Drosophila simulans is more abundant under colder and drier montane habitats in the western Himalayas as compared to its sibling D. melanogaster but the mechanistic bases of such climatic adaptations are largely unknown. Previous studies have described D. simulans as a desiccation sensitive species which is inconsistent with its occurrence in temperate regions. We tested the hypothesis whether developmental plasticity of cuticular traits confers adaptive changes in water balance-related traits in the sibling species D. simulans and D. melanogaster. Our results are interesting in several respects. First, D. simulans grown at 15 °C possesses a high level of desiccation resistance in larvae (~39 h) and in adults (~86 h) whereas the corresponding values are quite low at 25 °C (larvae ~7 h; adults ~13 h). Interestingly, cuticular lipid mass was threefold higher in D. simulans grown at 15 °C as compared with 25 °C while there was no change in cuticular lipid mass in D. melanogaster. Second, developmental plasticity of body melanisation was evident in both species. Drosophila simulans showed higher melanisation at 15 °C as compared with D. melanogaster while the reverse trend was observed at 25 °C. Third, changes in water balance-related traits (bulk water, hemolymph and dehydration tolerance) showed superiority of D. simulans at 15 °C but of D. melanogaster at 25 °C growth temperature. Rate of carbohydrate utilization under desiccation stress did not differ at 15 °C in both the species. Fourth, effects of developmental plasticity on cuticular traits correspond with changes in the cuticular water loss i.e. water loss rates were higher at 25 °C as compared with 15 °C. Thus, D. simulans grown under cooler temperature was more desiccation tolerant than D. melanogaster. Finally, desiccation acclimation capacity of larvae and adults is higher for D. simulans reared at 15 °C but quite low at 25 °C. Thus, D. simulans and D. melanogaster have evolved different strategies of water conservation consistent with their adaptations to dry and wet habitats in the western Himalayas. Our results suggest that D. simulans from lowland localities seems vulnerable due to limited acclimation potential in the context of global climatic change in the western Himalayas. Finally, this is the first report on higher desiccation resistance of D. simulans due to developmental plasticity of both the cuticular traits (body melanisation and epicuticular lipid mass) when grown at 15 °C, which is consistent with its abundance in temperate regions.
Previous studies on two tropical Drosophila species (D. malerkotliana and D. bipectinata) have shown lower resistance to stress-related traits but the rapid colonization of D. malerkotliana in the past few decades is not consistent with its sensitivity to desiccation and cold stress. We tested the hypothesis that developmental acclimation at two growth temperatures (17 and 25°C) can confer adaptations to desiccation and thermal stresses. We found divergence in developmental plastic effects on cuticular traits, i.e. a significant increase of body melanisation (~2-fold) and of cuticular lipid mass (~3-fold) in D. malerkotliana but only 1.5-fold higher cuticular lipid mass in D. bipectinata when grown at 17°C compared with 25°C. A comparison of the water budget of these two species showed significantly higher effects of developmental acclimation on body water content, rate of water loss and dehydration tolerance resulting in higher desiccation resistance in D. malerkotliana than in D. bipectinata. When grown in cooler conditions (17°C), D. malerkotliana had greater resistance to cold as well as desiccation stress. In contrast, heat resistance of D. bipectinata was higher when grown at 25°C. These laboratory observations are supported by data on seasonally varying populations. Furthermore, adult D. malerkotliana acclimated to different stresses showed greater resistance to those stresses than D. bipectinata adults. Thus, significant increase in stress resistance of D. malerkotliana through developmental acclimation may be responsible for its invasion and ecological success on different continents compared with D. bipectinata.
Water balance mechanisms have been investigated in desert Drosophila species of the subgenus Drosophila from North America, but changes in mesic species of subgenus Drosophila from other continents have received lesser attention. We found divergent strategies for coping with desiccation stress in two species of immigrans group--D. immigrans and D. nasuta. In contrast to clinal variation for body melanization in D. immigrans, cuticular lipid mass showed a positive cline in D. nasuta across a latitudinal transect (10°46'-31°43'N). Based on isofemale lines variability, body melanization showed positive correlation with desiccation resistance in D. immigrans but not in D. nasuta. The use of organic solvents has supported water proofing role of cuticular lipids in D. nasuta but not in D. immigrans. A comparative analysis of water budget of these two species showed that higher water content, reduced rate of water loss and greater dehydration tolerance confer higher desiccation resistance in D. immigrans while the reduced rate of water loss is the only possible mechanism to enhance desiccation tolerance in D. nasuta. We found that carbohydrates act as metabolic fuel during desiccation stress in both the species, whereas their rates of utilization differ significantly between these two species. Further, acclimation to dehydration stress improved desiccation resistance due to increase in the level of carbohydrates in D. immigrans but not in D. nasuta. Thus, populations of D. immigrans and D. nasuta have evolved different water balance mechanisms under shared environmental conditions. Multiple measures of desiccation resistance in D. immigrans but reduction in water loss in D. nasuta are consistent with their different levels of adaptive responses to wet and dry conditions on the Indian subcontinent.
In the Western Himalayas, Drosophila nepalensis is more abundant during the colder and drier winter than the warmer rainy season but the mechanistic bases of such adaptations are largely unknown. We tested effects of developmental plasticity on desiccation-related traits (body size, body melanization and water balance traits) that may be consistent with changes in seasonal abundance of this species. D. nepalensis grown at 15°C has shown twofold higher body size, greater melanization (∼15-fold), higher desiccation resistance (∼55 h), hemolymph as well as carbohydrate content (twofold higher) as compared with corresponding values at 25°C. Water loss before succumbing to death was much higher (∼16%) at 15°C than 25°C. Developmental plastic effects on body size are associated with changes in water balance-related traits (bulk water, hemolymph and dehydration tolerance). The role of body melanization was evident from the analysis of assorted darker and lighter flies (from a mass culture of D. nepalensis reared at 21°C) which lacked differences in dry mass but showed differences in desiccation survival hours and rate of water loss. For adult acclimation, we found a slight increase in desiccation resistance of flies reared at lower growth temperature, whereas in flies reared at 25°C such a response was lacking. In D. nepalensis, greater developmental plasticity is consistent with its contrasting levels of seasonal abundance. Finally, in the context of global climate change in the Western Himalayas, D. nepalensis seems vulnerable in the warmer season due to lower adult as well as developmental acclimation potential at higher growth temperature (25°C).
Laboratory selection experiments have evidenced storage of energy metabolites in adult Xies of desiccation and starvation resistant strains of D. melanogaster but resource acquisition during larval stages has received lesser attention. For wild populations of D. melanogaster, it is not clear whether larvae acquire similar or diVerent energy metabolites for desiccation and starvation resistance. We tested the hypothesis whether larval acquisition of energy metabolites is consistent with divergence of desiccation and starvation resistance in darker and lighter isofemale lines of D. melanogaster. Our results are interesting in several respects. First, we found contrasting patterns of larval resource acquisition, i.e., accumulation of higher carbohydrates during 3rd instar larval stage of darker Xies versus higher levels of triglycerides in 1st and 2nd larval instars of lighter Xies. Second, 3rd instar larvae of darker Xies showed »40 h longer duration of development at 21°C; and greater accumulation of carbohydrates (trehalose and glycogen) in fed larvae as compared with larvae nonfed after 150 h of egg laying. Third, darker isofemale lines have shown signiWcant increase in total water content (18%); hemolymph (86%) and dehydration tolerance (11%) as compared to lighter isofemale lines. Loss of hemolymph water under desiccation stress until death was signiWcantly higher in darker as compared to lighter isofemale lines but tissue water loss was similar. Fourth, for larvae of darker Xies, about 65% energy content is contributed by carbohydrates for conferring greater desiccation resistance while the larvae of lighter Xies acquire 2/3 energy from lipids for sustaining starvation resistance; and such energy diVerences persist in the newly eclosed Xies. Thus, larval stages of wild-caught darker and lighter Xies have evolved independent physiological processes for the accumulation of energy metabolites to cope with desiccation or starvation stress.
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