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Circadian Rhythmicity CIRCADIAN RHYTHMICITY Proceedings ofth e International Symposium on Circadian Rhythmicity, Wageningen, theNetherlands , 26-29 April 1971 Centre for Agricultural Publishing and Documentation, Wageningen 1972 S:/ BIILIOTHEEB PER y^NDBOUWHOGESCHeet. WAGENINGEN ISBN 9022 00356 6 © Centrefo r AgriculturalPublishin g andDocumentatio n (Pudoc),Wageningen , 1972 No part ofthi sboo k mayb ereproduce d and/or published in anyform , byprint ,pho ­ toprint, microfilm or any other meanswithou t written permission from thepublishers . Printed in the Netherlands by H. Veenman &Zone n N.V., Wageningen. The Organizing Committee J. F. Bierhuizen, Chairman G. W. Ankersmit, Secretary M. van Albada Miss B. Baggerman W. H. van Dobben A. M. Frens H. Klomp P. J. A. L. de Lint E. C. Wassink J. de Wilde Preface Once every three years the Agricultural University of Wageningen organizes a sym­ posium on a fundamental aspect of natural science. In 1971 the topic was Orcadian Rhythmicity. Its influence in biology ranges from thewhol e organism to the subcellular level. The symposium attempted to cover all these aspects. Diurnal rhythms were first observed long ago. In plants, for instance, diurnal movements of leguminous plants were noticed in the time of Alexander the Great. A century ago such rhythms were observed in animals, and interest in rhythmicity in man is now rapidly increasing with the advent of jets and space flight. Circadian rhythms are very important in ecology. Many organisms adapt themselves to cyclic external conditions such as day and night, and seasonal trends. Even tidal movements and moon phases seem important in terrestrial life. There is also a close interrelation between various species in a community. Production of pollen and nectar, and opening and closing of flowers are often diurnal in rhythm. Flight of bees is syn­ chronized with such plant rhythms. Some circadian rhythms may have great influence on survival, since they may synchronize activities with the most favourable moment of the day or season. A circadian rhythm is endogenous when rhythms continue for at least a few days under constant environmental conditions. Research has advanced rapidly since it became possible to control environmental conditions. Temperature changes of 1 °C and radiation for only a few seconds are now known to modify results. Besides such close control of the environment, research has been promoted by the precision, with which rhythms can be measured, by the sophistication of mathematical models, and by use of the computer. This book records the proceedings of a symposium on Circadian Rhythmicity held in Wageningen, the Netherlands, 26-29 April 1971 during which many new aspects were discussed of circadian rhythms in plants and animals at the cellular and subcel­ lular level, kinetic models, biochemical processes and the effect of external factors on rhythms. We thank the Board of the University for financial support, our contributors for their excellent reviews, the International Agricultural Centre for technical assistance and Pudoc for publishing the proceedings. J. F. Bierhuizen Contents E. Bünning, Symptoms, problems and common features of circadian rhythms in plants and animals 11 B. G. Cumming, The role of circadian rhythmicity in photoperiodic induction in plants t . 33 A. D. Lees, The role of circadian rhythmicity in photoperiodic induction in animals 87 J. W. Truman, Circadian rhythms and physiology with special reference to neuroendocrine processes in insects Ill B. M. Sweeney, Circadian rhythms in unicellular organisms 137 H. G. Schweiger, Circadian rhythms: subcellular and biochemical aspects . 157 K. Hoffmann, Biological clocks in animal orientation and in other functions. 175 E. C. Wassink, Concluding remarks: botanical aspects 207 J. de Wilde, Concluding remarks: zoologica l aspects 210 Proc. int. Symp. circadian Rhythmicity (Wageningen, 1971) 11-31 Symptoms, problems, and common features of circadian rhythms in plants and animals E. Bünning Institut für Biologie der Universität Tübingen, Tübingen, West Germany Introduction In my Opening Address at the 1960 Cold Spring Harbor Symposium on 'Biological Clocks', I summarized the history of research on endogenous diurnalrhythms . I will not therefore repeatmyself ,excep tt omentio n onepiec eo fhistory/Tha t isth ewor ki n the Netherlands of Anthonia Kleinhoonte. Kleinhoonte published her thesis in 1928 inDutc han dpublishe d worki nGerma n ayea rlate rabou tth etim eI becam eintereste d in theseproblems . Sheworke d in the Laboratory for Technical Botany inDelft , using the same dark room that Brouwer had used a few years earlier (1926) and the same species of plant. Brouwer believed heha d shown the control of thediurna llea fmove ­ ments by some unknown external factor. He was not the only biologist considering biological rhythms to be synchronized by a factor X, ascribed variously to electric charges in the air, cosmic rays and radioactive radiations. History credits Kleinhoontefo r showing that it isonl ynecessar y to improve control of the well known external factors such as temperature and light for the endogenous nature of thecircadia n rhythms tobecom eclear . Kleinhoonte showed that even pulses of weak light, not more than a minute long, effectively cause delays or advances of circadian cyclesan d synchronize them. These wereals o thefirs t steps in studyingwha t isno w known as phase-response curves of circadian rhythms. Unfortunately, even in later work, authors came to wrong conclusions by ignoring the synchronizing effects of weak control lights and other factors. When, under so- called constant conditions, we find an accumulation of physiological peaks around certain hours of the day or an exact 24-h periodicity, weshoul d ask ourselves: wher e is the mistake in the experimental conditions? (Fig. 1) I raised this question when I began work together with Kurt Stern in 1928.W e found that the mistake underlying previousresearc h wasth e overlooking ofth e synchronizing effect ofver ywea k control light. Recent evidence against control of rhythms by unknown external factors comes from Enright's (1965) careful analysis of several published experiments. His analysis gave no support for a role of subtle geophysical factors in synchronization. It was also left to Kleinhoonte to draw attention to earlier papers by Pfeffer. His publications from 1907-15 were neglected in papers on diurnal periodicity in plants 11 Fig. 1.Exampl e of experiments believed to showth esynchronizin g effect of 'factor X'. Diurnal leaf movements of Phaseolus in a darkroom. Night-peaks (maximum of dawnward movement) mostly occur about 3h after midnight in spiteo f 'constant con­ ditions'. After experiments of Stoppel (1916).Th e 'factor X' waslate r shownt o be weak red control light, used in the pre­ ceding morning hours to start the experi­ ment, and to water the plants. 6-8 8-10 10-12 12-K 14-16 16-18 18-20 20-22 22-2« TIME OF DAY and animals published before Kleinhoonte's work. Actually Pfeffer clearly demon­ strated an endogenous component in circadian leaf movements. His only mistake was to publish his mass of results, filling about 500 pages, in Abhandlungen der mathema­ tisch-physischen Klasse der Königlich Sächsischen Gesellschaft der Wissenschaften. This sounds very distinguished and aristocratic, but it is, as we say in Germany, 'ein Staats­ begräbnis erster Klasse' (a first class state funeral). Terminology and some basic phenomena Sometimes, any type of 24-h cycle iscalle d a circadian cycle.However , thister m should be restricted to cycles, which continue in constant temperature and in the absence of diurnal light-dark cycles. Under these conditions, circadian cycles are no longer exactly 24-h periods but vary between about 22 and 28 h and usually between 23 and 25 h. This period varies not only from species to species, but also from one individual plant or animal to the other. Only a few cases are known, where it becomes difficult to find significant deviations from the exact 24-h period when diurnal temperature and light- dark cycles are excluded. Two examples are the fiddler crab Uca and the Canavalia studied by Kleinhoonte. However many strictly exogenous diurnal physiological cycles do occur. These should be called 'daily rhythms' or '24-h rhythms', but not 'circadian rhythms' (Wurtman, 1966). It is not always easy to detect the participation of a circadian rhythrnicity in a daily rhythm. Continuous light or darkness are often so strongly disturbing or damaging to the organism, that the overt rhythrnicity disappears immedi­ ately. If, for example, bean plants are brought from the greenhouse into continuous darkness or continuous incandescent light, the circadian leaf movements will no longer continue. This is due to the far-red component of that light. Therefore the earlier experiments with beans were always carried out with plants grown in dark rooms, and thus adapted to this condition. Nowadays this is no longer a problem: greenhouse plants continue with their circadian leaf movements for their whole life time in the light of fluorescent tubes which lacks far-red. 12 A similar problem may occur with animals. Sometimes the circadian rhythmicity disappears quickly after bringing the animals into continuous darkness or continuous light. Then the application of continuous dim light can be advantageous. Diversity of circadianrhythm s It is tempting to assume an identical cellular basis for all circadian rhythms, from unicellular organisms up to higher plants and animals. However, this hypothesis isno t substantiated byth efacts .W ekno wabou tman ydevelopmenta l cyclesshowin gperiod s of about 24h which apparently are self sustaining cycles in the sense, that each of the several morphological steps is comparable with part of the mechanism of a clock. Mitotic cyclesca n also havefeature s of developmental cycles. Contrary to this, in the 'classical' cases of circadian rhythms, the cycles are maintained without overt mor- phogenetical steps. Microscopic or submicroscopic changes can be associated with the running of the circadian clock.
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