Table Of ContentThe Handbooks of Aging
Consisting of Three Volumes
Critical comprehensive reviews of
research knowledge, theories, concepts, and issues
Editor-in-Chief
James E. Birren
Handbook of the Biology of Aging
Edited by Edward L. Schneider and John W. Rowe
Handbook of the Psychology of Aging
Edited by James E. Birren and K. Warner Schaie
Handbook of Aging and the Social Sciences
Edited by Robert H. Binstock and Linda K. George
Handbook of
T he B i o l o gy of A g i ng
Third Edition
Editors
Edward L. Schneider and John W. Rowe
Associate Editors
Caleb E. Finch, George F. Martin,
and Edward J. Masoro
Academic Press, Inc.
Harcourt Brace [ovanovich, Publishers
San Diego New York Berkeley Boston London Sydney Tokyo Toronto
Copyright © 1990 by Academic Press, Inc.
All Rights Reserved.
No part of this publication may be reproduced or transmitted in any
form or by any means, electronic or mechanical, including photo
copy, recording, or any information storage and retrieval system,
without permission in writing from the publisher.
Academic Press, Inc.
San Diego, California 92101
United Kingdom Edition published by
Academic Press Limited
24-28 Oval Road, London NW1 7DX
Library of Congress Cataloging-in-Publication Data
Handbook of the biology of aging. -- 3rd ed. / edited by Edward L.
Schneider, John W. Rowe.
p. cm. -- (Handbook on aging)
Includes bibliographies and indexes.
ISBN 0-12-627870-9 (alk. paper)
1. Aging-Handbooks, manuals, etc. I. Schneider, Edward L.
II. Rowe, John W. (John Wallis), Date. III. Series.
[DNLM: 1. Aging. WT 104 H236]
QP86.H35 1985
574.3'72-dc20
DNLM/DLC
for Library of Congress 89-15060
CIP
Printed in the United States of America
89 90 91 92 9 8 7 6 5 4 3 2 1
Contributors
Numbers in parentheses indicate the pages on which the authors' contributions begin.
Lynne M. Ausman (384), School of Nutri David B. Danner (97), Laboratory of Mo
tion and USDA Human Nutrition Re lecular Genetics, National Institute on
search Center on Aging, Tufts Univer Aging, National Institutes of Health,
sity, Boston, Massachusetts 02111 Baltimore, Maryland 21224
Linda M. Bartoshuk (429), Department of Kenneth L. Davis (306), Department of
Surgery, Section of Otolaryngology, Psychiatry, Mt. Sinai School of Medi
Yale University School of Medicine, cine, and Alzheimer's Disease Research
New Haven, Connecticut 06510 Center, Bronx Veterans Administration
Medical Center, New York, New York
Dwight B. Brock (3), Epidemiology, De
10029
mography and Biometry Program, Na
tional Institute on Aging, National In Dariush Elahi (63), Division on Aging,
stitutes of Health, Bethesda, Maryland Harvard Medical School, and Geron
20892 tology Division, Department of Medi
cine, Beth Israel Hospital, Boston, Mas
Jacob A. Brody (3), School of Public Health,
sachusetts 02115
University of Illinois, Chicago, Illinois
60612
Andrew P. Goldberg (407), Department of
Anthony Cerami (116), Laboratory of Med Medicine, Division of Geriatrics, Johns
ical Biochemistry, The Rockefeller Uni Hopkins University, and Francis Scott
versity, New York, New York 10021 Key Medical Center, Baltimore, Mary
land 21224
Carl W. Cotman (255), Department of Psy-
chobiology, School of Biological Sci Jack M. Guralnik (3), Epidemiology, De
ences, University of California, Irvine, mography and Biometry Program, Na
California 92717 tional Institute on Aging, National In
stitutes of Health, Bethesda, Maryland
Barry J. Cusack (349), Geriatrics Section,
20892
Veterans Administration Medical Cen
ter, Boise, Idaho 83702, and Department James M. Hagberg (407), Center on Aging,
of Medicine, University of Washington, University of Maryland, College Park,
Seattle, Washington 98195 Maryland 20742, and Department of
xi
xii Contributors
Medicine, Division of Geriatrics, Johns Thomas H. Norwood (131), Department of
Hopkins University, and Francis Scott Pathology, University of Washington,
Key Medical Center, Baltimore, Mary Seattle, Washington 98195
land 21224
Gary S. Richardson (275), Endocrine Divi
Nikki J. Holbrook (97), Laboratory of Mo sion, Brigham and Women's Hospital,
lecular Genetics, National Institute on Boston, Massachusetts 02115
Aging, National Institutes of Health,
John W. Rowe (63), Division on Aging, De
Baltimore, Maryland 21224
partment of Medicine, Harvard Medical
1
Thomas B. Horvath (306), Department of School, Boston, Massachusetts 02115
Psychiatry, Mt. Sinai School of Medi
Robert M. Russell (384), Human Nutrition
cine, and Department of Psychiatry,
Research Center on Aging, School of
Bronx Veterans Administration Medical
Medicine, School of Nutrition, Tufts
Center, New York, New York 10029
University, Boston, Massachusetts
Thomas E. Johnson (45), Department of 02111
Psychology and Institute for Behavioral
Robert M. Sapolsky (330), Department of
Genetics, University of Colorado, Boul
Biological Sciences, Stanford Univer
der, Colorado 80309
sity, Stanford, California 94305
Edward G. Lakatta (181), Laboratory of
James R. Smith (131), Department of Vi
Cardiovascular Science, Gerontology
rology and Epidemiology, Baylor Col
Research Center, National Institute on
lege of Medicine, Houston, Texas 77030
Aging, National Institutes of Health;
Department of Medicine, Johns Joan Smith-Sonneborn (24), Program in
Hopkins Hospital; and Department of Aging and Human Development, Zoolo
Physiology, University of Maryland gy and Physiology Department, Univer
School of Medicine, Baltimore, Mary sity of Wyoming, Laramie, Wyoming
land 21224 82071
Annette T. Lee (116), Laboratory of Medi Gretchen H. Stein (131), Department of
cal Biochemistry, The Rockefeller Uni Molecular, Cellular, and Developmen
versity, New York, New York 10021 tal Biology, University of Colorado,
Boulder, Colorado 80309
Edward J. Masoro (72), Department of
Physiology, The University of Texas Robert E. Vestal (349), Clinical Pharma
Health Science Center, San Antonio, cology and Gerontology Research Unit,
Texas 78284 Veterans Administration Medical Cen
ter, Boise, Idaho 83702, and Depart
Patrick C. May (219), Department of
ments of Medicine and Pharmacology,
Gerontology, University of Southern
University of Washington, Seattle,
California, Los Angeles, California Washington 98195
90089
San Y. Wang (63), Division on Aging, Har
Richard A. Miller (157), Department of Pa
vard Medical School, and Gerontology
thology, Boston University School of
Division, Department of Medicine, Beth
Medicine, Boston, Massachusetts 02118
Israel Hospital, Boston, Massachusetts
David G. Morgan (219), Departments of 02115
Gerontology and Biological Sciences,
University of Southern California, Los Present address: Mount Sinai School of Medicine,
Angeles, California 90089 Mount Sinai Hospital, New York, New York 10029
Contributors xiii
James M. Weiffenbach (429), Clinical In National Institutes of Health, Bethesda,
vestigation and Patient Care Branch, Maryland 20892
National Institute of Dental Research,
Foreword
The present volume is one of three hand entific personnel, graduate students, and
books now in their third edition: Hand professionals will find the volumes useful.
book of the Biology of Aging, Handbook of The availability of the information in the
the Psychology of Aging, and Handbook of convenient form of the handbooks also
Aging and the Social Sciences. Because of may stimulate new courses of instruction
the growth in research on aging, there has and seminars on aging as well as provide
been an accelerated need to collate and easy access to the research literature.
interpret existing information. Thus, the The series editor wishes to thank the edi
decision was made to accelerate the pub tors of the individual volumes, Robert H.
lication of the third edition of these hand Binstock, Linda K. George, John W. Rowe,
books to reflect this expansion in research. K. Warner Schaie, Edward L. Schneider, and
The growth of the National Institute on the associate editors, Caleb E. Finch, Mar
Aging and sponsorship of research by the garet Gatz, Victor W. Marshall, George M.
National Institute of Mental Health and Martin, Edward J. Masoro, George C.
the Administration on Aging has stimu Myers, Timothy A. Salthouse, Carmi
lated needed research. Phenomena of Schooler, and James H. Schulz. The series
aging cut across many scientific fields. It is editor is grateful to these editors for their
impossible to represent all the material cooperation in the successful completion
that is relevant in one volume. We focus of this publication adventure. Thanks are
on three major sources of influence on owed to Donna E. Deutchman for develop
aging: the biological, the psychological, ing the plan for the third edition and for
and the social. Perhaps in a strict sense, coordinating the relationship with the
one should view aging as an ecological publisher.
phenomenon dependent upon the influ There is little doubt from the reading of
ences of genetics, physical and social en these volumes that the subject matter of
vironments, and individual behavior. aging has become more sophisticated and
It is hoped that these volumes will be also mainstream in many scientific disci
consulted across scientific areas to trace plines. It is hoped that the handbooks'
some of the pathways of aging through the publication will motivate continued at
matrix of scientific information and disci tention to research on aging and the well-
plinary orientation. It is expected that sci being of the elderly in our society.
James E. Birren
XV
Preface
The third edition of the Handbook of the in the biology of aging. In order to keep this
Biology of Aging continues the tradition of volume to a reasonable length, we have
providing a general overview to a wide sci omitted covering certain areas. However,
entific audience of some of the most im we have tried to focus on those subjects
portant topics in biomedical gerontology. which have either had the greatest pro
This third edition is intended to both up gress or which are of the widest interest to
date and extend the two previous editions the readership.
published in 1978 and 1985. There are new We wish to extend special thanks to the
sections on methodology for biological Associate Editors, Caleb E. Finch, George
aging studies and on animal models to F. Martin, and Edward J. Masoro, for their
help scientists entering this emerging extensive reviews and revisions of the
field. In addition, there are new chapters chapters.
on protein modifications with aging, spe As with previous editions, we would
cial senses, circadian rhythms, and the like to express our appreciation to our col
adrenocortical axis. The remaining chap leagues who served as outside reviewers:
ters are extensive updates and extensions S. Allen, V. Monnier, K. Aufderheide, R.
of specific chapters from the second edi Russell, G. Burmer, R. Sprott, E. Dekker,
tion. Since knowledge in biomedicine has K. Swisshelm, P. Garry, N. Talal, J. John
a doubling rate of approximately three son, and J. Thompson.
years, it is timely to review these key areas
Edward L. Schneider
John W. Rowe
xvii
One
Demography and Epidemiology of Aging
in the United States
Dwight B. Brock, Jack M. Guralnik, and Jacob A. Brody
I. Introduction projections of 12 million and 3.7% for the
85 and older group (Table II). These projec
One of the remarkable demographic devel tions are based on the assumption that
opments of the twentieth century in the mortality rates will not continue the rapid
United States has been the aging of the declines experienced almost continually
population. In 1900 there were 3.1 million since 1968, but will decline more modes
Americans age 65 and older, representing tly in the future. In an alternative set of
approximately 4% of the total population. projections, based on the assumption that
By 1986, according to Census Bureau esti mortality will continue to decline at the
mates, that segment of the population had same rate as in recent years (about 2%
grown to 29.2 million persons, or 12.1 % of annually), Guralnik, Yanagishita, and
the total population (Table I). Such in Schneider (1988) estimate that the 65 and
creases in the older population are un older segment will total some 87 million
precedented in the history of the United persons (27% of the total) and the 85 and
States, and most population projections older group will number almost 24 million
for the twenty-first century indicate a con (7.3%). Given that mortality has decreased
tinuation of that growth until at least the about 1.3% per year since the beginning of
middle of that century. Furthermore, this century—which included a period of
those 85 and older, termed by some the 14 years (1954-1968) in which there was a
"oldest old/' will continue to be the most slight increase in mortality among men—
rapidly growing segment of the popula the 2% decline assumption is not unre
tion, both in terms of their actual numbers alistic when one considers the possibility
and as a percentage of the total population. of future medical breakthroughs in such
The Social Security Administration areas as heart disease, cancer, and other
(SSA, 1988) projects that the older popula life-threatening chronic conditions. There
tion will number more than 69 million is, therefore, uncertainty in creating popu
persons and represent almost 22% of the lation projections, but what is clear from
population by 2040, with corresponding the range of projections which have been
Handbook of the Biology of Aging, Third Edition
Copyright © 1990 by Academic Press, Inc. All rights of reproduction in any form reserved.
3
4 Brock, Guralnik, and Brody
Table I
Estimates of the Population 65 Years and Older and Percentages by Race, Sex, and Age:
0
1986 (Numbers in Thousands)
Percentage of Population
Percentage of Total Population 65 and Older
TToottaall PPooppuullaattiioonn
PPooppuullaattiioonn 6655 aanndd 65 and 85 and 85 and
All Ages Older Older 65-74 75-84 Older 65-74 75-84 Older
All races 241,096 29,168 12.1 7.2 3.8 1.2 59.4 31.1 9.5
Male 117,365 11,819 10.1 6.5 2.9 0.7 64.7 28.7 6.7
Female 123,730 17,349 14.0 7.8 4.6 1.6 55.8 32.7 11.5
White 204,312 26,292 12.9 7.6 4.0 1.2 59.1 31.3 9.6
Male 99,810 10,644 10.7 6.9 3.1 0.7 64.6 28.8 6.6
Female 104,501 15,648 15.0 8.3 4.9 1.8 55.3 33.0 11.7
All other 36,784 2,875 7.8 4.9 2.3 0.7 62.5 28.9 8.5
Male 17,555 1,175 6.7 4.4 1.9 0.5 65.6 27.6 6.8
Female 19,229 1,700 8.8 5.3 2.6 0.9 60.4 29.9 9.7
Source: U.S. Bureau of the Census, Current Population Reports, Series P-25, No. 1000, 1987b.
a
Figures may not add to totals because of rounding.
made is that there will be a substantial number of very old persons does not in
number of very old and very vulnerable crease, and the period of illness and dis
persons in our society. ability of those persons decreases, leading
In terms of the general health and well- to a reduced period of need for medical care
being of older people, there has been con in late life. Others (Brody, 1985; Gruen-
siderable debate about what the future berg, 1977; Kramer, 1980; Manton, 1982;
holds. Fries (1980) has stated a theory of Schneider & Brody, 1983; Schneider &
compression of morbidity in which the Guralnik, 1987) have argued that the fu-
Table II
Population Projections by Age and Sex for the Years 2000, 2020, and 2040
0
(Alternative II Populations in Thousands)
2000 2020 2040
Percentage Percentage Percentage
of Total of Total of Total
Age and Sex Population Population Population Population Population Population
Both sexes
65 and older 35,480 12.8 52,026 16.9 69,650 21.7
85 and older 4,534 1.6 6,426 2.1 11,753 3.7
Male
65 and older 14,598 10.7 22,460 14.8 29,891 19.0
85 and older 1,263 0.9 1,896 1.3 3,724 2.4
Female
65 and older 20,882 14.9 29,566 19.0 39,759 24.3
85 and older 3,271 2.3 4,530 2.9 8,028 4.9
Source: Social Security Area Projections, Actuarial Study No. 102, 1988.
a
Alternative Π projections are prepared using the "middle" series of assumptions regarding future fertility, migration, and
mortality patterns.
