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THE BIOLOGY AND
TREATMENT OF
CANCER
 THE BIOLOGY AND
TREATMENT OF
CANCER
Understanding Cancer

Arthur B. Pardee
Department of Biological Chemistry
Harvard University
Dana-Farber Cancer Institute

Gary S. Stein
Department of Cell Biology
University of Massachusetts Medical School
and UMASS Memorial Cancer Center

A JOHN WILEY & SONS, INC., PUBLICATION

Copyright # 2009 by John Wiley & Sons, Inc. All rights reserved.

Published by John Wiley & Sons, Inc., Hoboken, New Jersey

Published simultaneously in Canada

Wiley-Blackwell is an imprint of John Wiley & Sons, formed by the merger of Wiley’s global Scientific,
Technical, and Medical business with Blackwell Publishing.

No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by
any means, electronic, mechanical, photocopying, recording, scanning, or otherwise, except as permitted under.
Section 107 or 108 of the 1976 United States Copyright Act, without either the prior written permission of
the Publisher, or authorization through payment of the appropriate per-copy fee to me Copyright Clearance
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http: //www. wiley.com/go/permission.

Limit of Liability/Disclaimer of Warranty: While the publisher and author have used their best efforts in preparing
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Library of Congress Cataloging-in-Publication Data:

Pardee, Arthur B. (Arthur Beck), 1921–

The biology and treatment of cancer : understanding cancer / Arthur B.
Pardee, Gary S. Stein.
p. cm.
ISBN 978-0-470-00958-1 (pbk.)
1. Cancer – Popular works. 2. Cancer – Treatment – Popular works. I.
Stein, Gary S. II. Title.
RC263.P276 2006
616.990 4 – dc22
2008001368

Printed in the United States of America

10 9 8 7 6 5 4 3 2 1
 CONTENTS

Contributors vii

Preface ix

PART I: INTRODUCTION

1 WHAT GOES WRONG IN CANCER 3

Arthur B. Pardee, Gary S. Stein, and Elizabeth A. Bronstein

PART II: CLINICAL PERSPECTIVE

2 CANCER AS A DISEASE: TYPES OF TUMORS, THEIR

FREQUENCIES, AND THEIR PROGRESSION 23
Kenneth J. Pienta
3 ENVIRONMENTAL, GENETIC, AND VIRAL
CAUSES OF CANCER 35
Rami I. Aqeilan, Nicola Zanesi, and Carlo M. Croce
4 DIAGNOSIS AND TREATMENT OF MALIGNANT DISEASES 57
F. Marc Stewart and Jessica A. Stewart
5 CLINICAL CHALLENGES FOR TREATMENT AND A CURE 83
Alan Rosmarin

PART III: CANCER BIOLOGY

6 UNDERSTANDING THE BIOLOGY OF CANCER 103

Laura A. Lambert, Donald H. Lambert, and Khandan Keyomarsi
7 MUTATIONS AND CELL DEFENSES 123
Otto S. Gildemeister, Jay M. Sage, and Kendall L. Knight

PART IV: CANCER DIAGNOSIS AND TREATMENT

8 CANCER DETECTION AND BIOMARKERS 145

Arthur B. Pardee and Peng Liang

v
vi CONTENTS

9 CLINICAL CHALLENGES FOR TREATMENT AND A CURE 155

Eleni Efstathiou and Christopher J. Logothetis
10 CLINICAL TRIALS IN ONCOLOGY 169
Konstantin H. Dragnev and Mark A. Israel
11 THE DEVELOPMENT OF DRUGS: CURRENT
CONCEPTS AND ISSUES 183
Barry S. Komm and Christopher P. Miller
12 A NEW GENERATION OF DRUGS IN CANCER
TREATMENT: MOLECULARLY TARGETED THERAPIES 193
Thomas O0 Hare, Christopher A. Eide, and Michael W. Deininger
13 EPIDEMIOLOGY: IDENTIFYING CANCER’S CAUSES 223
James R. Hebert
14 CONSUMER HEALTH INFORMATION 257
David Shepro and Catherine N. Norton

Index 267
 PREFACE

We and our families and friends, like too many others, have experienced cancers of
various kinds. Some patients recovered, but sadly, many died. From these trying
events and from our professional studies as cancer researchers, we decided that a
source which briefly summarizes our current knowledge about cancer and its treat-
ment in nontechnical language would be useful. We hope to provide explanations of
cancer treatment and biology that are meaningful to cancer patients and their
families. It is intended for an informed audience who are interested in learning about
cancer and who are not specialists. We acknowledge a forerunner with this theme,
John Cairns’ Cancer Science and Society, published in 1978 by W.H. Freeman and
Company, San Francisco.
This book is developed to explain the nature of the disease, current options for
treatment, and emerging strategies for cancer detection and therapy, as provided
from clinical experience and by spectacular advances in our understanding of
cells, genes, and molecular biology. Insights have been gained into fundamental
regulatory mechanisms that are faulty in cancers and that reveal prospects for new
treatments.
The presentations are not encyclopedic. We cannot do better than to paraphrase
the preface of The Evolution of Physics (1938) by Albert Einstein and Leopold
Infeld, a book on an even more difficult subject. ‘‘We have not a written a textbook
of physics. Our intention was rather to sketch in broad outline the attempts of the
human mind to find connections. But our presentation had to be simple. Through
the maze of facts and concepts we had to choose some highways. Some essential
lines of thought have been left out because they do not lie along the road we have
chosen.’’

Arthur B. Pardee
Gary S. Stein

ix
 CONTRIBUTORS

Rami I. Aqeilan, Department of Molecular Virology, Immunology, and Medical Genetics,

Division of Human Cancer Genetics and Comprehensive Cancer Care, Ohio State
University, Columbus, Ohio
Elizabeth A. Bronstein, UMASS Memorial Cancer Center, University of Massachusetts
Medical School, Worcester, Massachusetts
Carlo M. Croce, Department of Molecular Virology, Immunology, and Medical Genet-
ics, Division of Human Cancer Genetics and Comprehensive Cancer Care, Ohio State
University, Columbus, Ohio
Michael W. Deininger, Oregon Health & Science University, Center for Hematologic
Malignancies, Portland, Oregon
Konstantin H. Dragnev, Norris Cotton Cancer Center, Darthmouth-Hitchcock Medical
Center, Lebanon, New Hampshire
Eleni Efstathiou, Department of Genitourinary Medical Oncology, M.D. Anderson
Cancer Center, University of Texas, Houston, Texas
Christopher A. Eide, Howard Hughes Medical Institute, Oregon Health & Science
University, Cancer Institute, Portland, Oregon
Otto S. Gildemeister, Department of Biochemistry and Molecular Parmacology,
University of Massachusetts Medical School, Worcester, Massachusetts
James R. Hebert, Department of Epidemiology and Biostatistics, Arnold School of Public
Health, South Carolina Statewide Cancer Prevention and Control Program, University
of South Carolina, Columbia, South Carolina
Mark A. Israel, Norris Cotton Cancer Center, Darthmouth–Hitchcock Medical Center,
Lebanon, New Hampshere
Khandan Keyomarsi, Department of Experimental Radiation Oncology, M.D. Anderson
Cancer Center, University of Texas, Houston, Texas
Kendall L. Knight, Department of Biochemistry and Molecular Pharmacology, Univer-
sity of Massachusetts Medical School, Worcester, Massachusetts
Barry S. Komm, Wyeth Research, Collegeville, Pennsylvania
Donald H. Lambert, Department of Anesthesia, Boston University Medical School,
Boston Medical Center, Boston, Massachusetts
Laura A. Lambert, Department of Surgical Oncology, M.D. Anderson Cancer Center,
University of Texas, Houston, Texas.

vii
viii CONTRIBUTORS

Peng Liang, Vanderbilt–Ingram Cancer Center, Vanderbilt University, Nashville, Tennessee

Christopher J. Logothetis, Department of Genitourinary Medical Oncology, M.D.
Anderson Cancer Center, University of Texas, Houston, Texas
Christopher P. Miller, Radius Health, Cambridge, Massachusetts
Catherine N. Norton, MBLWHO1 Library, Marine Biological Laboratory, Woods Hole,
Massachusetts
Thomas O’Hare, Howard Hughes Medical Institute, Oregon Health & Science University
Cancer Institute, Portland, Oregon
Arthur B. Pardee, Harvard Medical School, Dana–Farber Cancer Institute, 44 Binney
Street, Boston, Massachusetts
Kenneth J. Pienta, Internal Medicine and Urology Departments, University of Michigan
Comprehensive Cancer Center, Ann Arbor, Michigan
Alan Rosmarin, Division of Hematology/Oncology and UMASS Memorial Cancer
Center, University of Massachusetts Medical School, Worcester, Massachusetts
Jay M. Sage, Department of Biochemistry and Molecular Pharmacology, University of
Massachusetts Medical School, Worcester, Massachusetts
David Shepro, Biology Department, Boston University, Boston, Massachusetts and
Marine Biological Laboratory, Woods Hole, Massachusetts
Gary S. Stein, UMASS Memorial Cancer Center, University of Massachusetts Medical
School, Worcester, Massachusetts
F. Marc Stewart, Seattle Cancer Care Alliance, Fred Hutchinson Cancer Research Center,
University of Washington, Seattle, Washington
Jessica A. Stewart, Seattle Cancer Care Alliance, Fred Hutchinson Cancer Research
Center, University of Washington, Seattle, Washington
Nicola Zanesi, Department of Molecular Virology, Immunology, and Medical Genetics,
Division of Human Caner Genetics and Comprehensive Cancer Care, Ohio State
University, Columbus, Ohio
Figure 1.1 Cancer-related chromosomal aberrations. In early stages of cancer, chromosomes
break and join with segments of other chromosomes that are not adjacent in normal cells
(chromosomal translocations). As a consequence, genes that control cell growth and specialized
properties of cells are frequently rearranged. Other cancer-related alterations that result from
reorganization are in cell adhesion and motility as well as in capacity to invade and grow in
tissues at distant sites from the initial tumor (metastasis). (A) normal chromosomes (note that
each chromosome is a single color); (B) chromosomes in cancer cells that contain fused segments
of multiple chromosomes (note multicolored chromosomes).
 Transmembrane
Receptor

Chromosomes

Nuclear
Cell Membrane
Membrane

Nucleoli Extracellular
Signal
Molecules

Endoplasmic
Mitochondria Reticulum
with Ribosomes

(A)

(B)

Figure 1.2a,b Structure and organization of regulatory machinery in (A) normal and (B) cancer
cells. The organization and location of machinery that controls genes is modified during the
onset and progression of cancer. (See text for full description).
 EXTRACELLULAR
ENVIRONMENT

Extracellular
Transmembrane Signal Molecule
Receptor

Cell Membrane
Scaffold
Protein Relay and
Adapter
Proteins

CYTOPLASM Intracellular
Signaling
Proteins

Nuclear Membrane

Nuclear Pores

NUCLEUS
Target Protein

Gene (DNA) Signal Response Element

Figure 1.3 Cell signaling. Cells communicate and respond to the extracellular environment
through a process designated signal transduction. Signal molecules bind to transmembrane
receptors that span the cell membrane. The interaction of signal molecules with components of
receptors located outside the cell modifies the intracellular components of the receptors. An
environmental signal is thereby transduced into a cascade of regulatory steps that control genes
which control cell proliferation and specialized properties of cells.
In some signaling pathways, scaffold proteins assemble signaling molecules into complexes for
the initial passage of information from the transmembrane receptor to relay and adaptor
proteins. Subsequent steps in the signaling process amplify and integrate signals. A chain of
intracellular signaling proteins processes regulatory information through the cytoplasm and into
the cell nucleus to activate or suppress genes. In other signaling pathways the regulatory cascades
are abbreviated. The transduction of regulatory information from the intracellular component of
the transmembrane receptor is more direct, circumventing intermediary steps in information
transfer. At an early stage in the signaling process a signaling protein enters the nucleus and
interacts directly with genes to modify expression.
Many cancer cells exhibit defects in one or more steps of signaling cascades that alter control of
cell growth, specialized cell properties, cell–cell communication, cell motility, and cell adhesion.
The components of signaling pathways that are modified in tumor cells are targets for treatments
that are effective and specific.
 Differentiation
Cell Specialization
G0

Metaphase
Checkpoint G1

M
G1
Cyclins
cycD/
G2 CDK4,6
Checkpoint
G1
M Cyclins R Point R Checkpoint
cycB/ Cyclins
CDK1 cycE/
G2 CDK2

S
Cyclins
cycA/
CDK1

(A)

PROPHASE METAPHASE ANAPHASE TELOPHASE

(B)

Figure 1.4a,b (A) The stages of the cell cycle. G1 is the period following cell division (M; mitosis)
and precedes the S phase, the period when genes (DNA) are duplicated to provide an identical set
of genes for progeny cells. (See text for full description).

Figure 2.3 Summary of the steps necessary for successful metastasis. Cancer cells must break
down the tissue surrounding them, invade their environment, and attract a blood supply. (See
text for full description).
Figure 2.4 Formation of tumor. As the normal cells mutate and divide, they start to pile up
(hyperplasia): (A) artists drawing; (B) actual pictures of this occurring in the breast. Over time,
these cells develop added mutations and begin to look abnormal (atypical dysplasia). If this
process occurs in an organ with ducts, such as the breast, they fill in the normal breast ducts
before they come invasive (ductal carcinoma in situ). Overall, this process is thought to take years
to occur. [(A) From ref. 40.]

Mutagens and Carcinogens

(Radiations, chemicals, biological
agents)
n , G A
tio tion n hy ene ctiv
t iva uta tio po tic ati
ac m la m m on
In tic ethy et ut
e hy at
en rm la ion
G ype tio ,
h n

Tumor Suppressor Oncogenes

p53, Rb,Fhit, Wwox CANCER Ras, Myc, Tcl1
STOP! GO!

miR 15/16 miR 155

MicroRNA

Figure 3.1 Schematic representation showing the complex relationships among genetics, epi-
genetics, and environmental factors (radiations, chemicals, and microorganisms) in causing
cancer. Oncogenes and tumor suppressors are protein-coding genes, whereas microRNA genes
are noncoding genes (see the text).
 (A) Tumor
TSG
Gene Therapy
KO
G- Mutant Cured
Healthy TS

On Tumor
Wild type co anti-oncogenic
-Tg
Therapy
Mutant Cured

(B) (C)
TS1-KO TS-KO

TS1,2-KO KO-Tg
Healthy Healthy

TS2-KO Tumor Onco-Tg Tumor in 6 months

Healthy Tumor in 1 year

Figure 3.3 Examples of cancer genetics experiments using engineered mice. (See text for full
description).

Figure 4.2a–d (A) Surgeon operates from a console located adjacent to the patient and the
operative field. (B,C) Robotic arms are used to transect the tumor. Hand controls offer stability
and precision in movement. (D) Simulated surgical approach to prostate cancer: robotic arms
allow careful separation of nerves away from site of tumor/prostate resection, preserving urinary
and sexual function. (Images C 2008 Intuitive Surgical, Inc.)
Figure 4.5 Comparison of three-dimensional conformal radiation therapy with intensity-
modulated radiation therapy for prostate cancer (red). IMRT utilizes greater number of beams
(yellow) tailored with various shapes and intensities to deliver more radiation to the tumor and
less to the normal tissues.

Figure 4.7a–c Brachytherapy for breast cancer. (A) Special balloon and catheter prior to
insertion into tumor site following lumpectomy. (B,C) Radioactive seeds introduced into balloon
through the catheter and removed after various time intervals. Extent of radiation effects show
by concentric circles. (Image 7A used with permission of The University of Toledo Medical Center.
Images 7B and 7C courtesy of Hologic.)
Figure 4.8 Microarray analysis used to detect abnormal expression of ErbB2. Flurorescent-
labeled cDNA for the ErbB2 gene combines specifically with a patient’s mRNA containing ErbB2.
If a patient with breast cancer (right) has a large amount of ErbB2 in her cells, it will appear as a
bright fluorescent spot on the array plate.

Figure 4.9 Comparison of photon delivery by linear accelerator and proton therapy by cyclotron
to a brain tumor. (A) Photon radiation passes through normal tissue, tumor (center, circular
structure), and more normal tissue. (B) Proton beam released in the tumor stops at the far edge if
the tumor, sparing normal tissue.
 Figure 4.12 Philadelphia chromosome translocation. (See text for full description).

Figure 4.15a,b (A) Colon cancer cell: extracellular growth factor (ECGF) stimulates mutated
receptor on cell. The tyrosine kinase enzyme (blue) located on the inner aspect of the receptor is
mutated. (See text for full description).
Figure 4.16 Colon cancer cell: ECGF is the normal growth factor stimulating normal cell growth.
Colon cancer cells that express abnormal amounts of the ECGFR may be blocked by an antibody
designed to attach specifically to the receptor. This antibody prevents stimulation by ECGF and
reduces the ability of the cell to divide and grow.

= antibody = 131I radioactive isotope = radiation

131
Figure 4.17 Radioimmunotherapy. I-labeled antibodies (radioactive antibodies) attach to
surface antigens on tumor cells. Radiation crossfires to other cells.
 Receptor
Extracellular
Matrix

Mitochondrion Peroxisome

Plasma
membrane
ER
Nucleus

Cytoplasm

Cytoskeleton

Lysosome Golgi
Extracellular

Channel

Figure 6.1 Overview of a cell. The most common structures of a cell, including a variety of
organelles (i.e., mitochodria, peroxisome, Golgi apparatus). (See text for full description).

Figure 6.9 Protection of normal cells against the toxic affect of chemotherapy. (See text for full
description).
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