation
GERM CELL PRODUCTION AND RADIATION EFFECTS ON
FERTILITY
REVIEW OF BASIC GENETICS
MUTATIONS
Mendelian
Chromosomal Changes
Multifactorial
RADIATION-INDUCED HERITABLE EFFECTS IN FRUIT FLIES
RADIATION-INDUCED HERITABLE EFFECTS IN MICE
RADIATION-INDUCED HERITABLE EFFECTS IN HUMANS
INTERNATIONAL
COMMISSION
ON
RADIOLOGICAL
PROTECTION ESTIMATES OF HEREDITARY RISKS
MUTATIONS IN THE CHILDREN OF THE A-BOMB SURVIVORS
CHANGING CONCERNS FOR RISKS
EPIGENETICS
Imprinted Genes
17

SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
12 Effects of Radiation on the Embryo and Fetus
HISTORICAL PERSPECTIVE
OVERVIEW OF RADIATION EFFECTS ON THE EMBRYO AND
FETUS
DATA FROM MICE AND RATS
Preimplantation
Organogenesis
The Fetal Period
EXPERIENCE IN HUMANS
Survivors of the A-Bomb Attacks on Hiroshima and Nagasaki Irradiated
In Utero
Exposure to Medical Radiation
COMPARISON OF HUMAN AND ANIMAL DATA
CANCER IN CHILDHOOD AFTER IRRADIATION IN UTERO
OCCUPATIONAL EXPOSURE OF WOMEN
THE PREGNANT OR POTENTIALLY PREGNANT PATIENT
SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
13 Radiation Cataractogenesis
CATARACTS OF THE OCULAR LENS
LENS OPACIFICATION IN EXPERIMENTAL ANIMALS
RADIATION CATARACTS IN HUMANS
THE LATENT PERIOD
DOSE–RESPONSE RELATIONSHIP FOR CATARACTS IN HUMANS
SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY

18


14 Radiologic Terrorism
POSSIBLE SCENARIOS FOR RADIOLOGIC TERRORISM
AVAILABILITY OF RADIOACTIVE MATERIAL
HEALTH EFFECTS OF RADIATION
EXTERNAL EXPOSURE TO RADIATION AND CONTAMINATION
WITH RADIOACTIVE MATERIALS
EXTERNAL CONTAMINATION
INTERNAL CONTAMINATION
MEDICAL MANAGEMENT
RADIOLOGIC TERRORISM

ISSUES

IN

THE

EVENT

OF

FURTHER INFORMATION
SUMMARY OF PERTINENT CONCLUSIONS
BIBLIOGRAPHY
15

Doses and Risks in Diagnostic Radiology, Interventional Radiology and

Cardiology, and Nuclear Medicine
DOSES FROM NATURAL BACKGROUND RADIATION
Cosmic Radiation
Natural Radioactivity in the Earth’s Crust
Internal Exposure
Areas of High Natural Background
COMPARISON OF RADIATION DOSES FROM NATURAL SOURCES
AND HUMAN ACTIVITIES
DIAGNOSTIC RADIOLOGY
Dose
Effective Dose
Collective Effective Dose
INTERVENTIONAL RADIOLOGY AND CARDIOLOGY
Patient Doses and Effective Doses
Dose to Personnel
19


NUCLEAR MEDICINE
Historical Perspective
Effective Dose and Collective Effective Dose
Principles in Nuclear Medicine
Positron Emission Tomography
The Therapeutic Use of Radionuclides
MEDICAL IRRADIATION OF CHILDREN AND PREGNANT WOMEN
Irradiation of Children
Irradiation of Pregnant Women
DOSES TO THE EMBRYO AND FETUS
RECOMMENDATIONS ON BREASTFEEDING INTERRUPTIONS
SUMMARY

SUMMARY OF PERTINENT CONCLUSIONS
Computed Tomography
Effective Dose and Cancer
Interventional Procedures
Nuclear Medicine
Medical Radiation of Children and Pregnant Women
Summary
BIBLIOGRAPHY
16 Radiation Protection
DE MINIMIS DOSE AND NEGLIGIBLE INDIVIDUAL DOSE
RADIATION DETRIMENT
NATIONAL COUNCIL ON RADIATION PROTECTION AND
MEASUREMENTS AND THE INTERNATIONAL COMMISSION ON
RADIOLOGICAL PROTECTION COMPARED
THE HISTORY OF THE CURRENT DOSE LIMITS
DOSE RANGES

20


SUMMARY OF PERTINENT CONCLUSIONS
GLOSSARY OF TERMS
BIBLIOGRAPHY
THE ORIGINS OF RADIATION PROTECTION
ORGANIZATIONS
QUANTITIES AND UNITS
Dose
Radiation Weighting Factor
Equivalent Dose
Effective Dose

Committed Equivalent Dose
Committed Effective Dose
Collective Equivalent Dose
Collective Effective Dose
Collective Committed Effective Dose
Summary of Quantities and Units
TISSUE REACTIONS AND STOCHASTIC EFFECTS
PRINCIPLES OF RADIATION PROTECTION
BASIS FOR EXPOSURE LIMITS
LIMITS FOR OCCUPATIONAL EXPOSURE
Stochastic Effects
Tissue Reactions (Formerly Known as Deterministic Effects)
AS LOW AS REASONABLY ACHIEVABLE
PROTECTION OF THE EMBRYO/FETUS
EMERGENCY OCCUPATIONAL EXPOSURE
EXPOSURE OF PERSONS YOUNGER THAN 18 YEARS OF AGE
EXPOSURE OF MEMBERS OF THE PUBLIC (NONOCCUPATIONAL
LIMITS)
21


EXPOSURE TO INDOOR RADON

SECTION II For Students of Radiation Oncology
17 Molecular Techniques in Radiobiology
HISTORICAL PERSPECTIVES
THE STRUCTURE OF DNA
RNA AND DNA
TRANSCRIPTION AND TRANSLATION
THE GENETIC CODE

AMINO ACIDS AND PROTEINS
RESTRICTION ENDONUCLEASES
VECTORS
Plasmids
Bacteriophage λ
Bacterial Artificial Chromosomes
Viruses
LIBRARIES
Genomic Library
cDNA Library
HOSTS
Escherichia Coli
Yeast
Mammalian Cells
DNA-MEDIATED GENE TRANSFER
AGAROSE GEL ELECTROPHORESIS
POLYMERASE CHAIN REACTION
Polymerase Chain Reaction–mediated Site-directed Mutagenesis
GENE-CLONING STRATEGIES
GENOMIC ANALYSES
22


Mapping
Contiguous Mapping
DNA Sequence Analyses
Polymorphisms or Mutations
Restriction Fragment Length Polymorphisms
Comparative Genome Hybridization
GENE KNOCKOUT STRATEGIES

Clustered Regularly Interspaced Short Palindromic Repeats and CRISPR
Associated Protein
Homologous Recombination to Knockout Genes
Knockout Mice
GENE EXPRESSION ANALYSIS
Northern Blotting
RNA Interference
Reverse Transcription Polymerase Chain Reaction
Quantitative Real-Time Polymerase Chain Reaction
Genetic Reporters
Promoter Bashing
Chromatin Immunoprecipitation
Protein–DNA Interaction Arrays (Chromatin ImmunoprecipitationChips)
Microarrays to Assay Gene Expression
RNA-Seq to Assay Gene Expression
Chromatin Immunoprecipitation-Seq
PROTEIN ANALYSIS
Western Blotting
Antibody Production
Immunoprecipitation

23


Far Western Blotting
Fluorescent Proteins
Two-Hybrid Screening
Split Luciferase Complementation Assay
Proteomics
Two-Dimensional Electrophoresis

DATABASES AND SEQUENCE ANALYSIS
SUMMARY OF PERTINENT CONCLUSIONS
GLOSSARY OF TERMS
BIBLIOGRAPHY
18 Cancer Biology
MECHANISMS OF CARCINOGENESIS
ONCOGENES
MECHANISMS OF ONCOGENE ACTIVATION
Retroviral Integration through Recombination
DNA Mutation of Regulatory Sites
Gene Amplification
Chromosome Translocation
MUTATION AND INACTIVATION OF TUMOR SUPPRESSOR GENES
The Retinoblastoma Paradigm
The Li–Fraumeni Paradigm
Familial Breast Cancer, BRCA1 and BRCA2
SOMATIC HOMOZYGOSITY
THE MULTISTEP NATURE OF CANCER
FUNCTION OF ONCOGENES AND TUMOR SUPPRESSOR GENES
Dysregulated Proliferation
Failure to Respond to Growth-Restrictive Signals
Failure to Commit Suicide (Apoptosis)
24