Table Of ContentPERSPECTIVES
IN
EXPERIMENTAL BIOLOGY
Volume 1 Zoology
Proceedings of the Fiftieth Anniversary
Meeting of the Society for Experimental
Biology
Edited by P. SPENCER DAVIES
With the editorial assistance of
A. PUNT, G. M. HUGHES, S. H. P. MADDRELL,
J. E. TREHERNE, D. B. SATTELLE,
A. P. M. LOCKWOOD, J. D. ROBERTSON
and E. R. TRUEMAN
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First edition 1976
Library of Congress Catalog Card No. 75-34712
Printed in Great Britain by A. Wheaton & Co., Exeter
ISBN 0 08 018767 6 (Volume 1)
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LIST OF CONTRIBUTORS
ALEXANDER, R. MCN., Department of Pure and Applied Zoology, University of Leeds.
ANDERSEN, SVEND OLAV, August Krogh Institute, University of Copenhagen, Denmark.
ANWYL, R., Department of Zoology, The University, Glasgow, U.K.
ATKINSON, R. J. Α., Department of Marine Biology, University of Liverpool, Port Erin,
Isle of Man, U.K.
BATESON, P. P. G.,' University of Cambridge, Sub-Department of Animal Behaviour,
Madingley, Cambridge, U.K.
ΒΕΝΝΕΤ-CLARK, H. C, Department of Zoology, University of Edinburgh, Edinburgh,
U.K.
BITENSKY, LUCILLE, Division of Cellular Biology, The Mathilda and Terence Kennedy
Institute of Rheumatology, Bute Gardens, London, W6 7DW, U.K.
BOWLER, K., Department of Zoology, University of Durham, Durham, U.K.
BROWN, H. MACK, Department of Physiology, University of Utah Medical Centre,
Salt Lake City, Utah, U.S.A.
BURROWS, MALCOLM, Department of Zoology, University of Oxford, U.K.
BURTON, R. F., Institute of Physiology, University of Glasgow, Glasgow, U.K.
CHAYEN, J., Division of Cellular Biology, The Mathilda and Terence Kennedy Institute
of Rheumatology, Bute Gardens, London, W6 7DW, U.K.
CLARK, R. B., Department of Zoology, University of Newcastle upon Tyne, U.K.
CRISP, D. J., N.E.R.C. Unit of Marine Invertebrate Biology, Marine Science Laboratories,
Menai Bridge, U.K.
CROGHAN, P. C, School of Biological Sciences, University of East Anglia, U.K.
DAVIES, P., Division of Cell Pathology, M.R.C. Clinical Research Centre, Watford Road,
Harrow, Middx., U.K.
DORSETT, D. Α., Marine Science Laboratories, University College of North Wales,
Bangor, U.K.
DUNCAN, C. J., Department of Zoology, University of Liverpool, U.K.
FLETCHER, C. R., Department of Pure and Applied Zoology, University of Leeds, U.K.
GOLDSWORTHY, G. J., Department of Zoology, University of Hull, U.K.
ix
χ LIST OF CONTRIBUTORS
GUPTA, BRU L., Department of Zoology, Downing Street, Cambridge, U.K.
HUDDART, HENRY, Department of Biological Sciences, Lancaster University, Lancaster,
U.K.
HUGHES, G. M., Research Unit for Comparative Animal Respiration, Bristol University,
Bristol, U.K.
HUTCHISON, J. B., MRC Unit on the Development and Integration of Behaviour,
University Sub-Department, Madingley, Cambridge, U.K.
JOHANSEN, KJELL, Department of Zoophysiology, University of Aarhus, DK-8000
Aarhus C, Denmark.
LOCKWOOD, A. P. M., Department of Oceanography, University of Southampton, U.K.
MADDRELL, S. H. P., A.R.C. Unit of Invertebrate Chemistry and Physiology, Cambridge,
U.K.
MAETZ, J., Departement de biologie du C.E.A., Station Zoologique, 06230 Villefranche
sur Mer, France
MEECH, ROBERT W., A.R.C. Unit of Invertebrate Chemistry and Physiology, Depart-
ment of Zoology, Cambridge, U.K.
MILLER, PETER L., Department of Zoology, Oxford, U.K.
MILLS, PAULINE S., Department of Zoology, Oxford, U.K.
NAYLOR, E., Department of Marine Biology, University of Liverpool, Port Erin, Isle of
Man, U.K.
PAYAN, P., Departement de biologie du C.E.A., Station Zoologique, 06230 Villefranche
sur Mer, France.
PHILLIPS, J. E., Department of Zoology, University of British Columbia, Vancouver 8,
B.C., Canada.
PICHON, YVES, Unite de physiologie de l'Insecte, Laboratoire de Neurobiologie Cellulaire,
Centre National de la Recherche Scientifique, 91 190 Gif sur Yvette, France.
PIDDINGTON, R. W., A.R.C. Unit of Invertebrate Chemistry and Physiology, Department
of Zoology, University of Cambridge, Downing Street, Cambridge, U. K.
POTTS, W. T. W., Department of Biological Sciences, University of Lancaster, U.K.
PRINCE, W. T., A.R.C. Unit of Invertebrate Chemistry & Physiology, Department of
Zoology, University of Cambridge, U.K.
RENZIS, G. DE, Departement de biologie du C.E.A., Station Zoologique, 06230 Ville-
franche sur Mer, France.
SCHOFFENIELS, E., Laboratory of General and Comparative Biochemistry, University of
Liege, 17 Place Delcour, B-4000 Liege, Belgium.
LIST OF CONTRIBUTORS xi
SHELTON, G., School of Biological Sciences, University of East Anglia, Norwich, U.K.
SLATER, T. F., Biochemistry Department, Brunei University, Uxbridge, Middlesex, U.K.
SLEIGH, MICHAEL Α., Department of Biology, University of Southampton, U.K.
SMITH, C. L. Department of Zoology, University of Liverpool, U.K.
SMITH, R. I., Department of Zoology, University of California, Berkeley, California,
94720, U.S.A.
SUTCLIFFE, D. W., F.B.A., Windermere, U.K.
TREHERNE, J. E., A.R.C. Unit of Invertebrate Chemistry and Physiology, Department of
Zoology, Cambridge, U.K.
TRUEMAN, E. R., Zoology Department, University of Manchester, U.K.
USHERWOOD, P. N. R., Department of Zoology, University of Nottingham, Nottingham,
U.K.
WAREHAM, A. C, Department of Physiology, Medical School, University of Manchester,
Manchester, U.K.
WEBB, J. E., Department of Zoology, Westfield College, London, U.K.
WEBER, ROY E., Department of Zoophysiology, University of Aarhus, DK-8000 Aarhus
C, Denmark.
WEIS-FOGH, TORKEL, Department of Zoology, Downing Street, Cambridge, U.K.
WELLS, G. P., Department of Zoology, University College, Go wer Street, London, U.K.
WELLS, M. J., Zoology Department, University of Cambridge, U.K.
PREFACE
The Fiftieth Anniversary of the inauguration of the Society for Experimental Biology
was celebrated by a special meeting held at the University of Cambridge from July 16th
to 19th, 1974. Prominent members of the Society were each invited to present a paper on
their own specialized field of research, providing a historical review, a statement of
current thought in that area and, if possible, a predictive element. In the belief that the
material thus presented would be of considerable use to other research workers, the
speakers were invited to contribute their papers to these two volumes. The chapters
therefore are each a personal affirmation, and reflect the approaches, opinions and styles
of their individual authors.
For convenience, the papers have been collected into one zoological and one botanical
volume. Within each volume papers on a particular discipline have been grouped together,
although these are not necessarily the groupings that were used when the papers were
read at the meeting. This perhaps underlines the difficulty of classification in such a
diversified field as is here presented. However, this diversity gives an indication of the
breadth of the Society's interests and of the topics which tend to appear most frequently
in its programmes. The diversity too, emphasizes the strength and aims of the Society;
to provide a forum for research workers in both animal and plant sciences to discuss
current research, to present work which is often still in progress and incomplete, and
above all, to encourage the cross-fertilization of ideas between specialized disciplines.
It is hoped that the material assembled in these two volumes will provide for young
research workers a perspective of the field of experimental biology and for under-
graduates a source of reference to a wide range of biological topics.
Several invited speakers were unable to provide a manuscript and we wish to record
our thanks to them for their contribution to the Anniversary meeting. We are also
indebted to those distinguished members of the Society who acted as chairmen to the
sessions of the meeting. They were T. Weis-Fogh, R. Brown, G. M. Hughes, A. Punt,
P. W. Brian, Sir Vincent Wigglesworth, J. Chayen, R. D. Keynes, F. R. Whatley, E. W.
Simon, G. E. Fogg, J. Heslop-Harrison, J. W. S. Pringle, Sir Rutherford Robertson,
J. S. Kennedy, L. C. Beadle, D. A. Boulter, G. P. Wells, J. D. Robertson and Helen K.
Porter. Finally, particular thanks are due to the local organizing team: Helen Skaer,
D. A. Hanke, C. G. Gill and the able Local Secretary, D. B. Sattelle.
Ν. SUNDERLAND, P. SPENCER DAVIES,
John Innes Institute, Department of Zoology,
Colney Lane, The University,
Norwich, U.K. Glasgow, U.K.
xiii
T HE EARLY DAYS OF T HE S.E.B.
G. P. WELLS
Department of Zoology, University College* Gower Street, London, U.K.
THE Society for Experimental Biology was called into being in December 1923, to
support a Journal, then known as the British Journal of Experimental Biology.
The Society has always had botanists among its members, but in its early years it was
dominated by zoologists. The first published Part of the Journal contained six papers,
all zoological. The programme of the Conference at which the Society was set up,
included twenty papers, only three of which were botanical. Later, the botanical side
grew very rapidly until it came to match the zoological, but in the beginning it was the
younger zoologists who most urgently needed a Society, and a Journal, to cater for the
experimental aspects of their subject. The reason is that a great darkness had settled on
the majority of British zoologists in the early years of this century. They became obsessed
by comparative anatomy and descriptive embryology, and by the possible evolutionary
relationships of the animals whose corpses they studied.
Lancelot Hogben (1970) has traced for us the slow reawakening of experimental
zoology in Britain in the years before World War I, and its more vigorous expansion in
the early twenties. But even in the mid-twenties, there were still many centres of learning
in which the idea that zoology could be an experimental science had hardly dawned.
A fair way of showing what the word 'Zoology' then meant, to most of the people
who used it, is to quote the syllabus of the B.Sc. courses in Zoology in London in 1924.
London, besides being a great university, was, then more than now, a great examining
body, controlling, through its External Degree examinations, a number of important
teaching centres that have since become Universities in their own right. The syllabus that
I am about to sketch out determined the form and content of the subject for hundreds of
students, many of whom were destined to become teachers in their turn, so its influence
extended far and wide.
Suppose then that, fifty years ago, you were interested in zoology, and set out to get a
London B.Sc. degree. You could take a General or a Special B.Sc. Either would keep you
busy for at least 2 years after you had passed the Intermediate Examination. The General
Degree Course consisted of three science subjects of equal weight, one of which, in your
case, would be zoology. The Special Degree Course would consist mainly of intensive
teaching in zoology, with a lesser amount of one other science subject : its syllabus was
an elaboration of that for the General Course.
Here is what you must master, if you wanted to pass in zoology at the General level in
1924:
The main facts and principles of Zoology as exemplified by a study of the characteristic structure
and development of typical members of all the larger groups of the animal kingdom, with specia
1
2 G. P. WELLS
reference to those contained in the syllabus of practical work which follows; the chief facts and the
theories of heredity and evolution; the classification of the animal kingdom and the distinguishing
characters of the larger groups.
Then follows the list of types which is to form the basis of the practical course. It
consists of 258 named animals. Slightly more than half of these are distinguished by
asterisks; this 'signifies external form or demonstration specimen only'. The remaining
121 you must know pretty well. The types range from Amoeba to Man, but Man has an
asterisk. In addition to this, you must know twenty-three named larval forms, and the
early stages in the development of Amphioxus, frog, chick and rabbit.
This ends the syllabus.
If you aspired to Special Honours in zoology you had to know all this,
together with a more extended study of the more modern aspects of zoological work with special
reference to (1) Evolution and its evidences, (2) Bionomics, (3) Heredity (Genetics), (4) Experimental
Morphology and Embryology, (5) the meaning of sex and the mechanism of its determination.
You also studied a limited topic, known as a Special Subject, 'as far as possible from
original publications and actual material'. This topic you chose in consultation with
your teachers.
Notice that these courses contained no physiology, no behaviour, very little ecology
unless there was some in your Special Subject. Even those parts of the prescribed course
which are essentially experimental—genetics and experimental embryology—were
usually taught by lectures alone. In his practical examination, the candidate was required
to do three things : to dissect animals, to make sections and other permanent preparations,
and, confronted by rows of specimens, to indentify, draw and classify them, 'giving your
reasons'.
Meanwhile, the Botany students had a much better time. Their syllabus included
substantial amounts of physiology and ecology; their material lived; they made experi-
ments in the teaching laboratory and in the examination room. But animal physiology
stood apart from zoology and was taught as a separate subject with a strong medical
bias. In those days it was generally restricted to the mammal, coupled with the central
nervous system, heart and hind legs of the frog.
This was the situation from which the experimentally-minded zoologists were breaking
away.
One of the most active centres of the new movement was in Cambridge, and its leader
was James Gray. Though not yet Professor, he became, in Hogben's words, 'the mentor
of a new generation of Cambridge zoologists'. Others, too, were experimenting in the
zoology department—notably Munro Fox and J. T. Saunders—and experimental
questions began to appear in both written and practical examination papers. Comparative
physiology was also attracting attention elsewhere—in the Department of Physiology
for example, where Joseph Barcroft was extending his studies on haemoglobin to
invertebrates. One can learn something of the extent and variety of the movement by
turning over the pages of the first volume of the Biological Proceedings of the Cambridge
Philosophical Society, a volume which covered the 2 years from 1923 to 1925. It contains
contributions in other fields—in palaentology and botany, for example—but nearly two
thirds of its twenty-nine papers are about comparative physiology and experimental
THE EARLY DAYS OF THE S.E.B. 3
zoology. This, of course, is not because the followers of these lines of enquiry were more
numerous, or more productive, than their colleagues. It is because the workers in other
fields had other journals in which they could readily publish. It was one of the chief
reasons why the Biological Proceedings was started, to provide an outlet for the experi-
mentally-minded zoologists.
However, it was not in Cambridge that our Society began. We were conceived in
Edinburgh and born in London, at Birkbeck College. On the other hand, the British
Journal of Experimental Biology, for whose support we were created, was born in Edin-
burgh but conceived in London—more precisely, in the head of Hogben, on whose
first-hand account (1970) I have based my narrative of these events.
Lancelot Hogben, a man of tremendous energy and breadth of vision, was a Cam-
bridge graduate who became, for a few years, a lecturer at the Royal College of Science
in South Kensington. Here he had what he has modestly described as 'a by no means
epoch-making flirtation with genetical cytology'—a flirtation that resulted in six papers
and an invitation to move to Edinburgh.
The invitation came from Frank Crew, an Edinburgh graduate of medicine with a
strong interest in genetics, who had abandoned medical practice after demobilization
from World War I to teach zoology in his old University. Presently there was set up in
Edinburgh a new institute—the Animal Breeding Research Department—and Crew
became its first director in 1921. He was anxious to have a cytologist around. So Hogben
was invited to go north, and joined Crew as his second-in-command. By that time he was
turning from cytology to endocrinology and the physiology of colour change, and he
combined his position at the Animal Breeding Research Station with a part-time
lectureship in Comparative Physiology in the Department of Zoology.
While still at the Royal College of Science, Hogben had begun to think of a biological
journal for the publication of experimental research. When he reached Edinburgh, he
found Crew's Department, in his own words 'temporarily housed by Crew's inspired
opportunism in a dilapidated and abandoned Infirmary in a back street of Old Edin-
burgh'. But, in spite of its shabbiness, many distinguished biologists came for working
visits. Among them were Julian Huxley and J. B. S. Haldane, and these two completed
the quartet of, in Hogben's phrase, 'the Founding Fathers of the S.E.B.'.
With the other three, Hogben discussed his idea for a new Journal. Crew at once
declared that he had in hand enough cash from compensation for war wounds to finance
the initial project, and that he would be delighted to use it in this way. For other reasons,
and not because of his generous offer, the other three prevailed on Crew to become the
first Managing Editor and a supporting Editorial Board of eleven names was organized,
including a Professor of Botany, Ruggles Gates. Crew persuaded Oliver & Boyd to print
and publish the new journal in Edinburgh. Hogben undertook the circularization of
libraries, University departments, and Institutes. Finally, in October 1923, the first Part
of the British Journal of Experimental Biology saw the light.
Crew was then 35 years old, Huxley 36, Haldane 31 and Hogben 28.
It seemed to the Founding Fathers that a Society was needed, as a secure base for the
B.J.E.B. If one belonged to such Societies as the Physiological, the Biochemical or the
Linnaean in those days, one paid a handsome subscription and automatically received
the Society's journal, which could thus rely on financial support. Moreover, one generally
4 G. P. WELLS
sent one's manuscripts to one's journal. To provide the Β J.E.B, with these advantages,
the Founding Fathers (overcoming some resistance from other members of the Editorial
Board) announced a Conference to inaugurate the new Society, to take place at Birkbeck
College in the Christmas vacation, 1923. The invitations went to University departments
of botany, zoology and physiology and to research institutes, and in the event the
attendance exceeded the most extravagant hopes of the promoters.
That inaugural conference, as anybody who was there will remember, was a tremend-
ously exciting occasion. From then on, the success of the S.E.B, was assured—except in
one respect. It had been invented to give support to the B.J.E.B., and this it failed to do.
Paradoxically perhaps, its failure was due to the peculiar virtues which ensured its
success as a Society.
These virtues were two. In the first place, the great majority of its members were
young. In the second place, its interests were not restricted to a single specialized field.
It brought together experimental biologists from a range of different specializations, to
bring about, in Hogben's phrase, 'the cross-fertilization between widely separated
compartments of biological enquiry'. But the young were not opulent; the members of
specialist societies who joined us were already paying subscriptions elsewhere; and so it
was decided that one could belong to the S.E.B, for 12s. 6d. per annum. If, in addition,
one took the journal, the subscription was £2. But it turned out that only a minority of
members chose the latter option.
So what was to be done, to ensure the financial stability of the B.J.E.B. ? Its difficulties
were increased by the appearance, 2 months before its first issue came out, of another
journal about which I have already spoken—the Biological Proceedings of the Cambridge
Philosophical Society. Experimental zoology certainly needed an outlet, but not, at that
stage, two. So there were conversations, and meetings, and a solution was found. The
B.J.E.B. had included from the first, besides papers reporting on individual researches, a
certain proportion in which the writers reviewed recent progress in their fields. This
function was taken over by the Cambridge journal, which became Biological Reviews, and
under the editorship of Munro Fox, at once achieved international importance. Papers
on particular experimental researches became the business of the B.J.E.B.
But still the financial position of the B.J.E.B. was insecure, until the problem was
solved by the intervention of G. P. Bidder. He was a Cambridge zoologist with sub-
stantial private means and business interests, an expert on sponges, a familiar figure at
Naples and at Plymouth, who had assisted progressive developments in the biological
sciences more often, I suspect, than is generally known. His contribution to our story
was the invention of the Company of Biologists Limited, as a stable financial foundation
for the journal. He induced forty or fifty biologists to take shares in the Company, and
himself guaranteed any overdraft up to a sufficient limit. With the funds so raised, the
B.J.E.B. was bought from Oliver & Boyd, and thereafter was published for the Company
by the Cambridge University Press. The Company appoints the Editor of the journal,
but does not meddle with what he does. Wisely, they persuaded James Gray to undertake
the Editorship. Their business is managed by a Board of Directors, consisting at first of
Crew, Huxley, Hogben, Saunders and Bidder, who was also the Secretary. Bidder
resigned from the Secretaryship after 3 years, to be replaced by Saunders. The word
British' was later dropped from the title of the Journal, because of a short-lived plan to
