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COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY
VOLUME XXV
COLD SPRING HARBOR SYMPOSIA
ON Q UANTITA T I V E BIOLOGY
Founded in 1933
by 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 o/ 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 XXV
Biological Clocks
THE BIOLOGICAL LABORATORY
COLD SPRING HARBOR, L. I., NEW YORK
1960
COPYRIGHT �9 1961 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.
69166
PRINTED BY
WAVERLY PRESS, INC., BALTIMORE, MARYLAND, U. S. A.
Foreword
While daily rhythms have been studied systematically for many years, there has been a most remarkable outburst of interest in the subject during the past ten years. This growth of interest is due, almost entirely, to the elegant experimental demonstrations that birds, bees, and many other animals orient or "navigate" using the position of the sun as a guidepost, and compensating for its movement in time with the use of an internal chronometer. Rapid advances have been seen in studies of rather diverse phenomena in a wide variety of organisms from single cells through higher plants and animals.
By bringing together leading investigators from throughout the world who are intimately concerned with the broad array of phenomena classified under the general topic "Biological Clocks", it is hoped that this year's Symposium may serve as a unifying influence on the entire field of study.
The program this year was organized by a committee consisting of C. S. Pitten- drigh (Chairman), J. Aschoff, V. G. Bruce, E. Biinning, D. R. Griffin, and J. W. Hastings. Serving as chairmen of the program sessions were: C. S. Pittendrigh, E. Bfinning, J. Aschoff, S. B. Hendricks, V. G. Bruce, H. Kalmus, O. H. Schmitt, A. D. Lees, K. V. Thimann, C. P. Richter, F. Halberg, R. Bfinsow, D. R. Griffin, K. S. Rawson, K. Hoffmann, A. D. Hasler, and F. A. Brown, Jr. The Laboratory is indeed grateful to all of these gentlemen for their efforts in organizing and con- ducting the program.
The Laboratory is pleased to acknowledge support of the Symposium by The Carnegie Corporation of New York, The Rockefeller Foundation, Institute of Oceanography and Marine Biology, National Science Foundation, Office of Naval Research under the auspices of the American Institute of Biological Sciences, United States Public Health Service, and the United States Air Force monitored by the Office of Scientific Research of the Air Research and Development Com- mand.
This year, preparation of the volume for publication was supervised by our editor, Leonora Frisch, assisted by Ruth Schalet and Ethel Burtch. The very difficult task of indexing was handled by Dr. C. S. Pittendrigh and his colleagues at Princeton University. To all of these people, the Laboratory owes a debt of gratitude.
The meetings, held from the 5th through the 14th of June, 1960, were attended by 150 scientists, including 31 from abroad.
ARTHUR CHOVNICK
LIST OF PREVIOUS VOLUMES
Volume I (1933) Surface Phenomena, 239 pp. Volume II (1934) Aspects of Growth, 284 pp. Volume III (1935) Photochemical Reactions, 359 pp. Volume IV (1936) Excitation Phenomena, 376 pp. Volume V (1937) Internal Secretions, 433 pp. Volume VI (1938) Protein Chemistry, 395 pp. Volume VII (1939) Biological Oxidations, 463 pp. Volume VIII (1940) Permeability and the Nature of Cell Membranes, 285 pp. Volume IX (1941) Genes and Chromosomes: Structure and Organization, 315 pp. Volume 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 XII I (1948) Biological Applications of Tracer Elements, 222 pp. Volume XIV (1949) Amino Acids and Proteins, 217 pp. Volume XV (1950) Origin and Evolution of Man, 425 pp. Volume XVI (1951) Genes and Mutations, 521 pp. Volume XVII (1952) The Neuron, 323 pp. Volume XVIII (1953) Viruses, 301 pp. Volume XIX (1954) The Mammalian Fetus: Physiological Aspects of Development,
225 pp. Volume XX (1955) Population Genetics: The Nature and Causes of Genetic Varia-
bility in Populations, 346 pp. Volume XXI (1956) Genetic Mechanisms: Structure and Function, 392 pp. Volume XXII (1957) Population Studies: Animal Ecology and Demography, 437 pp. Volume XXI I I (1958) Exchange of Genetic Material : Mechanisms and Consequences,
450 pp. Volume XXIV (1959) Genetics and Twentieth Century Darwinism, 321 pp.
vi
SOME SYMPOSIUM PARTICIPANTS
Top row: H. M. Webb, H. Kalmus; W. Braemer, K. Hoffmann, A. D. Hasler; F. Halberg, A. Sollberger, H. Kalmus, A. D. Lees.
Second row; K. Klotter, Th. Hellbrtigge, J. Aschoff, C. S. Pittendrigh, D. S. Lehrman; A. Wolfson, A. J. Marshall; W. Braemer, B. J. Oetting, H. J. Ball, G. Birukow.
Third row: Th. Helbriigge, E. Segal, B. Schwemmle; H. M. Webb, J. Harker, M. Sandeen; F. A. Brown, Jr., M. B. Wilkins, G. Birukow, E. Biinning.
Bottom row: A. Barnett, B. Sweeney, M. Lindauer; K. S. Rawson, O. H. Schmitt; S. B. Hendricks, J. E. Flynn, K. V. Thimann.
SOME SYMPOSIUM PARTICIPANTS
Top row: F. W. Went, J. Aschoff; S. K. Roberts, P. DeCoursey; C. S. Pittendrigh, A. Chovnick, E. Bfinning. Second row: L. Frisch, V. G. Bruce; F. A. Brown, Jr., G. Birukow; A. D. Lees, C. E. Jenner. Th i rd row: H. 0. Schwassmann, A. Chovnick, M. B. Wilkins, M. Renner, H. Hayes; D. Kennedy, M. B. Wilkins, M. C.
Lobban; H. G. Wallraff, D. M. A. Mercer, A. Sollberger, F. A. Brown, Jr. Bot tom row: K. Hoffmann, C. Hauenschild, P. DeCoursey, P. S. Corbet, J. Aschoff, J. L. Cloudsley-Thompson;
H. Kalmus, A. Chovnick, 0. H. Schmitt, H. E. Savely, J. T. Enright; S. Mori, M. Kato.
(Photographs by Robert Beaumont)
List of Those Attending the Symposium ADLER, HELMUT E., American Museum of Natural
History, New York City AKEY, ALICE, Adelphi College, Garden City, New
York ALEXANDER, MARY L., University of Texas, Houston,
Texas ANDERSON, PAUL, Columbia University, New York
City ASCHOFF, J~RGEN, Max-Planck-Institut ffir Verhaltens-
physiologic, Erling-Andechs/Obb., Germany BALL, HAROLD J., University of Nebraska, Lincoln,
Nebraska BARLOW, JOHN S., Massachusetts Institute of Tech-
nology, Cambridge, Massachusetts and Massachu- setts General Hospital, Boston, Massachusetts
BARNETT, AUDREY, Indiana University, Bloomington, Indiana
BARNWELL, FRANKLIN H., Northwestern University, Evanston, Illinois
BARTELL, CLELMER K., Duke University, Durham, North Carolina
BARTH, ROBERT, Harvard University, Cambridge, Massachusetts
BARTON, ALEXANDER, The Stony Brook School, Stony Brook, New York
BENNETT, MIRIAM F., Sweet Briar College, Sweet Briar, Virginia
BIANCHI, CARLA, Northwestern University, Evanston, Illinois
BIRUKOW, GEORG, Zoologisches Institut der Universi- titt, GSttingen, Germany
BLAKESLEE, ALTON, The Associated Press, New York City
BLISS, DOROTHY E., The American Museum of Natural History, New York City
BRAEMER, WOLFGANG, Max-Planck-Institut, Seewiesen, fiber Starnberg/Obb., Germany
BRANDT, WILLIAM H., Oregon State College, Corvallis, Oregon
BREHME, KATHERINE, Adelphi College, Garden City, New York
BROWN, CLIFFORD, National Film Board of Canada, Montreal, Canada
BROWN, FRANK A., JR., Northwestern University, Evanston, Illinois
BRUCE, VICTOR G., Princeton University, Princeton, New Jersey
B?3NNING, ERWlN, Botanisches Institut, Universitiit Tiibingen, Germany
B?3NSOW, ROBERT C., Pflanzenphysiologisches Institut der Universitiit, G6ttingen, Germany
BUTNER, ALFRED, Jamaica, New York CARR, ARCHIE, University of Florida, Gainesville,
Florida ix
CAVANAGH, JAMES, Garden City, New York CHOVNICK, ARTHUR, Biological Laboratory, Cold
Spring Harbor, New York CLOUDSLEY-THOMPSON, J. L., University of London,
Ydng's College, London, England COLLINS, WALTER, Colorado State University, Fort
Collins, Colorado CONKLIN, MARIE E., Adelphi College, Garden City,
New York CORBET, PHILIP S., East Africa Virus Research Insti-
tute, Entebbe, Uganda, East Africa DECouRSEY, PATRICIA, University of Wisconsin, Madi-
son, Wisconsin EHRET, CHARLES F., Argonne National Laboratory,
Leniont, Illinois ENGELMANN, FRANZ, Albert Einstein College of Medi-
cine, New York City ENRIGHT, J. T., University of California, LaJolla,
California FAY, ANTHONY F., Flushing, New York FINGERMAN, MILTON, Tulane University, New Orleans,
Louisiana FLYNN, JOHN E., Office of Naval Research, New York
City FOLK, G. EDGAR, State University of Iowa, Iowa City,
Iowa FRISCH, LEONORA, Biological Laboratory, Cold Spring
Harbor, New York GALINSKY, IRVING, Hofstra College, Hempstead, New
York GALLER, S. R., Omce of Naval Research, Washington,
D.C. GAY, HELEN, Carnegie Institution of Washington, Cold
Spring Harbor, New York GOULD, EDWIN, Tulane University, New Orleans,
Louisiana GRACE, EDWIN J., Grace Clinic Group, Brooklyn, New
York GRIFFIN, DONALD R., Harvard University, Cambridge,
Massachusetts GUYSELMAN, J. BRUCE, Carleton College, Northfield,
Minnesota HALBERG, ERNA, University of Minnesota Medical
School, Minneapolis, Minnesota HALBERG, FRANZ, University of Minnesota Medical
School, Minneapolis, Minnesota HAMILTON, WILLIAM J., California Academy of Sci-
ences, San Francisco, California HAMNER, KARL C., University of California, Los
Angeles, California HAN, JACK, State University of New York, Syracuse,
New York
X LIST OF THOSE ATTENDING THE SYMPOSIUM
HARKER, JANET E., Zoological Laboratory, Cambridge, England
HASLER, ARTHUR D., University of Wisconsin, Madi- son, Wisconsin
HASTINGS, ANTOINETTE, American Association for the Advancement of Science, Westbury, New York
HASTINGS, J. WOODLAND, University of Illinois, Urbana, Illinois
HAUENSCHILD, C., Zoologisches Institut der Universi- t~t, Freiburg/Br., Germany
HAYES, HELEN, Office of Naval Research, Washington. D.C.
HELLBRiJGGE, THEODOR, Kinderpoliklinik der Universi- t/it, M0.nchen, Mr:richeR, Germany
HELMS, CARL W., Massachusetts Audubon Society, South Lincoln, Massachusetts
HENDRICKS, STERLING B., V. S. Department of Agri- culture, Beltsville, Maryland
HIGHKIN, n. R., California Institute of Technology, Pasadena, California
HILLS, ALICIA, Life Magazine, New York City HOAGLAND, HUDSON, The Worcester Foundation for
Experimental Biology, Shrewsbury, Massachusetts HOFFMANN, KLAUS, Max-Planck-Institut ftir Verhalt-
ensphysiologie, Erling-Andechs/Obb., Germany, and Princeton University, Princeton, New Jersey
HoPwooD, JOSEPH, Colorado State University, Fort Collins, Colorado
HUBBARD, HENRY, Newsweek, New York City ISAAC, DONALD, University of California, Berkeley,
California JACOBS, GEORGE, National Aeronautics and Space Ad-
ministration, Washington, D. C. JENNER, CHARLES E., University of North Carolina,
Chapel Hill, North Carolina KAHN, DAVID, Newsday, Garden City, New York KALMUS, HANS, University College London, London,
England KARAKASHIAN, MARLENE W., University of California,
Los Angeles, California KATO, MASARU, Washington State University, Pullman,
Washington KAUFMANN, BERWIND N., University of Mississippi
Medical Center, Jackson, Mississippi KAUFMANN, BERWIND P., Carnegie Institution of Wash-
ington, Cold Spring Harbor, New York KENNEDY, DONALD, Syracuse University, Syracuse,
New York KEBNAGHAN, ROY PETER, Biological Laboratory, Cold
Spring Harbor, New York KLOTTER, KARL, Institut fflr Angewandte Mechanik,
Darmstadt, Germany KRAMER, SOL, State University College on Long Island,
Oyster Bay, New York KRAUSS, MARIAN, Biological Laboratory, Cold Spring
Harbor, New York
KRIS, E. CHRISTINE, Cambridge, Massachusetts LARSEN, VICTOR, Adelphi College, Garden City, New
York LEES, ANTHONY D., Agricultural Research Council,
Cambridge, England LEHRMAN, DANIEL S., Rutgers University, Newark,
New Jersey LEINWEBER, FRANZ J., Johns Hopkins University,
Baltimore, Maryland LEINWEBER, LUPE, Baltimore, Maryland LIENAU, CARL, Columbia University, New York City LINCOLN, RICHARD G., Long Beach State College, Long
Beach, California LINDAUER, MARTIN, Zoologisches Institut der Universi-
tiit M0.nchen, Miinchen, Germany LOBBAN, MARY C., Medical Research Council :Labora-
tories, London, England MACDOWELL, E. C., Biological Laboratory, Cold Spring
Harbor, New York MALM, MIGNON, Princeton University, Princeton, New
Jersey MARSHALL,'A. J., Monash University, Victoria, Aus-
tralia MENAKER, MICHAEL, Harvard University, Cambridge,
Massachusetts MERCER, DERWENT ~. A., Physical Laboratory, The
University, Southampton, England MILLMAN, EDWARD, Grumman Aircraft Engineering
Corporation, Bethpage, New York MINIS, DOROTHY, Princeton University, Princeton, New
Jersey MIYAKE, AKIO, Osaka City University, Osaka, Japan MOE, GORDON K., Masonic Medical Research Labora-
tory, Utica, New York MORI, SYUITI, Zoological Institute, Kyoto University,
Kyoto, Japan MUKAI, FRANK, Carnegie Institution of Washington,
Cold Spring Harbor, New York NOVICK, ALVIN, Yale University, New Haven, Con-
necticut NOWOSIELSKI, JOZEF, Cornell University, Ithaca, New
York 0ETTING, BONNALIE J., Northwestern University,
Evanston, Illinois 0L~VE, JOHN R., American Institute of Biological
Sciences, Washington, D. C. PALMER, JOHN, Northwestern University, Evanston,
Illinois PETROPULOS, STEPHEN F., Princeton University, Prince-
ton, New Jersey PITTENDRIGH, COLIN S., Princeton University, Prince-
ton, New Jersey PROKOP, BARBARA, Biological Laboratory, Cold Spring
Harbor, New York RAWSON, KENNETH S., Swarthmore College, Swarth-
more, Pennsylvania
LIST OF THOSE ATTENDING THE SYMPOSIUM xi
RAWSON, KENNETH S., MRs., Swarthmore College, Swarthmore, Pennsylvania
RENh'ER, MAX, Zoologisches Institut der Universitiit, Miinchen, Germany
RI(?ttTER, CURT P., The Johns Hopkins Hospital, Balti- more, Maryland
ROBERTS, SHEPHERD K., Princeton University, Prince- ton, New Jersey
SAND EEN, MURIEL I., Duke University, Durham, North Carolina
SAVELY, HARVEY E., U. S. Air Force, Washington, D.C.
SCHALET, RUTH, Biological Laboratory, Cold Spring Harbor, New York
SCHARRER, BERTA, Albert Einstein College of Medicine, New York City
SCHARRER, ERNST, Albert Einstein College of Medicine, New York City
SCHI'~ELE, GEORGE A., Princeton University, Princeton, New Jersey
SCHEFANO, ANTHONY, Hofstra College, Hempstead, New York
SC~IDT-KOENIG, KLAUS, Max-Planck-Institut far Verhaltensphysiologie, Wilhehnshaven, Germany, and Duke University, Durham, North Carolina
SCHMITT, OTTO H., University of Minnesota, Minne- apolis, Minnesota
SCHMITT, OTTO H., MRS., University of Minnesota, Minneapolis, Minnesota
SCHWASSMANN, HORST 0., University of Wisconsin, Madison, Wisconsin
SCH~VEMMLE, BERTHOLD, Pflanzenphysiologisches Insti- rut der Universiti~t, GSttingen, Germany
SEGAL, EARL, Kansas State Teachers College, Emporia, Kansas
SHAPIRO, ARTHUR, New York State University, Brook- lyn, New York
SHAPIRO, SEYMOUR, Brookhaven National Laboratory, Upton, New York
SIMON, EDWARD, Carnegie Institution of Washington, Cold Spring Harbor, New York
SIROHI, G. S., University of California, Los Angeles, California
SMOLKER, ROBERT, State University of New York, Oyster Bay, New York
SOKOLOFF, ALEXANDER, William H. Miner Agricultural Research Institute, Chazy, New York
SOKOLOFF, ALEXANDER, MB.S., William H. Miner Agri- cultural Research Institute, Chazy, New York
SOLLBERGER, ARNE, Caroline Institute, Stockholm, Sweden
SUCKLING, EUSTACE, State University of New York Medical College, Brooklyn, New York
SWADE, RICHARD H., Princeton University, Princeton, New Jersey
SWEENEY, BEATRICE M., University of California, LaJolla, California
TALL, BENITA, American Institute of Biological Sci- ences, Washington, D. C.
THIMANN, KENNETH V., Harvard University, Cam- bridge, Massachusetts
THORNBURG, WAYNE, Dartmouth Medical School, Hanover, New Hampshire
TOLLES, WALTER, Institute of Oceanography and Marine Biology, Oyster Bay, New York
~VALLRAFF, HANS G., Max-Planck-Institut ftir Verhalt- ensphysiologie, Wilhelmshaven, Germany
WEBB, H. MARGUERITE, Stazione Zoologica, Napoli, Italy
WEITZMAN, MARY, Albert Einstein College of Medicine, New York City
WELLS, G. P., University College London, London, England
WENT, F. W., Missouri Botanical Garden, St. Louis, Missouri
WEVER, Ri2TGER, Max-Planck-Institut ftir Verhaltens- physiologic, Erling-Andechs/Obb, Germany
WILDER, JOSEPH, Society for Biological Rhythms, Stockholm, Sweden
WILKINS, MALCOLM B., University of London King's College, London, England
WISPY, WARREN, University of Miami, Miami, Florida WOLF, WILLIAM, New York City WOLFSON, ALBERT, Northwestern University, Evans-
ton, Illinois WULFF, VERNER J., Masonic Medical Research Lab-
oratory, Utica, New York
Contents
Foreword . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v List of Participants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix BttiNrNO, ERWIN. Opening Address: Biological Clocks . . . . . . . . . . . . . . . . . . . . . 1
GENERAL PROPERTIES OF CIRCADIAN RHYTHMS
ASCHOFF, J. Exogenous and Endogenous Components in Circadian Rhythms. 11 BRUCE, V. G. Environmental Entra inment of Circadian Rhythms . . . . . . . . . . 29 DECouRsEY, P. Phase Control of Activity in a Rodent . . . . . . . . . . . . . . . . . . . . 49 BROWN, F. A., JR. Response to Pervasive Geophysical Factors and the Biological
Clock Problem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 MERCER, D. M. A. Analytical Methods for the Study of Periodic Phenomena
Obscured by Random Fluctuations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 S~VEENEY, B., AND J. W. HASTINGS. Effects of Temperature upon Diurnal
Rhythms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 RAWSON, K. S. Effects of Tissue Temperature on Mammalian Activity
Rhythms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 WILKINS, M. B. The Effect of Light upon Plant Rhythms . . . . . . . . . . . . . . . . . . 115
METABOLIC ASPECTS OF CIRCADIAN RHYTHMS
HASTINGS, J. W. Biochemical Aspects of Rhythms: Phase Shifting by Chem- icals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
SWEENEY, B. The Photosynthetic Rhy thm in Single Cells of Gonyaulax
polyedra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145 EHRET, C. F. Action Spectra and Nucleic Acid Metabolism in Circadian
Rhythms at the Cellular Level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
MODELS FOR CIRCADIAN RHYTHMS
PITTENDRIGH, C. S. Circadian Rhythms and the Circadian Organization of Living Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
KLOTTER, K. General Properties of Oscillating Systems . . . . . . . . . . . . . . . . . . . . 185 KLOTTER, K. Theoretical Analysis of Some Biological Models . . . . . . . . . . . . . . . 189 WEVER, R. Possibilities of Phase-control, Demonstrated by an Electronic
Model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197 SCHMITT, O. H. Biophysical and Mathematical Models of Circadian Rhythms 207 KALMUS, H., AND L. A. WIGGLESWORTH. Shock Excited Systems as Models for
Biological Rhythms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211 EHRET, C. F., AND J. S. BARLOW. Toward a Realistic Model of a Biological
Period-measuring Mechanism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
CIRCADIAN RHYTHMS IN PHOTO- AND THERMOPERIODISM
WENT, F. W. Photo- and Thermoperiodic Effects in Plant Growth . . . . . . . . . . 221 HIGHKIN, H. R. The Effect of Constant Temperature Environments and of
Continuous Light on the Growth and Development of Pea Plants . . . . . . 231 SCHWEMMLE, B. Thermoperiodic Effects and Circadian Rhythms in Flowering
of Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239 I-IENDRICKS, S. B. Rates of Change of Phytochrome as an Essential Factor
Determining Photoperiodism in Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 BiJNNING, E. Circadian Rhythms and the Time Measurement in Photo-
periodism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249 xii
CONTENTS xiii
B?)NSOW, R. C. The Circadian R h y t h m of Photoperiodic Responsiveness in K a l a n c h o ~ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 257
LEES, A. D. Some Aspects of Animal Photoperiodism . . . . . . . . . . . . . . . . . . . . . 261 HAMNER, K. C. Photoperiodism and Circadian Rhy thms . . . . . . . . . . . . . . . . . . . 269
CIRCADIAN RHYTHMS IN ANIMAL PHYSIOLOGY
HARKER~ J. Endocrine and Nervous Factors in Insect Circadian R h y t h m s . . . 279 HALBERG, F. Temporal Coordination of Physiologic Function . . . . . . . . . . . . . . 289 HELLBR~GGE, TIt. The Development of Circadian Rhy thms in Infants . . . . . . 311 LOBBAN, M. C. The Ent ra inment of Circadian Rhy thms in Man . . . . . . . . . . . 325 MoRI, S. Analysis of Environmental and Physiological Factors on the Daily
Rhythmic Activi ty of a Sea Pen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333
ECOLOGICAL ASPECTS OF CIRCADIAN RHYTHMS
CLOUDSLEY-THoMPSON, J. L. Adapt ive Functions of Circadian Rhy thms . . . . 345 CORBET, P. S. Pat terns of Circadian Rhy thms in Insects . . . . . . . . . . . . . . . . . . 357 RENNER, M. The Contribution of the Honey Bee to the Study of Time-Sense
and Astronomical Orientation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361
CHRONOMETRY IN CELESTIAL ORIENTATION
Dedication to Gus tav Kramer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 369 LINDAUER, M. Time-Compensated Sun Orientation in Bees . . . . . . . . . . . . . . . . 371 HOFFMANN, K. Experimental Manipulat ion of the Orientational Clock in
Birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379 SCHMIDT-KOENIG, K. Internal Clocks and Homing . . . . . . . . . . . . . . . . . . . . . . . . 389 PARDI, L. Innate Components in the Solar Orientation of Littoral Amphipods. 395 BIRUKOW, G. Innate Types of Chronometry in Insect Orientation . . . . . . . . . . 403 BRAEMER, W. A Critical Review of the Sun-Azimuth Hypothesis . . . . . . . . . . . 413 HASLER, A. D., AND H. O. SCHWASSMANN. Sun Orientation of Fish a t Different
Latitudes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429 SCHWASSMANN, H. O. Environmental Cues in the Orientation R h y t h m of Fish. 443 WALLRAFF, H. G. Does Celestial Navigat ion Exist in Animals? . . . . . . . . . . . . . 451 SAUER, F., AND E. SAUER. Star Navigat ion of Nocturnal Migrating Birds . . . . 463 PAl)I, F. Orientation by Night: The Moon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475
LONGER PERIODICITIES
FINGERMAN, M. Tidal Rhythmic i ty in Marine Organisms . . . . . . . . . . . . . . . . . . 481 HAUENSCHILD, C. Lunar Periodicity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 491 MARSHALL, A. J. Annual Periodicity in the Migrat ion and Reproduction of
Birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 499 WOLFSON, A. Regulation of Annual Periodicity in the Migration and Repro-
duction of Birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 507