COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY

VOLUME XXII

COLD S P R I N G H A R B O R S Y M P O S I A

ON Q U A N T I T A T I VE BIOLOG Y

Founded in 1933

REGINALD G. HARRIS

Director of the Biological Laboratory

1924 to 1936

The Symposia were organized and managed by Dr. Harris until his death. Their continued use- fulness is a tribute to the soundness of his vision.

The Symposium volumes

are published by the Long Island Biological Association

as a part of the work of the Biological Laboratory

Cold Spring Harbor, L. I., New York

COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY

VOLUME XXII

Population Studies: Animal Ecology and Demography

THE BIOLOGICAL LABORATORY

COLD SPRING HARBOR, L. I., NEW YORK

1957

COPYRIGHT 1958 BY

THE BIOLOGICAL LABORATORY

LONG ISLAND BIOLOGICAL ASSOCIATION, INC.

All rights reserved, This book may not be reproduced

in whole or in part except by reviewers for the public

press without written permission from the publisher.

P R I N T E D BY

WAVERLY PRESS, INC., BALTiMOrE, MARYL~D, U. S. A.

Foreword The purpose of this year's Symposium was to bring together in the informal

setting of the Biological Laboratory ecologists, demographers, anthropologists, population geneticists, and mathematicians to discuss research problems, experi- mental procedures, and analytical techniques of common interest to the fields of ecology and demography. Following an introductory survey by Professor LaMont C. Cole of Cornell University, daily sessions were devoted to a variety of problems. In order that problems confronting demographers would be seen in proper perspec- tive, Professors R. J. Braidwood of Chicago University, F. S. Hulse of the Uni- versity of Washington, J. B. Birdsell of the University of California, and G. T. Trewartha of the University of Wisconsin discussed the historical aspects of human populations.

The presentation of demographic theory was made by Drs. A. J. CoMe of Prince- ton University, N. Ryder of the University of Wisconsin, J. Hajnal of the London School of Economics, and L. Henry of the National Institute of Demographic Studies in Paris. It was generally agreed that demographic models, although capable of short-range predictions, are insufficient to cope with gross changes in human attitudes or behavior. A number of persons discussed mathematical models of the growth of animal populations and experimental tests of these models. In the case of simple populations, such as those of bacteria described by Dr. H. Moser of the Biological Laboratory, the mathematical models and population growth agree. In the case of more complex organisms, the agreement is far from perfect, although some features of the changes in population size can be predicted. Simple logistic curves are suitable in certain instances; in more complicated situations populations may oscillate according to a number of different types of patterns.

The closing days were devoted to discussions of communities of human and nonhuman populations--the association of diverse types of individuals or, on the subhuman level, species into groups of fairly constant patterns. The effect of these groupings on the evolution of the interbreeding populations was discussed by Drs. Th. Dobzhansky of Columbia University, R. C. Lewontin of North Carolina State College, and H. Shapiro of the American Museum of Natural History. The Symposium closed with a concluding survey by Professor G. E. Hutchinson of Yale University, who showed how some of the newer mathematical concepts are useful in clarifying the statement of problems that are under investigation. On the other hand, he pointed out that demography is faced with the problem of anticipating decisions made by individual human beings. Although these decisions can be antici- pated in part, there is no technique by which a person can be prevented from chang- ing his mind, and consequently altering his behavior.

The meetings were held from June 3 to June 12, 1957, and were attended by about 150 scientists. Approximately 40 of the participants were from overseas. The Biological Laboratory was able to finance the conference with funds obtained from the Population Council, Inc., the Milbank Memorial Fund, the U. S. Atomic Energy Commission, the U. S. Public Health Service, the National Science Founda- tion, and the Carnegie Corporation of New York. I wish to acknowledge this sup- port and express our gratitude to the organizations that provided it. I wish also to record our appreciation and thanks to the committee who planned the program: L. C. Birch (chairman), Th. Dobzhansky, Frank Lorimer, and Bruce Wallace.

M. DEMEREC

Executive Editor: KATHERINE BREHME WARREN, Associate Professor of Biology, Adelphi College, Garden City, New York

V

LIST OF PREVIOUS VOLUMES

Volume Volume Volume Volume Volume Volume Volume Volume Volume Volume

I (1933) Surface Phenomena, 239 pp. II (1934) Aspects of Growth, 284 pp. III (1935) Photochemical Reactions, 359 pp. IV (1936) Excitation Phenomena, 376 pp. V (1937) Internal Secretions, 433 pp. VI (1938) Protein Chemistry, 395 pp. VII (1939) Biological Oxidations, 463 pp. VIII (1940) Permeability and the Nature of Cell Membranes, 285 pp. IX (1941) Genes and Chromosomes: Structure and Organization, 315 pp. X (1942) The Relation of Hormones to Development, 167 pp.

Volume XI (1946) Heredity and Variation in Microorganisms, 314 pp. Volume XII (1947) Nucleic Acids and Nucleoproteins, 279 pp. Volume XIII (1948) Biological Applications of Tracer Elements, 222 pp. Volume Volume Volume Volume Volume Volume

Volume

Volume

XIV (1949) Amino Acids and Proteins, 217 pp. XV (1950) Origin and Evolution of Man, 425 pp. XVI (1951) Genes and Mutations, 521 pp. XVII (1952) The Neuron, 323 pp. XVIII (1953) Viruses, 301 pp. XIX (1954) The Mammalian Fetus: Physiological Aspects of Development,

225 pp. XX (1955) Population Genetics: The Nature and Causes of Genetic Varia-

bility in Populations, 346 pp. XXI (1956) Genetic Mechanisms: Structure and Function, 393 pp.

vi

SOME SYMPOSIUM PARTICIPANTS

T o p row: H. N. Kluyver, L. Henry, J. Sutter (background, L. Sanghvi); A. J. Nicholson, L. C. Birch; M. Demerec, Dr. and Mrs. B. Colombo.

Second row: C. G. Sibley, L. C. Cole, P. J. Wangersky; F. W. Notestein, J. Hajnal; H. Ozeki, L. Cavalli-Sforza, S. Utida.

T h i r d row: Dr. and Mrs. W. F. Blair, C. H. Waddington, R. C. Lewontin; H. L. Shapiro, C. A. Reed; N. Dobzhansky, E. L. Scott, J. Yerushalmy, J. Neyman, A. Polnik.

Bot tom row: J. N. Spuhler, R. J. Braidwood; C. Malogolowkin, B. Wallace, Th. Dobzhansky; C. A. Reed, L. C. Dunn.

SOME SYMPOSIUM PARTICIPANTS

Top row: H. N. Kluyver, M. Udvardy; A. D. Pickett, H. G. Andrewartha; L. Slobodkin, A. J. Nicholson, D. Chitty; F. Lorimer, M. Demerec, F. Osborn.

Second row: M. Demerec, K. Maramorosch; A. Milne; L. Cavalli-Sforza, J. B. Birdsell. T h i r d row: J. A. Moore, J. B. Birdsell; Th. Dobzhansky, A. Buzzati-Traverso; C. C. Lindegren, A. P. James. Bot tom row: F. Lorimer, J. Hajnal; D. Kirk, G. Trewartha; G. E. Hutchinson, J. Neyman.

(The photographs were taken by Drs. L. C. Cole, K. Maramorosch, A. Sokoloff, and K. B. Warren. The art work was done by Mr. Harry Jones.)

List of those attending the Symposium

ANDREWARTHA, H. G., University of Adelaide, Adelaide, Australia BEARDMORE, JOHN, Columbia University, New York (University of Sheffield,

Sheffield, England) BERGER, AGNES, Columbia University, New York BIRCH, L. C., The University of Sydney, New South Wales, Australia BIRDSELL, JOSEPH B., University of California, Los Angeles, California BLAIR, W. FRANK, University of Tesas, Austin, Texas BLAIR, MRS. W. FRANK, University of Texas, Austin, Texas BOYES, J. WALLACE, McGill University, Montrel, Canada BRAIDWOOD, ROBERT J., University of Chicago, Chicago, Illinois BRAIDWOOD, MRS. ROBERT J., University of Chicago, Chicago, Illinois BRUCK, DAVID, Columbia University, New York BUZZATI-TRAvERso, ADRIANO, University of California, La Jolla, California CALHOUN, EDITH, Kensington, Maryland CALHOUN, JOHN B., National Institute of Mental Health, Bethesda, Maryland CARTER, T. C., MRC. Radiobiological Research Unit, Berkshire, England CASPARI, ERNST, Wesleyan University, Middletown, Connecticut CAVALLI-SFORZA, L. L., University of Parma, Milan, Italy CHITTY, DENNIS, Oxford University, Oxford, England CHITTY, HELEN, Oxford University, Oxford, England CHRISTIAN, JOHN, Johns Hopkins University, Bethesda, Maryland COALE, ANSLEY J., Princeton University, Princeton, New Jersey COLE, LAMONT C., Cornell University, Ithaca, New York COLOMBO, BERNARDO, Instituto Universitario di Venezia, Venice, Italy COLOMBO, MRS. BERNARDO, Instituto Universitario di Venezia, Venice, Italy CUNNINGHAM, W. J., Yale University, New Haven, Connecticut DAVIS, DAVID E., Johns Hopkins University, Baltimore, Maryland DE, DEEPESH, Carnegie Institution of Washington, Cold Spring Harbor, New

York (Calcutta University, Calcutta, India) DELZELL, DAVID E., University of Michigan, Ann Arbor, Michigan DEMEREC, M., Carnegie Institution of Washington, and Biological Laboratory, Cold

Spring Harbor, New York DOBZHANSKY, TH., Columbia University, New York DOBZHANSKY, N., Columbia University, New York DUNCAN, OTIS DUDLEY, University of Chicago, Chicago, Illinois DUNN, L. C., Columbia University, New York ENGLESBERG, ELLIS, Biological Laboratory, Cold Spring Harbor, New York ERRINGTON, PAUL L., Iowa State College, Ames, Iowa EVANS, FRANCIS C., University of Michigan, Ann Arbor, Michigan FALEK, ARTHUR, Columbia University, New York FALLS, J. BRUCE, University of Toronto, Toronto, Canada FIRSCHEIN, I. LESTER, Columbia University, New York FRIEDMAN, SHEILA, Biological Laboratory, Cold Spring Harbor, New York GAY, HELEN', Carnegie Institution of Washington, Cold Spring Harbor, New York GHENT, A. W., Canada Department of Agriculture, Sault Ste. Marie, Canada GRACE, EDWIN J., The Grace Clinic, Brooklyn, New York GRACE, THOMAS, Rockefeller Institute, New York HAIRSTON, NELSON G., University of Michigan, Ann Arbor, Michigan

ix

x LIST OF THOSE ATTENDING SYMPOSIUM

HAJNAL, JOHN, London School of Economics and Political Science, London, England HASKINS, CARYL P., Carnegie Institution of Washington, Washington, D. C. HASKINS, EDNA FERRELL, Washington, D. C. HENRY, LOUIS, Institut National d't~tudes D~mographiques, Paris, France HIRSCH, JOAN M., University of Chicago, Chicago, Illinois HODGE, CHARLES, Temple University, Philadelphia, Pennsylvania HONEYMAN, M. S., National Cancer Institute, Bethesda, Maryland HULSE, FREDERICK S., University of Washington, Seattle, Washington HUTCHINSON, G. EVELYN, Yale University, New Haven, Connecticut JAMES, ALLEN P., Atomic Energy of Canada, Chalk River, Canada KAPLAN, ARNOLD R., New York State Psychiatric Institute, New York KAUFMANN, BERWIND P., Carnegie Institution of Washington, Cold Spring Har-

bor, New York KING, JAMES C., Biological Laboratory, Cold Spring Harbor, New York KIRK, DUDLEY, Population Council, New York KISER, CLYDE V., Milbank Memorial Fund, New York KLUYVER, H. N., Institute for Ecological Research, Arnhem, Holland KNAPP, BRYON, Columbia University, New York KOHN, ALAN J., Yale University, New Haven, Connecticut KONTKANEN, PAAVO, Museum Zoologicum, Helsinki, Finland KOZINSKI, A. W., Polish Academy of Science, Warsaw, Poland LAHR, ERNEST L., Carnegie Institution of Washington, Cold Spring Harbor, New

York KUSUKAWA, AKIRA, Kyushu University, Fukuoka, Japan LEVINS, RICHARD, Columbia University, New York LEVITAN, MAX, Woman's Medical College, Philadelphia, Pennsylvania LEWONTIN, R. C., North Carolina State College, Raleigh, North Carolina LINDEGREN, CARL C., Southern Illinois University, Carbondale, Illinois LORIMER, FRANK, American University, Washington, D. C. MALOGOLOWKIN, CHANA, Columbia University, New York (University of Brazil,

Rio de Janeiro, Brazil) MARAMOROSCH, KARL, Rockefeller Institute, New York MAULDIN, W. PARKER, The Population Council, New York MAYR, ERNST, Harvard University, Cambridge, Massachusetts MILNE, A., Durham University, Newcastle upon Tyne, England MOORE, JOHN A., Columbia University, New York MOSER, HERMANN Biological Laboratory, Cold Spring Harbor, New York MUHSAM, HALMUT, V., United Nations, New York NEYMAN, JERZY, University of California, Berkeley, California NICHOLSON, A. J., Commonwealth Scientific and Industrial Research Organization,

Canberra, Australia NOTESTEIN, FRANK W., Princeton University, Princeton, New Jersey 0SBORN, FREDERICK, The Population Council, New York OSBORNE, RICHARD H., Columbia University, New York OZEKI, H., Carnegie Institution of Washington, Cold Spring Harbor, New York PATRICK, RUTH, Academy of Natural Sciences, Philadelphia, Pennsylvania PICKETT, A. D., Canada Department of Agriculture, Kentville, Canada PIMENTEL, DAVID, Cornell University, Ithaca, New York PITELKA, FRANK A., University of California, Berkeley, California POINTING, P. J., Canada Department of Agriculture, Sault Ste. Marie, Canada POLNIK, AMELIA, University of Chicago, Chicago, Illinois POTTER, ROBERT G., JR., Princeton University, Princeton, New Jersey

LIST OF THOSE ATTENDING SYMPOSIUM xi

REED, CHARLES A., University of Illinois, Chicago, Illinois REYNOLDSON, T. B., University College of North Wales, Bangor, Wales RYDER, NORMAN B., University of Wisconsin, Madison, Wisconsin SALZANO, FRANCISCO, University of Michigan, Ann Arbor, Michigan (University

of Rio Grande do Sul, Porto Alegre, Brazil) SANGHVI, L. D., India Cancer Research Institute, Bombay, India SANK, DIANE, New York State Psychiatric Institute, New York SCHLAGER, GUNTHER, University of Kansas, Lawrence, Kansas SCOTT, ELIZABETH L., University of California, Berkeley, California SENGUN, ATIF, Carnegie Institution of Washington, Cold Spring Harbor, New

York (University of Istanbul, Istanbul, Turkey) SHAPIRO, HARRY L., American Museum of Natural History, New York SIBLEY, CHARLES G., Cornell University, Ithaca, New York SKAAR, DAVID, Carnegie Institution of Washington, Cold Spring Harbor, New

York SKELLAM, JOHN G., The Nature Conservancy, London, England SLOBODKIN, LAWRENCE, Univerity of Michigan, Ann Arbor, Michigan SOKAL, ROBERT R., University of Kansas, Lawrence, Kansas SOKOLOFF, ALEXANDER, Hofstra College, Hempstead, New York SPRUGEL, GEORGE, Jr., National Science Foundation, Washington, D. C. SPUHLER, HELEN, University of Michigan, Ann Arbor, Michigan SPUHLER, JAMES N., University of Michigan, Ann Arbor, Michigan STOLNITZ, GEORGE J., Indiana University, Bloomington, Indiana SUTTER, JEAN, Institut National d'Etudes D6mographiques, Paris, France TASHIAN, RICHARD, Columbia University, New York TIETZE, CHRISTOPHER, Falls Church, Virginia TREWARTHA, GLENN T., University of Wisconsin, Madison, Wisconsin UDVARDY, MIKLOS D. F., University of British Columbia, Vancouver, Canada UELAND, ERIC, Yale University, New Haven, Connecticut UTIDA, SYUNRO, Kyoto University, Kyoto, Japan VAN VALEN, LEIGH, Columbia University, New York VETUKHIV, MICHAEL, Columbia University, New York VILKOMERSON, HILDA, University of Pennsylvania, Philadelphia, Pennsylvania VON WETTSTEIN, DITER, Forest Research Institute, Stockholm, Sweden WADDINGTO.N, C. H., Institute of Animal Genetics, Edinburgh, Scotland WALKER, NORMA FORD, University of Toronto, Toronto, Canada WALLACE, BRUCE, Biological Laboratory, Cold Spring Harbor, New York WANGERSKY, PETER J.,Yale University, New Haven, Connecticut WARREN, CHARLES 0., Commonwealth Fund, New York WARREN, KATHERINE BREHME, Biological Laboratory, Cold Spring Harbor, New

York; Adelphi College, Garden City, New York WASSERMANN, F., New York University Medical College, New York WHEATLAND, SARAH, Yale University, New Haven, Connecticut WOODELL, STANLEY, London, England YERUSHALMY, JACOB, University of California, Berkeley, California

Contents

Foreword . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v Frontispiece . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii List of Participants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix

POPULATION STUDIES: ANIMAL ECOLOGY AND DEMOGRAPHY

INTRODUCTORY SURVEY

COLE, LAMONT C. Sketches of General and Comparative Demography . . . . . . 1

HUMAN POPULATIONS: HISTORICAL STUDY

LORIMER, FRANK. Introductory Remarks of the Chairman . . . . . . . . . . . . . . . . . 17 BRMDWOOD, ROBERT J., and REED, CHARLES A. The Achievement and Early

Consequences of Food-Production: A Consideration of the Archeological and Natural-Historical Evidence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

HULSE, FREDERICK S. Some Factors Influencing the Relative Proportions of Human Racial Stocks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

BIRDSELL, JOSEPH B. Some Population Problems Involving Pleistocene Man. 47 TREWARTHA, GLENN T., and GOSAL, GURDEV. The Regionalism of Population

Change in India . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71

DEMOGRAPHIC THEORY

COALE, ANSLEY J. How the Age Distribution of a Human Population is Deter- mined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

RYDER, N. ]~. The Conceptualization of the Transition in Fertility . . . . . . . . . 91 HAJNAL, J. Mathematical Models in Demography . . . . . . . . . . . . . . . . . . . . . . . 97 HENRY, LouIs. Theoretical Research and Demography . . . . . . . . . . . . . . . . . . . . 105

ANIMAL POPULATIONS, EXPERIMENTAL AND THEORETICAL

NEYMAN, JERZY, and SCOTT, ELIZABETH L. On a Mathematical Theory of Populations Conceived as a Conglomeration of Clusters . . . . . . . . . . . . . . . 109

MOSER, H. Structure and Dynamics of Bacterial Populations Maintained in the Chemostat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I21

UTIDA, SYUNRO. Population Fluctuation, an Experimental and Theoretical Approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

NIC~IOLSON, A. J. The Self-adjustment of Populations to Change . . . . . . . . . . . . 153

GROWTH OF HUMAN POPULATIONS

POTTER, ROBERT G. JR. Farris' Formula for Predicting Fertile Days . . . . . . . . 175 STOLNITZ, G. J. The Old and the New in Recent International Patterns of

Population Growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187 COLOMBO, BERNARDO. On the Sex Ratio in Man . . . . . . . . . . . . . . . . . . . . . . . . . . 193

ECOLOGY OF NATURAL POPULATIONS, I

BIRCH, L. C. The Role of Weather in Determining the Distribution and Abund- ance of Animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203

ANDREWARTHA, H. G. The Use of Conceptual Models in Population Ecology.. 219 PITELKA, FRANK A. Some Aspects of Population Structure in the Short-term

Cycle of the Brown Lemming in Northern Alaska . . . . . . . . . . . . . . . . . . . . . 237

X l l l

xiv CONTENTS

ECOLOGY OF NATURAL POPULATIONS, II

MILNE, A. Theories of Natural Control of Insect Populations . . . . . . . . . . . . . . 253 BLAIR, W. FRANK. Changes in Vertebrate Populations under Conditions of

Drought . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273 CHITTY, DENNIS, Self-regulation of Numbers through Changes in Viability.. 277 KLUYVER, H. N. Roosting Habits, Sexual Dominance and Survival in the

Great Tit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281 ERRINGTON, PAUL L. Of Population Cycles and Unknowns . . . . . . . . . . . . . . . . 287

ECOLOGY OF NATURAL POPULATIONS, III

UDVARDY, MIKLOS D. F. An Evaluation of Quantitative Studies in Birds . . . . 301 REYNOLDSON, T. B. Population Fluctuations in Urceolaria mitra (Peritricha)

and Enchytraeus albidus (Oligochaeta) and their Bearing on Regulation.. 313 WANGERSKY, P. J., and CUNNINGHAM, W. J. Time Lag in Population Models. 329

THE STRUCTURE OF COMMUNITIES

CALHOUN, JOHN B. Social Welfare as a Variable in Population Dynamics . . . . 339 DUNCAN, OTIS DUDLEY, Population Distribution and Community Structure.. 357 KONTKANEN, PAAVO. On the Delimitation of Communities in Research on

Animal Biocoenotics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 373 SUTTER, JEAN, and TRAN-NGoc-ToAN. The Problem of the Structure of Iso-

lates and of their Evolution among Human Populations . . . . . . . . . . . . . . . 379

THE POPULATION AS A UNIT OF EVOLUTION

DOBZHANSKY, THEODOSIUS. Mendelian Populations as Genetic Systems . . . . . 385 LEWONTIN, R. C. The Adaptations of Populations to Varying Environments. 395 SHAPIRO, HARRY L. The Population Unit and Culture . . . . . . . . . . . . . . . . . . . . . 409

CONCLUSION

HUTCHINSON, G. EVELYN. Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . 415