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.
BackgroundWe recently reported that the altitude of origin altered the photic and thermal sensitivity of the circadian pacemaker controlling eclosion and oviposition rhythms of high altitude Himalayan strains of Drosophila ananassae. The present study was aimed at investigating the effects of altitude of origin on the pacemaker controlling the adult locomotor activity rhythm of D. helvetica.MethodsLocomotor activity rhythms of the high altitude Himalayan (haH) strain (Hemkund-Sahib, 4,121 m above sea level) and the low altitude Himalayan (laH) strain (Birahi, 1,132 m a.s.l.) of D. helvetica were assayed by two experiments. The first experiment examined the natural entrainment pattern in light-dark (LD) cycles at the breeding site of each strain. The second experiment examined the entrainment parameters in LD 12:12 cycles and the period of free-running rhythm in constant darkness (DD) under controlled laboratory conditions.ResultsWhen entrained by natural or artificial LD cycles, the haH strain had an unimodal activity pattern with a single peak that commenced in the forenoon and continued till evening, while the laH strain had a bimodal activity pattern in which the morning peak occurred before lights-on and was separated by about 4 h from the evening peak. Unimodality of the haH strain was retained in DD; however, bimodality of the laH strain was abolished in DD since the evening peak disappeared immediately after the trasfer from LD 12:12 to DD. The period of the free-running rhythm of the haH strain was ~26.1 h, whereas that of the laH strain was ~21.7 h.ConclusionParameters of entrainment and free-running rhythm of the adult locomotor activity of the haH strain of D. helvetica were strikingly different from those of the laH strain and were likely due to ecological adaptations to the prevailing environmental conditions at the altitude where the species evolved.
The sensitivity of the circadian photoreceptors mediating entrainment of the eclosion rhythm and phase shifts of oviposition rhythm of the high altitude (HA) strain of Drosophila ananassae originating from Badrinath (5123 m above sea level) in the Himalayas was compared with the low altitude (LA) strain from Firozpur (179 m above sea level). Reduced photic sensitivity of the HA strain is regarded as the result of natural selection, which led to the weakening of the coupling mechanism between the circadian pacemaker and light at the high altitude of origin. The present study was designed to determine whether or not the photic entrainment of the oviposition rhythm of the HA strain of D. ananassae is also altered by the high altitude of its origin, and the results are compared with those of the LA strain. The effects of light intensity on the phase angle difference (Psi), degree of rhythmicity (R), the percent oviposition in photophase, the threshold light intensity (i.e., the intensity at which stable entrainment occurred), and the saturation light intensity (i.e., the intensity beyond which the values of Psi or amplitude of rhythm remained unaltered) were determined. Entrainment was studied in light-dark cycles in which the light intensity of 12 h of photophase varied from 1 to 1000 lux, and complete darkness prevailed in all scotophases. The oviposition rhythm of the HA strain was arrhythmic from 1 to 90 lux, weakly rhythmic at 95 lux, but rhythmic at or above 100 lux, while that of the LA strain was weakly rhythmic at 1 lux but rhythmic at or above 2 lux. Oviposition of the HA strain occurred mostly in the photophase, while that of the LA strain occurred in the scotophase; as a result, the oviposition medians of the HA strain were around the subjective forenoons while those of the LA strain were around the subjective evenings. The percent of oviposition in photophase increased from 68 to 98 in the HA strain and from 5 to 33 in the LA strain as light intensity increased from 1 to 1000 lux. In the HA strain, the Psi values were significantly less and values of R and percent oviposition in photophase were significantly more than those of the LA strain at each level of light intensity. Threshold and saturation intensities for Psi were 100 and 700 lux, respectively, for the HA strain, but just 2 and 45 lux, respectively, for the LA strain. The saturation intensity for R was 650 and 700 lux for the HA and LA strains, respectively. These results extend the confirmation that the reduced photic sensitivity of the HA strain might have been acquired through natural selection in response to environmental conditions at the high altitude of its origin.
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