Adult eclosion rhythm of the Indian meal moth Plodia interpunctella: response to various thermocycles with different means and amplitudes

In addition to photoperiod, thermoperiod (or thermocycle) might be an important Zeitgeber for entraining the circadian oscillator controlling adult eclosion rhythm in the Indian meal moth Plodia interpunctella Hübner (Lepidoptera: Pyralidae). This is confirmed by exposing larvae receiving diapause‐preventing treatments to various thermocycles with different means and amplitudes of temperature. The thermocycles investigated in the present study are TC 8 : 16 h, TC 12 : 12 h, TC 16 : 8 h and TC 20 : 4 h, where T and C represent thermophase (30 °C) and cryophase (20 °C), respectively. For all thermocycles, the peak of adult eclosion rhythm occurs at around the mid‐thermophase. This indicates that the larvae use both ‘temperature‐rise’ and ‘temperature‐fall’ signals to adjust the eclosion phase in each thermocycle. The absence (DD) or presence (LL) of light affects this time‐keeping system slightly under the given thermocycle. The rhythmic adult eclosion noted after exposure of larvae to 30 °C DD for 14 days is recorded in the thermocycles (TC 12 : 12 h, DD; mean temperature = 25 °C) with different amplitudes of 27.5/22.5 °C, 26.5/23.5 °C and 25.5/24.5 °C. The peak in adult eclosion advances in time as the amplitude of the temperature cycle decreases. In the temperature cycle of 25.5/24.5 °C, a peak occurs at the end of the cryophase, 2 h before the temperature‐rise. The adult eclosion rhythm is also observed under various thermocycles (TC 12 : 12 h, DD) consisting of different temperature levels (30 to 20 °C) with different amplitudes. It is found that the temporal position of the peak advances significantly when the amplitude of the thermocycle becomes lower.     

Effects of photoperiod and temperature on the rhythm and free‐running of adult eclosion in the Indian meal moth Plodia interpunctella

The rhythm of adult eclosion in the Indian meal moth Plodia interpunctella Hübner (Lepidoptera: Pyralidae) is investigated under various photoperiods and temperatures aiming to determine the nature of the temperature compensation and the free‐running period. Insects that are committed to a nondiapause larval development show diel rhythms of adult eclosion at 30, 25 and 20 °C. At 30 °C, the eclosion peak (i.e. the mean time of eclosion) occurs approximately 20 h after lights off under an LD 4 : 20 h photocycle, and at approximately 15 h under an LD 20 : 4 h photocycle. At 25 °C, the peak of eclosion occurs approximately 19 h after lights off under an LD 2 : 20 h photocycle and at approximately 16 h under an LD 20 : 4 h photocycle. At 20 °C, the eclosion peak is significantly advanced under long days of >12 h (i.e. approximately 20 h after lights off under an LD 4 : 20 h photocycle and approximately 9 h under an LD 20 : 4 h photocycle), indicating an effect of both lights‐off and lights‐on signals on the timing of the adult eclosion. To determine the involvement of a self‐sustained oscillator, the rhythm of adult eclosion is examined under darkness at different temperatures (30 to 21 °C). The mean free‐running periods are 22.4, 22.8, 22.0 and 22.5 h at 30, 24, 23 and 22 °C, respectively, indicating that the eclosion rhythm is temperature‐compensated. However, this rhythm does not free‐run under constant darkness at 21 °C. Because a clear diel rhythm is observed under 24‐h photocycles at 20 °C, the oscillator might be damped out within 24 h at the lower temperature.     

Effects of background light conditions on thermoperiodic eclosion rhythm of onion fly Delia antiqua

To elucidate the effects of light on thermoperiodic regulation of adult eclosion rhythm in the onion fly, Delia antiqua, the responses to two thermoperiods, 29°C (12 h):21°C (12 h) and 25.5°C (12 h):24.5°C (12 h), with different amplitude and same average temperature, were examined in continuous darkness (DD) and continuous light (LL). Irrespective of the temperature step between warm phase (W) and cool phase (C), temperature cycles effectively entrained the adult eclosion rhythm in both DD and LL. Eclosion peaks, however, varied with light conditions and temperature step between W and C. It advanced by approximately 2–3 h in DD than in LL and at smaller temperature step. Background light conditions and temperature step also affect the amplitude of eclosion rhythm. It became lower in LL than in DD and at smaller temperature steps. On transfer to constant temperature (25°C), eclosion rhythm was elicited earliest in the pupae at 8°C temperature step in DD and latest in those at 1°C temperature step in LL. Pupae at 1°C temperature step in DD and at 8°C temperature step in LL demonstrated intermediate responses, but the eclosion rhythm was elicited 1 day earlier in the former than in the latter. This might be ascribed to the interaction between background light and temperature step under thermoperiodic conditions. The results suggest that continuous light and a smaller temperature step weaken the coupling strength between eclosion rhythm and thermoperiod, but the light effect is stronger than the temperature step effect.     

Interacting effect of thermoperiod and photoperiod on the eclosion rhythm in the onion fly, Delia antiqua supports the two-oscillator model

Daily light and temperature cycles entrain adult eclosion rhythms in many insect species, but little is known about their interaction. We studied this problem in the onion fly, Delia antiqua. Pupae were subjected to various combinations of a photoperiod of 12L:12D and thermoperiods. The thermoperiods consisted of 12 h warm phase (W) and 12 h cool phase (C), giving a mean temperature of 25 °C with different temperature steps of 8, 4 and 1 °C. As the phase relation of the two Zeitgebers was varied, the phase of eclosion rhythm was shifted, depending on the phase angle with the light cycle and the amplitude of the temperature cycle. When the temperature step in the thermoperiod was 8 °C (WC 29:21 °C), the eclosion rhythm was entrained mainly to thermoperiod rather than photoperiod. In the regime with a 4 °C temperature step (WC 27:23 °C), both thermoperiod and photoperiod affected eclosion rhythm, and a phase jump of the eclosion rhythm occurred when the warm phase of thermoperiod was delayed 15-18 h from light-on. In regimes with a 1 °C temperature step (WC 25.5:24.5 °C), the eclosion rhythm was completely entrained to photoperiod. The observed interacting effect of light and temperature cycle on the eclosion rhythm in D. antiqua can be explained by the two-oscillator model proposed by Pittendrigh and Bruce (1959).     

Effect of photoperiod and thermoperiod on the eclosion rhythm ofTrichogramma evanescens

The dynamics of adult eclosion inTrichogramma evanescens Westw. were studied under (1) constant light and temperature of 20 °C, (2) photoperiods L12:D12 and L16:D8 at 20 °C, (3) thermoperiod 4 h 27 °/20 h 20 °C at constant light, (4) joint action of photo- and thermoperiod. The emergence was arhythmic in constant light combined with constant temperature, while a sharp monophasic rhythm was observed with the isolated action of photo- or thermoperiod. The ‘light-on’ and ‘temperature step-up’ signals were shown to act in one direction. When both signals were combined, they manifested themselveses competing entraining stimuli which, in turn, revealed an apparent individual variation in relative reactivity to the light and temperature signals. Some perspectives which follow from these observations are discussed.     

Temperature cycle amplitude alters the adult eclosion time and expression pattern of the circadian clock gene period in the onion fly

Soil temperature cycles are considered to play an important role in the entrainment of circadian clocks of underground insects. However, because of the low conductivity of soil, temperature cycles are gradually dampened and the phase of the temperature cycle is delayed with increasing soil depth. The onion fly, Delia antiqua, pupates at various soil depths, and its eclosion is timed by a circadian clock. This fly is able to compensate for the depth-dependent phase delay of temperature change by advancing the eclosion time with decreasing amplitude of the temperature cycle. Therefore, pupae can eclose at the appropriate time irrespective of their location at any depth. However, the mechanism that regulates eclosion time in response to temperature amplitude is still unknown. To understand whether this mechanism involves the circadian clock or further downstream physiological processes, we examined the expression patterns of period (per), a circadian clock gene, of D. antiqua under temperature cycles that were square wave cycles of 12-h warm phase (W) and 12-h cool phase (C) with the temperature difference of 8 °C (WC 29:21 °C) and 1 °C (WC 25.5:24.5 °C). The phase of oscillation in per expression was found to commence 3.5 h earlier under WC 25.5:24.5 °C as compared to WC 29:21 °C. This difference was in close agreement with the eclosion time difference between the two temperature cycles, suggesting that the mechanism that responds to the temperature amplitude involves the circadian clock.     

Effects of Temperature,Photoperiod, and Light Intensity on the Eclosion Rhythm of the High-Altitude Himalayan Strain of Drosophila ananassae

Eclosion rhythm of the high-altitude Himalayan strain of Drosophila ananassae from Badrinath (altitude 5123 m) was temperature-dependent and at 21°C, it was entrained by cycles of 12 h light: 12 h darkness (LD 12:12) and free-ran in constant darkness, however, it was arrhythmic at 13°C or 17°C under identical experimental conditions (Khare, P. V., Barnabas, R. J., Kanojiya, M., Kulkarni, A. D., Joshi, D. S. (). Temperature dependent eclosion rhythmicity in the high altitude Himalayan strains of Drosophila ananassae. Chronobiol. Int. 19:1041–1052). The present studies were designed to see whether or not these strains could be entrained at 13°C, 17°C, and 21°C by two types of LD cycles in which the photoperiod at 100 lux intensity varied from 6 h to 18 h, and the light intensity of LD 14:10 cycles varied from 0.001 lux to 1000 lux. All LD cycles entrained this strain at 21°C but not at 13°C or 17°C. These results demonstrate that the entrainment of eclosion rhythm depends on the ambient temperature and not on the photoperiod or light intensity of LD cycles. Thus the temperature has taken precedence over the light in the entrainment process of eclosion rhythm of the high altitude Himalayan strain of D. ananassae. This may be the result of natural selection in response to the environmental temperature at Badrinath that resembles that of the sub-Arctic region but the photoperiod or light intensity are of the subtropical region.     

Multi-Oscillatory Control of Eclosion and Oviposition Rhythms in Drosophila melanogaster: Evidence from Limits of Entrainment Studies

The eclosion and oviposition rhythms of flies from a population of Drosophila melanogaster maintained under constant conditions of the laboratory were assayed under constant light (LL), constant darkness (DD), and light/dark (LD) cycles of 10:10 h (T20), 12:12 h (T24), and 14:14 h (T28). The mean (±95% confidence interval; CI) free-running period (τ) of the oviposition rhythm was 26.34 ± 1.04 h and 24.50 ± 1.77 h in DD and LL, respectively. The eclosion rhythm showed a τ of 23.33 ± 0.63 h (mean ± 95% CI) in DD, and eclosion was not rhythmic in LL. The τ of the oviposition rhythm in DD was significantly greater than that of the eclosion rhythm. The eclosion rhythm of all 10 replicate vials entrained to the three periodic light regimes, T20, T24, and T28, whereas the oviposition rhythm of only about 24 and 41% of the individuals entrained to T20 and T24 regimes, respectively, while about 74% of the individuals assayed in T28 regimes showed entrainment. Our results thus clearly indicate that the τ and the limits of entrainment of eclosion rhythm are different from those of the oviposition rhythm, and hence this reinforces the view that separate oscillators may regulate these two rhythms in D. melanogaster.     

Changes in temperature, not photoperiod, control the pattern of adult eclosion in the tsetse, Glossina morsitans

Abstract. The timing of adult eclosion in tsetse, an event that normally occurs in mid-afternoon, is regulated by the daily cycle of temperature elevation. If a temperature cycle is maintained, the rhythm of eclosion persists under continuous light or continuous darkness. Artificially shifting the temperature peak to the scotophase results in a concomitant shift in the eclosion pattern. Daily temperature variations as small as 0.4°C are sufficient to establish the rhythm. Eclosion activity tracks the temperature peak, even if the pulses are of short duration (4 h) or with irregular frequencies of 12 or 36 h. The temperature-induced rhythm offers a simple mechanism for separating females and males. Individuals that pupariate on the same day eclose as adults over a 4–5 day period at 25°C, and in such collections, females are the first to eclose. This distinction makes it possible to collect samples of predominately one sex, a feature that may facilitate the collecting of males for use in the sterile-male technique.     

Effects of temperature on the development of overwintering immature stages of the near‐threatened butterfly Leptalina unicolor (Bremer & Grey) (Lepidoptera: Hesperiidae)

Overwintering larvae of multivoltine and univoltine populations of Leptalina unicolor were reared under various constant and fluctuating temperatures superimposed on a photoperiod of either 12 h of light and 12 h of darkness (LD 12:12) or LD 15:9. Diapause of the larvae terminated in midwinter (by early February). All the larvae of both populations pupated after two molts without feeding and the head capsule width of the final instar larvae was smaller than that of the penultimate instar ones. The photoperiod did not significantly affect larval development, but long‐day conditions (LD 15:9) hastened pupal development. The thermoperiod had a significant effect on the development of the multivoltine population. When multivoltine population larvae were kept under a low fluctuating temperature regime (cryophase/thermophase = 14/20°C), the period until adult eclosion was shorter than that under a constant temperature of 17°C. On the contrary, when larvae were kept under a high fluctuating temperature regime (24/30°C), the period until adult eclosion was longer than that under a constant temperature of 27°C. However, the univoltine population did not show such a reaction to the fluctuating temperature. The durations of final instar larva and pupa of the multivoltine population were shorter than those of the univoltine population. The developmental zeros of penultimate and final instar larvae and pupae of the univoltine population were lower than those of the multivoltine population. The head capsule width of penultimate instar larvae and the forewing length of adults of the univoltine population were larger than those of the multivoltine population for both sexes.     

Comparison of the circadian eclosion rhythm between non-diapause and diapause pupae in the onion fly, Delia antiqua: the change of rhythmicity

When pupae of Delia antiqua were transferred to constant darkness (DD) from light-dark (LD) cycles or constant light (LL), the sensitivity to light of the circadian clock controlling eclosion increased with age. The daily rhythm of eclosion appeared in both non-diapause and diapause pupae only when this transfer was made during late pharate adult development. When transferred from LL to DD in the early pupal stage, the adult eclosion was weakly rhythmic in non-diapause pupae but arrhythmic in diapause pupae. However, the sensitivity of the circadian clock to temperature cycles or steps was higher in diapause pupae than in non-diapause pupae; in the transfer to a constant 20 degrees C from a thermoperiod of 25 degrees C (12 h)/20 degrees C (12 h) on day 10 after pupation or from chilling (7.5 degrees C) in DD, the adult eclosion from diapause pupae was rhythmic but that from non-diapause pupae arrhythmic. In a transfer to 20 degrees C from the thermoperiod after the initiation of eclosion, rhythmicity was observed in both types of pupae. The larval stage was insensitive to the effect of LD cycle initiating the eclosion rhythm. In D. antiqua pupae in the soil under natural conditions, therefore, the thermoperiod in the late pupal stage would be the most important 'Zeitgeber' for the determination of eclosion timing.     

Multi-Oscillatory Control of Eclosion and Oviposition Rhythms in Drosophila melanogaster: Evidence from Limits of Entrainment Studies

The eclosion and oviposition rhythms of flies from a population of Drosophila melanogaster maintained under constant conditions of the laboratory were assayed under constant light (LL), constant darkness (DD), and light/dark (LD) cycles of 10:10 h (T20), 12:12 h (T24), and 14:14 h (T28). The mean (±95% confidence interval; CI) free-running period (τ) of the oviposition rhythm was 26.34 ± 1.04 h and 24.50 ± 1.77 h in DD and LL, respectively. The eclosion rhythm showed a τ of 23.33 ± 0.63 h (mean ± 95% CI) in DD, and eclosion was not rhythmic in LL. The τ of the oviposition rhythm in DD was significantly greater than that of the eclosion rhythm. The eclosion rhythm of all 10 replicate vials entrained to the three periodic light regimes, T20, T24, and T28, whereas the oviposition rhythm of only about 24 and 41% of the individuals entrained to T20 and T24 regimes, respectively, while about 74% of the individuals assayed in T28 regimes showed entrainment. Our results thus clearly indicate that the τ and the limits of entrainment of eclosion rhythm are different from those of the oviposition rhythm, and hence this reinforces the view that separate oscillators may regulate these two rhythms in D. melanogaster.     

LIGHT AT NIGHT ALTERS THE PARAMETERS OF THE ECLOSION RHYTHM IN A TROPICAL FRUIT FLY,DROSOPHILA JAMBULINA

We investigated the effects of natural light at night (LAN) in the field and artificial LAN in the laboratory on the circadian rhythm of pupal eclosion in a tropical wild type strain of Drosophila jambulina captured at Galle, Sri Lanka (6.1^oN, 80.2^oE). The influence of natural LAN, varying in intensity from 0.004 lux (starlight intensity) to 0.45 lux (moonlight intensity), on the entrainment pattern of the circadian rhythm of eclosion at 25^o?±?0.5^oC was examined by subjecting the mixed-aged pupae to natural cycles of light and darkness at the breeding site of this strain in the field. The eclosion peak was ～2?h prior to sunrise, and the 24?h rhythmicity was the most robust. Effects of artificial LAN at 25^o?±?0.5^oC were determined in the laboratory by subjecting pupae to LD 12:12 cycles in which the light intensity of the photophase was 500 lux in all LD cycles, while that of the scotophase was either 0 lux (complete darkness, DD), 0.5, 5, or 50 lux. In the 0 lux LAN condition (i.e., the control experiment), the eclosion peak was ～2?h after lights-on, and the 24?h eclosion rhythm was not as strong as in the 0.5 lux LAN condition. The entrainment pattern in 0.5 lux LAN was strikingly similar to that in the field, as the 0.5 lux LAN condition is comparable to the full moonlight intensity in the tropics. LAN at 0.5 lux dramatically altered both parameters of entrainment, as the eclosion peak was advanced by ～4?h and the 24?h eclosion rhythm was better than that of the control experiment. LAN at 5 lux, however, resulted in a weak eclosion rhythm that peaked in the subjective forenoon. Interestingly, the 50 lux LAN condition rendered the eclosion events unambiguously arrhythmic. After-effects of LAN on the period (τ) of the free-running rhythm and the nature of eclosion rhythm were also determined in DD by a single LD 12:12 to DD transfer. After-effects of the LAN intensity were observed on both the τ and nature of the eclosion rhythm in all four experiments. Pupae raised in 0.5 lux LAN exhibited the shortest τ (20.6?±?0.2?h, N?=?11 for this and subsequent values) and the most robust rhythm, while pupae raised in 50 lux LAN had the longest τ (29.5?±?0.2?h) and weakest rhythm in DD. Thus, these results demonstrate the intensity of LAN, varying from 0 to 50 lux, profoundly influences the parameters of entrainment as well as free-running rhythmicity of D. jambulina. Moreover, the observed arrhythmicity in LD 12:12 cycles caused by the 50 lux LAN condition appeared to be the masking effect of relatively bright light at night, as the LD 12:12 to DD transfer restored the rhythmicity, although it was rather weak. (Author correspondence: drdsjoshi08@gmail.com)     

Entrainment of Eclosion Rhythm in Drosophila melanogaster Populations Reared for More Than 700 Generations in Constant Light Environment

In this paper, we report the results of our extensive study on eclosion rhythm of four independent populations of Drosophila melanogaster that were reared in constant light (LL) environment of the laboratory for more than 700 generations. The eclosion rhythm of these flies was assayed under LL, constant darkness (DD) and three periodic light‐dark (LD) cycles (T20, T24, and T28). The percentage of vials from each population that exhibited circadian rhythm of eclosion in DD and in LL (intensity of approximately 100 lux) was about 90% and 18%, respectively. The mean free‐running period (τ) of eclosion rhythm in DD was 22.85 ± 0.87 h (mean ± SD). Eclosion rhythm of these flies entrained to all the three periodic LD cycles, and the phase relationship (ψ) of the peak of eclosion with respect to “lights‐on” of the LD cycle was significantly different in the three periodic light regimes (T20, T24, and T28). The results thus clearly demonstrate that these flies have preserved the ability to exhibit circadian rhythm of eclosion and the ability to entrain to a wide range of periodic LD cycles even after being in an aperiodic environment for several hundred generations. This suggests that circadian clocks may have intrinsic adaptive value accrued perhaps from coordinating internal metabolic cycles in constant conditions, and that the entrainment mechanisms of circadian clocks are possibly an integral part of the clockwork.     

Thermal responses in the citrus fruit fly,Dacus tsuneonis: evidence for a pupal diapause

Thermal responses controlling pupariation and adult eclosion in a citrus fruit fly,Dacus tsuneonis (Miyake), were studied to understand the winter biology of this species. When mature larvae were exposed to various temperature conditions, the highest percentage of pupariation was obtained at 15 °C, although the variance at this temperature was greater than at 20 °C or 25 °C. Pupariation occurred most rapidly at 20 °C and an alternating temperature with a mean of 15 °C. At constant 15 °C, pupae failed to emerge as adults. Pupae were characterized by a reduced respiration rate, which is typical of a diapausing pupa. When insects were stored at different temperatures for 45 days after pupariation, and then transferred to 25 °C, adult eclosion occurred earlier when the initial temperature was 10 °C than when it was 5 °C or 15 °C. Adult eclosion occurred most synchronously and pupal mortality was lowest when insects were stored at 15 °C for 90 days before incubation at 25 °C. These results strongly suggest thatD. tsuneonis enters a pupal diapause.     

Comparison of the circadian eclosion rhythm between non-diapause and diapause pupae in the onion fly,Delia antiqua: the effect of thermoperiod

When non-diapause and diapause pupae of Deliaantiqua were exposed to various thermoperiods where thermophase (T) was 25 °C and the cryophase (C) was 15 or 20 °C (TC₁₅ or TC₂₀) in constant darkness (DD), the majority of both types of flies emerged before the rise in temperature. Eclosion time was delayed at the lower cryophase temperature. Moreover, there was a significant difference in the time of adult eclosion between non-diapause and diapause pupae; diapause pupae eclosed earlier than non-diapause pupae. When the two types of pupae were transferred to a constant low temperature (15 or 20 °C) after having experienced TC₁₅ or TC₂₀ 12:12 h, they showed circadian rhythmicity in eclosion. The free-running period (τ) of the eclosion rhythm changed after transfer to constant low temperatures in both non-diapause and diapause pupae, suggesting that this change represents a transient cycle until the temperature-sensitive oscillator is coupled again to the temperature-insensitive pacemaker. However, diapause pupae tended to show a shorter τ than non-diapause pupae. This observation suggests that the difference in adult eclosion time under thermoperiodic conditions between non-diapause and diapause pupae is related to their different τ s.     

Towards a Characterization of the Locomotor Activity Rhythm of the Supralittoral Isopod Tylos europaeus

Freshly collected samples of Tylos europaeus from Korba beach (northeast of Tunisia) were housed in an environmental cabinet at controlled temperature (18°C?±?.5°C) and photoperiod. Locomotor activity was recorded under two photoperiodic regimens by infrared actography every 20?min by multichannel data loggers. One regimen simulated the natural light-dark cycle on the day of collection, whereas the second imposed a state of continuous darkness on all individuals. Under entraining conditions, the animals displayed rhythmic activity, in phase with the period of darkness, whereas in continuous darkness these isopods exhibited a strong endogenous rhythm with circadian and semidiurnal components at mean periods of τ (h:min)?=?25:09?±?01:02?h and τ?=?12:32?±?00:26?h, respectively. Under free-running conditions, this endogenous rhythm showed significant intraspecific variability. (Author correspondence: bohli.dhouha@gmail.com)     

Effect of temperature on circadian rhythm controlling the crepuscular activity of the burying beetle Nicrophorus quadripunctatus Kraatz (Coleoptera: Silphidae)

Locomotor activity rhythm was examined at various temperatures under a 16 h light : 8 h dark photoperiod (LD 16:8) or LD 12:12 using adults of the burying beetle Nicrophorus quadripunctatus. At 20°C, the locomotor activity of the beetles showed a bimodal daily pattern with two peaks around lights on and lights off under both photoperiods. This bimodal activity rhythm persisted under constant darkness; therefore, the activity of adult N. quadripunctatus is controlled by a circadian clock. Adults showed a bimodal activity pattern for temperatures ranging from 15 to 25°C. The evening peak of the daily activity was earlier at lower temperatures. These findings suggest that in the field, N. quadripunctatus shows crepuscular activity, and is active earlier in the afternoon in cooler seasons. In this species, therefore, temperature appears to play an important role in the determination of daily activity patterns.     

The Circadian Body Temperature Rhythm in the Elderly: Effect of Single Daily Melatonin Dosing

The present study is part of a more extensive investigation dedicated to the study and treatment of age‐dependent changes/disturbances in the circadian system in humans. It was performed in the Tyumen Elderly Veteran House and included 97 subjects of both genders, ranging from 63 to 91 yrs of age. They lived a self‐chosen sleep‐wake regimen to suit their personal convenience. The experiment lasted 3 wks. After 1 control week, part of the group (n=63) received 1.5 mg melatonin (Melaxen?) daily at 22:30 h for 2 wks. The other 34 subjects were given placebo. Axillary temperature was measured using calibrated mercury thermometers at 03:00, 08:00, 11:00, 14:00, 17:00, and 23:00 h each of the first and third week. Specially trained personnel took the measurements, avoiding disturbing the sleep of the subjects. To evaluate age‐dependent changes, data obtained under similar conditions on 58 young adults (both genders, 17 to 39 yrs of age) were used. Rhythm characteristics were estimated by means of cosinor analyses, and intra‐ and inter‐individual variability by analysis of variance (ANOVA). In both age groups, the body temperature underwent daily changes. The MESOR (36.38±0.19°C vs. 36.17±0.21°C) and circadian amplitude (0.33±0.01°C vs. 0.26±0.01°C) were slightly decreased in the elderly compared to the young adult subjects (p<0.001). The mean circadian acrophase was similar in both age groups (17.19±1.66 vs. 16.93±3.08 h). However, the inter‐individual differences were higher in the older group, with individual values varying between 10:00 and 23:00 h. It was mainly this phase variability that caused a decrease in the inter‐daily rhythm stability and lower group amplitude. With melatonin treatment, the MESOR was lower by 0.1°C and the amplitude increased to 0.34±0.01°C, a similar value to that found in young adults. This was probably due to the increase of the inter‐daily rhythm stability. The mean acrophase did not change (16.93 vs. 16.75 h), although the inter‐individual variability decreased considerably. The corresponding standard deviations (SD) of the group acrophases were 3.08 and 1.51 h (p<0.01). A highly significant correlation between the acrophase before treatment and the phase change under melatonin treatment indicates that this is due to a synchronizing effect of melatonin. Apart from the difference in MESOR, the body temperature rhythm in the elderly subjects undergoing melatonin treatment was not significantly different from that of young adults. The data clearly show that age‐dependent changes mainly concern rhythm stability and synchronization with the 24 h day. A single daily melatonin dose stabilizes/synchronizes the body temperature rhythm, most probably via hypothermic and sleep‐improving effects.     

Die Temperaturabhängigkeit semilunarer und diurnaler Schlüpfrhythmen bei der intertidalen MückeClunio marinus (Diptera,Chironomidae)

On Helgoland (North Sea), the imagines ofClunio emerge during two seasonal periods (late spring and summer) from water temperatures of 8°–18 °C. The temperature dependence of the known semilunar eclosion rhythm ofClunio (correlated in nature with the spring tides every 14–15 days) was tested in the laboratory. Between 15° and 23 °C the semilunar eclosion maxima varied by only one day within the artifical 15-day zeitgebercycle, below 15 °C they were delayed up to 8 days at 8 °C. However, the days of pupation were approximately independent of the temperature level. One can conclude the existence of a temperature-independent physiological switch inducing the pupation only within a few days of the semilunar zeitgeber-cycle. Moreover, a semilunar synchronized differentiation of the imaginal discs already starts in the preceding larval instar, indicating an additional physiological switch. A model is suggested in which the semilunar eclosion rhythm and its relatively slight temperature dependence is explained by the action of two physiological switches which are coupled with the endogenous temperature-compensated lunar timing mechanism on the same days of the 15-day zeitgeber-cycle. In the laboratory, the diurnal eclosion and its underlying circadian timing mechanism (correlated on Helgoland with the time of spring low water in the late afternoon) also proved to be temperature independent between 12° and 20 °C. A comparison of field and laboratory data showed very similar results at temperatures around 18 °C (summer swarming period). In contrast, the midges emerged on all days of the semimonthly cycle of springs and neaps during the spring swarming period. This lack of semilunar synchronization may be the consequence of fluctuating temperatures during the larval and pupal development in spring time due to a general rise in the water temperature (4°–8 °C) and to short temperature rises up to 18 °C during exposure of the intertidal habitat at about low tide. Since some higher parts of theClunio habitat suitable for egg deposition are exposed on almost every day of the semimonthly cycle, even such animals that undergo lunar unsynchronized metamorphosis can reproduce within the short imaginal life duration (ca 2 h) if they emerge just about the time of low water. In correspondence with the daily delay in the times of low water by about 50 min, the diurnal eclosion rhythm was in fact modified with the tides during the spring period resulting in shifts of the diurnal eclosion time of up to 12 hours within the semimonthly cycle of springs and neaps.