Table Of ContentACADEMIC PRESS RAPID MANUSCRIPT REPRODUCTION
Proceedings of a Satellite Symposium of the 140th
Meeting of the American Association for the Advance-
ment of Science Held in San Francisco, California,
February 25, 1974
CIRCANNIM.
CLOCKS
ANNUAL BIOLOGICAL RHYTHMS
EDITED BY
6RIC T P€NG€Ll£Y
Department of Biology
University of California, Riverside
ACADEMIC PRESS, INC
NEW YORK SAN FRANCISCO LONDON 1974
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Participants
Ivan Assenmacher, Laboratoire de Physiologie Animale, Faculté des Sciences,
Place Eugene Bataillon, Université de Montpellier, 34 Montpellier, France.
Sigfried P. Berthold, Vogelwarte Radolfzell am Max-Planck-Institut für Verhal-
tensphysiologie, 7761 Schloss Moggingen, West Germany.
Mary Anne Brock, Gerontology Research Center, Baltimore City Hospitals,
Baltimore, Maryland, 21224
Albert R. Dawe, Office of Naval Research, 536 S. Clark Street, Chicago, Illinois,
60605
James T. Enright, Scripps Institute of Oceanography, P. Ο. Box 1529, La Jolla,
California, 92037
Richard J. Goss, Division of Biological and Medical Sciences, Brown University,
Providence, Rhode Island, 02912
Bengt W. Johansson, Heart Laboratory, Department of Medicine, General
Hospital, S-21401 Malmo, Sweden.
Helmut Klein, Max-Planck-Institut für Verhaltensphysiologie, Seewiesen und
Erling-Andechs, 8131 Erling-Andechs, West Germany.
Michael Menaker, Department of Zoology, University of Texas, Austin, Texas,
78712
Nicholas Mrosovsky, Department of Zoology, University of Toronto, Toronto,
Ontario, M5S IA5, Canada.
Eric T. Pengelley, Department of Biology, University of California, Riverside,
California, 92502
Alain Reinberg, Laboratoire de Physiologie, Fondation A. de Rothschild, 29
Rue Manin, 75, Paris-19e, France.
James T. Rutledge, Department of Animal Physiology, University of California,
Davis, California, 95616
Robert Schwab, Department of Animal Physiology, University of California,
Davis, California, 95616
vii
PARTICIPANTS
Jerome Β. Senturia, Department of Biology and Health Sciences, The Cleveland
State University, Cleveland, Ohio, 44115
Wilma A. Spurrier, Department of Physiology, Stritch School of Medicine, 2160
South First Avenue, May wood, Illinois, 60153
ABSENT
Sally J. Asmundson, Department of Biology, University of California, Riverside,
California, 92502
Eberhard Gwinner, Max-Planck-Institut für Verhaltensphysiologie, Seewiesen und
Erling-Andechs, 8131 Erling-Andechs, West Germany.
viii
Preface
The purpose of this symposium is to summarize the present state of knowl-
edge on endogenous annual rhythms (circannual clocks), and to point out their
biological significance and importance. It is further hoped that the symposium
will give the phenomenon the attention it deserves, particularly since many
biologists are unaware of it. In addition we gathered our knowledge on the
subject under one heading, so that future investigators, particularly the young,
can use it as a starting point for future ideas and experiments. It is also hoped
that there will be a meeting of minds on the subject, with careful analysis of
each others work. This book is the outcome of these aims and objectives.
To the critical reader it will be perfectly obvious that our ignorance of the
basic nature of the subject is profound. This is due primarily to two factors,
namely that the discovery of the phenomenon is relatively new and that the
demands of the time involved make it very difficult to study. It must also be
stressed that we know of the existence of endogenous annual rhythms only in
very few species of animals, and it seems remarkable that they have apparently
never been explored or demonstrated in plants. Hopefully this will be under-
taken, as it may prove of great importance. It will be of help also to point out
that our terminology at this time has largely been borrowed from those people
who study circadian rhythms. The adaptations and meanings of this terminology
have been thoroughly discussed and explained in the papers by Dr. Alain
Reinberg and by Pengelley and Asmundson. The inexperienced reader may wish
to consult the introductions of these two papers first, in order to master the
terminology.
In both the organization of the symposium and the preparation of this volume
I have been greatly assisted by many people. First, I wish to acknowledge the
initiative and leadership of Dr. Richard Goss. He was a constant source of
encouragement and help when I needed it most. Second, I must thank Mrs.
Elisabeth Zeutschel of the AAAS administrative staff for her excellent organiza-
tion. She has somewhat restored my trust in administrators. I am indebted to
Dr. William Fuller for supplying me with the background material on Professor
Rowan, and to Mr. Robert Lister for writing his biography. I wish also to express
my gratitude to Drs. Alain Reinberg, Ivan Assenmacher, Bengt Johansson,
Helmut Klein, and Peter Berthold for traveling all the way from their respective
European countries to be with us in San Francisco. Finally I thank Mrs. Mary
Hickey who retyped the manuscripts for the final production process.
ix
Professor William Rowan (1891-1957)
Karsh, Ottawa
This volume is dedicated, by the Participants of the
Symposium, to the memory of Professor William Rowan,
in honor of the fiftieth anniversary of his classic work on
the Junco.
There follows a short biography of Professor Rowan,
and his original paper of 1925.
BIOGRAPHICAL SKETCH OF
PROFESSOR WILLIAM ROWAN
by
Robert Lister
Formerly Research Assistant to
Professor Rowan
Edmonton, Alberta
January 1974
PROFESSOR WILLIAM ROWAN was born in
Basle, Switzerland in 1891 of English-Swiss
parents. He was educated in England and
received his M.Sc. from University College,
London. For a short time he taught at
Bedford, but in 1919 accepted an appoint-
ment to the Zoology Department at the Uni-
versity of Manitoba. A year later he joined
the Biology Department of the University of
Alberta and when that department split in
1921, he became head of the new Department
of Zoology, a post he held until his retire-
ment in 1956.
Rowan's early field notes show his
keen interest in birds while still a boy,
but it was as a university student at
Blakeney Point on the north Norfolk coast of
England that he probably acquired his life-
long interest in bird migration.
The mud flats and saltings at Blakeney
teemed with migrating shorebirds while sand
and shingle annually hosted large colonies
of breeding terns.
When he arrived in Alberta he found
near Edmonton an area that was reminiscent
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BIOGRAPHICAL SKETCH
of Blakeney. Here again, at Beaverhills
Lake in spring, the lake flats swarmed with
countless waders on passage to the Arctic
and geese in their thousands rested to await
the breakup of the northern ice.
It was the remarkable regularity with
which the birds returned each spring that
fired Rowan 1 s imagination. The birds
arrived within a day or two of the same date
each year. Weather was variable and he
looked for some constant that might regulate
the spring migration. The lengthening days,
it seemed to him, might be the governing
factor. He decided to subject birds to
increasing periods of light to simulate the
lengthening daylight as a test of his hypo-
thesis.
His classical experiments on juncos
and crows are too well known to need eluci-
dation. It is sufficient to note here that
although we know today that light in itself
does not have the profound effect on bird
migration that Rowan thought it might, his
work pioneered a new field of avian biology.
For his outstanding work he received
his D.Sc. from the University of London and
in 1945 was awarded the Flavelle medal of
the Royal Society of Canada, Canada's
highest honour in the field of Science.
William Rowan's ornithological inte-
rests were by no means confined to migration
One of his earliest field studies was on the
ecology and breeding behavior of Merlins
(Faleo oolumbarius) on the Yorkshire moors.
During his early years in Alberta he con-
tributed to "British Birds" a series of
"Notes on Alberta Waders Included in the
British List". These, as many other of his
scientific and popular papers, were profuse-
ly illustrated with his delightful pen-and-
ink ske tches .
2
