The circadian pacemaker controlling the eclosion rhythm of the high altitude Himalayan strains of Drosophila ananassae captured at Badrinath (5123 m) required ambient temperature at 21 degrees C for the entrainment and free-running processes. At this temperature, their eclosion rhythms entrained to 12h light, 12h dark (LD 12:12) cycles and free-ran when transferred from constant light (LL) to constant darkness (DD) or upon transfer to constant temperature at 21 degrees C following entrainment to temperature cycles in DD. These strains, however, were arrhythmic at 13 or 17 degrees C under identical experimental conditions. Eclosion medians always occurred in the thermophase of temperature cycles whether they were imposed in LL or DD; or whether the thermophase coincided with the photophase or scotophase of the concurrent LD 12:12 cycles. The temperature dependent rhythmicity in the Himalayan strains of D. ananassae is a rare phenotypic plasticity that might have been acquired through natural selection by accentuating the coupling sensing mechanism of the pacemaker to temperature, while simultaneously suppressing the effects of light on the pacemaker.
Eclosion rhythm of the high-altitude Himalayan strain of Drosophila ananassae from Badrinath (altitude 5123 m) was temperature-dependent and at 21 degrees C, it was entrained by cycles of 12h light: 12h darkness (LD 12:12) and free-ran in constant darkness, however, it was arrhythmic at 13 degrees C or 17 degrees C under identical experimental conditions (Khare, P. V., Barnabas, R. J., Kanojiya, M., Kulkarni, A. D., Joshi, D. S. (2002). 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 degrees C, 17 degrees C, and 21 degrees C by two types of LD cycles in which the photoperiod at 100 lux intensity varied from 6h to 18h, 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 degrees C but not at 13 degrees C or 17 degrees 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.
The effect of altitude on four basic properties of the pacemaker controlling the circadian rhythm of oviposition in two strains of Drosophila ananassae was determined. The high altitude (HA) strain from Badrinath (5123 m above sea level) had a low amplitude peak in the forenoon while the low altitude (LA) strain from Firozpur (179 m a.s.l.) had a high amplitude peak after the lights-off of LD 12:12 cycles. Free running periods in continuous darkness were about 22.6 and 27.4 h in the HA and LA strains, respectively. The light pulse phase response curve (PRC) for the HA strain showed a low amplitude and a dead zone of 8h; the ratio for the advance to delay region (A/D) was less than 1, while the PRC for the LA strain had a high amplitude, which was devoid of a dead zone and showed a ratio of A/D > 1. The magnitude of the delay phase shifts at CT 18 evoked by light pulses of 1 h duration, but varying light intensity was significantly different in the HA and LA strain, which suggests that the photic sensitivity of the clock photoreceptors mediating the phase shifts had been affected by the altitude.
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(o)N, 80.2(o)E). 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(o)C 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 approximately 2 h prior to sunrise, and the 24 h rhythmicity was the most robust. Effects of artificial LAN at 25(o) +/- 0.5(o)C 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 approximately 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 approximately 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 (tau) 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 tau and nature of the eclosion rhythm in all four experiments. Pupae raised in 0.5 lux LAN exhibited the shortest tau (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 tau (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.
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