Spatially varying selection shapes life history clines among populations of Drosophila melanogaster from sub‐Saharan Africa

Clines in life history traits, presumably driven by spatially varying selection, are widespread. Major latitudinal clines have been observed, for example, in Drosophila melanogaster, an ancestrally tropical insect from Africa that has colonized temperate habitats on multiple continents. Yet, how geographic factors other than latitude, such as altitude or longitude, affect life history in this species remains poorly understood. Moreover, most previous work has been performed on derived European, American and Australian populations, but whether life history also varies predictably with geography in the ancestral Afro‐tropical range has not been investigated systematically. Here, we have examined life history variation among populations of D. melanogaster from sub‐Saharan Africa. Viability and reproductive diapause did not vary with geography, but body size increased with altitude, latitude and longitude. Early fecundity covaried positively with altitude and latitude, whereas lifespan showed the opposite trend. Examination of genetic variance–covariance matrices revealed geographic differentiation also in trade‐off structure, and Q_ST‐F_ST analysis showed that life history differentiation among populations is likely shaped by selection. Together, our results suggest that geographic and/or climatic factors drive adaptive phenotypic differentiation among ancestral African populations and confirm the widely held notion that latitude and altitude represent parallel gradients.     

Environmental stress and reproduction in Drosophila melanogaster : starvation resistance, ovariole numbers and early age egg production

Background  

The Y model of resource allocation predicts a tradeoff between reproduction and survival. Environmental stress could affect a tradeoff between reproduction and survival, but the physiological mechanisms underlying environmental mediation of the tradeoff are largely unknown. One example is the tradeoff between starvation resistance and early fecundity. One goal of the present study was to determine if reduced early age fecundity was indeed a robust indirect response to selection for starvation resistance, by investigation of a set of D. melanogaster starvation selected lines which had not previously been characterized for age specific egg production. Another goal of the present study was to investigate a possible relationship between ovariole number and starvation resistance. Ovariole number is correlated with maximum daily fecundity in outbred D. melanogaster. Thus, one might expect that a negative genetic correlation between starvation resistance and early fecundity would be accompanied by a decrease in ovariole number.     

Impact of body melanization on mating success in Drosophila melanogaster

Mating speed and copulation duration respond rapidly to laboratory selection in Drosophila melanogaster Meigen (Diptera: Drosophilidae), but there is a lack of data on the evolutionary response to natural selection in the wild. Further, it is not clear whether body melanization and mating behavior are correlated traits. Accordingly, we tested whether variation in body color impacts on mating latency, copulation duration, and fecundity in latitudinal populations of D. melanogaster. We observed geographical variation (cline) for mating propensity, i.e., mating speed as well as copulation duration increased along latitude. Phenotypic plastic responses for body melanization at 17 and 25 °C also showed significant correlations with mating latency and copulation duration. Within‐population analysis based on assorted dark and light flies of five geographical populations showed significant positive correlations of copulation duration and fecundity with body melanization. To assess the role of males and/or females on mating speed and copulation duration, we used atypical body color strains (i.e., dark and light males of D. melanogaster) for no‐choice mating tests. Our data showed a major influence of males for copulation duration and of females for mating speed. Furthermore, a difference in impact of body melanization on mating speed and copulation duration was demonstrated between species, i.e., low melanization in Drosophila ananassae Doleschall is correlated with lower mating speed and shorter copulation duration than in D. melanogaster. Geographical changes in mating propensity were significantly correlated with body melanization at three levels, i.e., within and between populations and between species. Thus, we have shown that a relationship exists between body melanization and mating success. Further, we found seasonal changes in temperature and humidity to confer selection pressures on mating‐related traits.     

Impact of body melanisation on contrasting levels of desiccation resistance in a circumtropical and a generalist <Emphasis Type="Italic">Drosophila</Emphasis> species

We investigated the role of cuticular lipids, body melanisation and body size in conferring contrasting levels of desiccation resistance in latitudinal populations of Drosophila melanogaster and Drosophila ananassae on the Indian subcontinent. Contrary to the well known role of cuticular lipids in water proofing in diverse insect taxa, there is lack of geographical variations in the amount of cuticular lipids per fly in both the species. In D. ananassae, quite low levels of body melanisation are correlated with lower desiccation resistance. By contrast, increased levels of desiccation resistance are correlated with quite high melanisation in D. melanogaster. Thus, species specific cuticular melanisation patterns are significantly correlated with varying levels of desiccation resistance within as well as between populations and across species. Role of body melanisation in desiccation resistance is further supported by the fact that assorted dark and light flies differ significantly in cuticular water loss, hemolymph and dehydration tolerance. However, similar patterns of body size variation do not account for contrasting levels of desiccation resistance in these two Drosophila species. Climatic selection is evidenced by multiple regression analysis with seasonal amplitude of thermal and humidity changes (Tcv and RHcv) along latitude on the Indian subcontinent. Finally, the contrasting levels of species specific distribution patterns are negatively correlated with RHcv of sites of origin of populations i.e. a steeper negative slope for D. ananassae corresponds with its desiccation sensitivity as compared with D. melanogaster. Thus, evolutionary changes in body melanisation impact desiccation resistance potential as well as distribution patterns of these two Drosophila species on the Indian subcontinent.     

Coadapted changes in energy metabolites and body color phenotypes for resistance to starvation and desiccation in latitudinal populations of D. melanogaster

In D. melanogaster, resistance to starvation and desiccation vary in opposite directions across a geographical gradient in India but there is lack of such clinal variation on other continents. However, it is not clear whether these resistance traits or other correlated traits are the target of natural selection. For resistance to starvation or desiccation in D. melanogaster, we tested the hypothesis whether body color phenotypes and energy metabolites show correlated selection response. Our results are interesting in several respects. First, based on within population analysis, assorted darker and lighter flies from a given population showed that darker flies store higher amount of trehalose and confer greater desiccation resistance as compared with lighter flies. By contrast, lighter flies store higher lipids content and confer increased starvation tolerance. Thus, there is a trade-off for energy metabolites as well as body color phenotypes for starvation and desiccation stress. Further, trait associations within populations reflect similar patterns in geographical populations. Second, we found opposite clines for trehalose and body lipids. Third, coadapated phenotypes have evolved under contrasting climatic conditions i.e. drier and colder northern localities select darker flies with higher trehalose as well as desiccation resistance while hot and humid localities favor lighter flies with higher lipids level and greater starvation tolerance. Thus, the evolution of coadapated phenotypes associated with starvation and desiccation resistance might have resulted due to specific ecological conditions i.e. humidity changes on the Indian subcontinent.     

Adaptation to fluctuating environments in a selection experiment with Drosophila melanogaster

A fundamental question in life‐history evolution is how organisms cope with fluctuating environments, including variation between stressful and benign conditions. For short‐lived organisms, environments commonly vary between generations. Using a novel experimental design, we exposed wild‐derived Drosophila melanogaster to three different selection regimes: one where generations alternated between starvation and benign conditions, and starvation was always preceded by early exposure to cold; another where starvation and benign conditions alternated in the same way, but cold shock sometimes preceded starvation and sometimes benign conditions; and a third where conditions were always benign. Using six replicate populations per selection regime, we found that selected flies increased their starvation resistance, most strongly for the regime where cold and starvation were reliably combined, and this occurred without decreased fecundity or extended developmental time. The selected flies became stress resistant, displayed a pronounced increase in early life food intake and resource storage. In contrast to previous experiments selecting for increased starvation resistance in D. melanogaster, we did not find increased storage of lipids as the main response, but instead that, in particular for females, storage of carbohydrates was more pronounced. We argue that faster mobilization of carbohydrates is advantageous in fluctuating environments and conclude that the phenotype that evolved in our experiment corresponds to a compromise between the requirements of stressful and benign environments.     

Female fecundity traits in wild populations of African annual fish: the role of the aridity gradient

The evolution of life history is shaped by life expectancy. Life‐history traits coevolve, and optimal states for particular traits are constrained by trade‐offs with other life‐history traits. Life histories contrast among species, but may also diverge intraspecifically, at the level of populations. We studied the evolution of female reproductive allocation strategy, using natural populations of two sympatric species of African annual fishes, Nothobranchius furzeri and Nothobranchius orthonotus. These species inhabit pools in the Mozambican savanna that are formed in the rainy season and persist for only 2–10 months. Using 207 female N. furzeri from 11 populations and 243 female N. orthonotus from 14 populations, we tested the effects of genetic background (intraspecific lineage) and life expectancy (position on the aridity gradient determining maximum duration of their temporary habitat) on female fecundity traits. First, we found that variation in female body mass was small within populations, but varied considerably among populations. Second, we found that fecundity was largely defined by female body mass and that females spawned most of their eggs in the morning. Third, we found that the trade‐off between egg size and egg number varied among lineages of N. furzeri and this outcome has been confirmed by data from two separate years. Overall, we demonstrate that local conditions were important determinants for Nothobranchius growth and fecundity and that eggs size in arid region was less limited by female fecundity than in humid region.     

Higher investment in flight morphology does not trade off with fecundity estimates in a poleward range‐expanding damselfly

1. Evolutionary increases in dispersal‐related traits are frequently documented during range expansions. Investment in flight‐related traits is energetically costly and a trade‐off with fecundity may be expected during range expansion. 2. However, in contrast to wing‐dimorphic species, this trade‐off is not general in wing‐monomorphic species. In the absence of a dispersal‐‐fecundity trade‐off, an increased investment in clutch size at the expansion front is expected possibly at a cost of reduced offspring size. 3. The study evaluated investment in female flight morphology and fecundity‐related traits (clutch size, hatchling size) and potential trade‐offs among these traits in replicated populations of the poleward range‐expanding damselfly Coenagrion scitulum. 4. Females at the expansion front had a higher relative thorax length, indicating an increased investment in flight; this can be explained by spatial sorting of dispersal ability or in situ natural selection at the expansion front. Edge females produced larger hatchlings, however, this pattern was totally driven by the population‐specific thermal larval regimes and could not be attributed to the range expansion per se. By contrast, clutch sizes did not differ between core and edge populations. There was no signal of a dispersal–fecundity trade‐off either for a trade‐off between clutch size and hatchling size. 5. These results indicate that evolution of a higher dispersal ability at the expansion front of C. scitulum does not trade off with investment in fecundity, hence a dispersal–fecundity trade‐off is unlikely to slow down range expansion of this species.     

Evolutionary genetics of Drosophila ananassae: evidence for trade-offs among several fitness traits

Correlated responses to bi‐directional selection on thorax length, examined on several life‐history traits and chromosome inversion polymorphisms, have revealed apparent novel trade‐offs in Drosophila ananassae. We provide evidence of trade‐offs between hatching time and pupal period, pupal period and egg‐pupa development time, and pupal period and larval development time (LDT). Body size shows positive correlations with ovariole number, LDT and DT (egg–fly). We provide evidence of sexual dimorphism for trade‐offs between longevity and body size and starvation and longevity in females only. Trade‐offs between wing/thorax (W/T) ratio and longevity, W/T ratio and starvation, and DT (egg‐ fly) and longevity are evident in males only. Sexual dimorphism is also evident for inversion polymorphism with body size and longevity. A longevity assay suggests that low line females outlived high line females whereas high line males outlived low line males. The mean longevity in males is negatively correlated with the 2L‐ST and 3R‐ST arrangement frequencies whereas the 3L‐ST arrangement frequency is positively correlated with the mean longevity in males but opposite arrangements are found in females. Absolute starvation resistance is negatively correlated with 2L‐ST and 3R‐ST chromosome arrangements and results in a trade‐off between longevity and absolute starvation resistance in females. Analyses of fecundity, hatchability, and viabilities based on age intervals in both G₁₀ and G₁₃ suggest that the early reproduction is favoured in D. ananassae. The productivity percentage is highest in the high line and there is no effect of late reproduction on it. Overall, we provide some unravelled trade‐offs and striking sex differences, which may help in understanding the life‐history evolution of the species. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90 , 669–685.     

Trade-off of energy metabolites as well as body color phenotypes for starvation and desiccation resistance in montane populations of Drosophila melanogaster

Storage of energy metabolites has been investigated in different sets of laboratory selected desiccation or starvation resistant lines but few studies have examined such changes in wild-caught populations of Drosophila melanogaster. In contrast to parallel selection of desiccation and starvation tolerance under laboratory selection experiments, opposite clines were observed in wild populations of D. melanogaster. If resistance to desiccation and starvation occurs in opposite directions under field conditions, we may expect a trade-off for energy metabolites but such correlated changes are largely unknown. We tested whether there is a trade-off for storage as well as actual utilization of carbohydrates (trehalose and glycogen), lipids and proteins in D. melanogaster populations collected from different altitudes (512-2500 m). For desiccation resistance, darker flies (> 50% body melanization) store more body water content and endure greater loss of water (higher dehydration tolerance) as compared to lighter flies (< 30% body melanization). Based on within population analysis, we found evidence for coadapted phenotypes i.e. darker flies store and actually utilize more carbohydrates to confer greater desiccation resistance. In contrast, higher starvation resistance in lighter flies is associated with storage and actual utilization of greater lipid amount. However, darker and lighter flies did not vary in the rate of utilization of carbohydrates under desiccation stress; and of lipids under starvation stress. Thus, we did not find support for the hypothesis that a lower rate of utilization of energy metabolites may contribute to greater stress resistance. Further, for increased desiccation resistance of darker flies, about two-third of total energy budget is provided by carbohydrates. By contrast, lighter flies derive about 66% of total energy content from lipids which sustain higher starvation tolerance. Our results support evolutionary trade-off for storage as well as utilization of energy metabolites for desiccation versus starvation resistance in D. melanogaster.     

Variation in adult life history and stress resistance across five species of<Emphasis Type="Italic">Drosophila</Emphasis>

Dry weight at eclosion, adult lifespan, lifetime fecundity, lipid and carbohydrate content at eclosion, and starvation and desiccation resistance at eclosion were assayed on a long-term laboratory population ofDrosophila melanogaster, and one recently wild-caught population each of four other species ofDrosophila, two from themelanogaster and two from theimmigrans species group. The relationships among trait means across the five species did not conform to expectations based on correlations among these traits inferred from selection studies onD. melanogaster. In particular, the expected positive relationships between fecundity and size/lipid content, lipid content and starvation resistance, carbohydrate (glycogen) content and desiccation resistance, and the expected negative relationship between lifespan and fecundity were not observed. Most traits were strongly positively correlated between sexes across species, except for fractional lipid content and starvation resistance per microgram lipid. For most traits, there was evidence for significant sexual dimorphism but the degree of dimorphism did not vary across species except in the case of adult lifespan, starvation resistance per microgram lipid, and desiccation resistance per microgram carbohydrate. Overall,D. nasuta nasuta andD. sulfurigaster neonasuta (immigrans group) were heavier at eclosion than themelanogaster group species, and tended to have somewhat higher absolute lipid content and starvation resistance. Yet, these twoimmigrans group species were shorter-lived and had lower average daily fecundity than themelanogaster group species. The smallest species,D. malerkotliana (melanogaster group), had relatively high daily fecundity, intermediate lifespan and high fractional lipid content, especially in females.D. ananassae (melanogaster group) had the highest absolute and fractional carbohydrate content, but its desiccation resistance per microgram carbohydrate was the lowest among the five species. In terms of overall performance, the laboratory population ofD. melanogaster was clearly superior, under laboratory conditions, to the other four species if adult lifespan, lifetime fecundity, average daily fecundity, and absolute starvation and desiccation resistance are considered. This finding is contrary to several recent reports of substantially higher adult lifespan and stress resistance in recently wild-caught flies, relative to flies maintained for a long time in discretegeneration laboratory cultures. Possible explanations for these apparent anomalies are discussed in the context of the differing selection pressures likely to be experienced byDrosophila populations in laboratory versus wild environments. This paper is dedicated to the memory of our friend and former colleague Dr Hans Raj Negi, who tragically passed away at a very young age in a road accident in November 2003.     

Comparison of potential fecundity models for walleye pollock Gadus chalcogrammus in the Pacific waters off Hokkaido,Japan

Potential fecundity models of walleye or Alaska pollock Gadus chalcogrammus in the Pacific waters off Hokkaido, Japan, were developed. They were compared using a generalized linear model with using either standard body length (L_S) or total body mass (M_T) as a main covariate along with Fulton's condition factor (K) and mean diameter of oocytes (D_O) as additional potential covariates to account for maternal conditions and maturity stage. The results of model selection showed that M_T was a better single predictor of potential fecundity (F_P) than L_S. The biological importance of K on F_P was obscure, because it was statistically significant when used in the predictor with L_S (i.e. length‐based model), but not significant when used with M_T (i.e. mass‐based model). Meanwhile, D_O was statistically significant in both length and mass‐based models, suggesting the importance of downregulation on the number of oocytes with advancing maturation. Among all candidate models, the model with M_T and D_O in the predictor had the lowest Akaike's information criterion value, suggesting its better predictive power. These newly developed models will improve future comparisons of the potential fecundity within and among stocks by excluding potential biases other than body size.     

Evolution of starvation resistance in an invasive insect species,Aethina tumida (Coleoptera: Nitidulidae)

Starvation resistance, or the ability to survive periods without food, can shed light on selection pressure imposed by food scarcity, including chances to invade new regions as a result of human transport. Surprisingly, little information is known about starvation resistance for invasive insect species. Given that native and invasive populations differ in starvation resistance, this would suggest different selection scenarios and adaptive shifts fostering invasion success. Here, we show striking differences in starvation resistance of adult small hive beetles Aethina tumida (SHB) between native and invasive populations. In the laboratory, starvation resistance of freshly emerged laboratory‐reared and field‐collected adult females and males was evaluated in the beetle's native African range and in their invasive North American range. SHB in their native African range survived longer than SHB in their invasive North American range. Across ranges, females survived longer than males. Field‐collected SHB survived in Africa longer than freshly emerged ones, but not in the invasive range. This suggests no selection for starvation resistance in the invasive range, possibly due to differences between African and European‐derived honey bee hosts facilitating a trade‐off scenario between reproduction and starvation resistance. The ability of adult females to survive up to two months without food appears to be one factor contributing to the invasion success of this species. Assuming food availability is usually high in the invasive ranges, and trade‐offs between starvation resistance and fecundity/reproduction are common, it seems as if selection for starvation resistance during transport could set up potential trade‐offs that enhance reproduction after invasion. It would be interesting to see if this is a possible general pattern for invasive insect species.     

The role of the tolerance–fecundity trade‐off in maintaining intraspecific seed trait variation in a widespread dimorphic herb

Coexistence of species with different seed sizes is a long‐standing issue in community ecology, and a trade‐off between fecundity and stress tolerance has been proposed to explain co‐occurrence in heterogeneous environments. Here we tested an intraspecific extension of this model: whether such trade‐off also explains seed trait variation among populations of widespread plants under stress gradients. We collected seeds from 14 populations of Plantago coronopus along the Atlantic coast in North Africa and Europe. This herb presents seed dimorphism, producing large basal seeds with a mucilaginous coat that facilitates water absorption (more stress tolerant), and small apical seeds without coats (less stress tolerant). We analysed variation among populations in number, size and mucilage production of basal and apical seeds, and searched for relationships between local environment and plant size. Populations under higher stress (higher temperature, lower precipitation, lower soil organic matter) had fewer seeds per fruit, higher predominance of basal relative to apical seeds, and larger basal seeds with thicker mucilaginous coats. These results strongly suggest a trade‐off between tolerance and fecundity at the fruit level underpins variation in seed traits among P. coronopus populations. However, seed production per plant showed the opposite pattern to seed production per fruit, and seemed related to plant size and other life‐cycle components, as an additional strategy to cope with environmental variation across the range. The tolerance–fecundity model may constitute, under stress gradients, a broader ecological framework to explain trait variation than the classical seed size–number compromise, although several fecundity levels and traits should be considered to understand the diverse strategies of widespread plants to maximise fitness in each set of local conditions.     

No apparent cost of evolved immune response in Drosophila melanogaster

Maintenance and deployment of the immune system are costly and are hence predicted to trade‐off with other resource‐demanding traits, such as reproduction. We subjected this longstanding idea to test using laboratory experimental evolution approach. In the present study, replicate populations of Drosophila melanogaster were subjected to three selection regimes—I (Infection with Pseudomonas entomophila), S (Sham‐infection with MgSO₄), and U (Unhandled Control). After 30 generations of selection flies from the I regime had evolved better survivorship upon infection with P. entomophila compared to flies from U and S regimes. However, contrary to expectations and previous reports, we did not find any evidence of trade‐offs between immunity and other life history related traits, such as longevity, fecundity, egg hatchability, or development time. After 45 generations of selection, the selection was relaxed for a set of populations. Even after 15 generations, the postinfection survivorship of populations under relaxed selection regime did not decline. We speculate that either there is a negligible cost to the evolved immune response or that trade‐offs occur on traits such as reproductive behavior or other immune mechanisms that we have not investigated in this study. Our research suggests that at least under certain conditions, life‐history trade‐offs might play little role in maintaining variation in immunity.     

LONG-TERM LABORATORY EVOLUTION OF A GENETIC LIFE-HISTORY TRADE-OFF IN DROSOPHILA MELANOGASTER. 2. STABILITY OF GENETIC CORRELATIONS

Experiments in laboratory populations of Drosophila melanogaster have shown a negative genetic correlation between early-life fecundity on the one hand and starvation resistance and longevity on the other. Selection for late-life reproductive success resulted in long-lived populations that had increased starvation resistance but diminished early-life fecundity relative to short-lived controls. This pattern of differentiation proved, however, to be unstable. When assayed in a standard high-fecundity environment, the relative early fecundity of the long- and short-lived stocks reversed over a decade. That is, the long-lived populations came to have greater relative early-life fecundity, late-life fecundity, longevity and starvation resistance. Nevertheless, when these populations were assayed in other assay environments, the original trade-off was still present. We investigated the genetic structure of the short- and long-lived populations, to ask whether the inconstancy of the trade-off, as inferred from among population comparisons, is reflected in the pattern of genetic correlations within populations. For this purpose, lines from each of the short- and long-lived populations that had been selected for starvation resistance were compared with unselected controls. The direct and correlated responses of these starvation selected populations suggest that (1) the original genetic trade-off was still present in the ancestral short- and long-lived populations, even when it was no longer apparent from their comparison; (2) the trade-off was present in both assay environments; and (3) selectable genotype × environment variation exists for early fecundity. We suggest that a failure of the pattern of differentiation among populations to reflect the pattern of genetic correlations, if common in natural populations, will prevent the reliable inference of genetic trade-offs from comparisons of most natural populations.     

Dietary polyunsaturated fatty acids affect volume and metabolism of Drosophila melanogaster sperm

Dietary fatty acids can accumulate in sperm and affect their function in vertebrates. As Drosophila melanogaster shares several pathways of lipid metabolism and shows similar lipid‐dependent phenotypes but lacks some hormones that in vertebrates regulate lipid metabolism, there is currently no clear prediction as to how dietary fatty acids affect the sperm of D. melanogaster. We manipulated the amount and identity of dietary polyunsaturated fatty acids (PUFA) in the food of D. melanogaster males (a treatment known to affect membrane fluidity) and measured changes in sperm parameters. We found that (a) males reared on food containing PUFA‐rich, plant‐derived lipids showed a slower increase in sperm volume over male age compared to males reared on yeast‐derived lipid food which is richer in saturated fatty acids. (b) The resistance of sperm membrane integrity to osmotic stress was not altered by dietary lipid treatment, but (c) food containing yeast‐derived lipids induced a 46% higher in situ rate of production of reactive oxygen species in sperm cells. These findings show that dietary lipids have similar effects on sperm parameters in Drosophila as in vertebrates, affect some, but not all, sperm parameters and modulate male reproductive ageing. In concert with recent findings of sex‐specific seasonal variation of diet choice in the wild, our results suggest a substantial dietary impact on the dynamics of male reproduction in the wild.     

Impact of body melanisation on desiccation resistance in montane populations of D. melanogaster: Analysis of seasonal variation

In the montane localities of subtropical regions, winter is the dry season and ectothermic drosophilids are expected to evolve desiccation resistance to cope with drier climatic conditions. An analysis of six montane populations (600–2226 m) of D. melanogaster showed variations for body melanisation (i.e. pigmentation) and desiccation resistance across seasons as well as along altitude. During winter season, plastic changes for melanisation of three posterior abdominal segments (5th + 6th + 7th) correspond with higher desiccation resistance. Thus, we analyzed genetic and plastic effects for these ecophysiological traits by comparing wild-caught and laboratory reared individuals of D. melanogaster for autumn as well as winter season. A ratio of slope values in wild vs. laboratory populations has shown a 1.64-fold plastic effect during autumn; and a two-fold effect during winter. For body melanisation and desiccation resistance, evolutionary response to altitudinal environmental gradient is similar to the phenotypic response across seasons. Thus, our observations are in agreement with the co-gradient hypothesis. Further, we tested the hypothesis whether a thicker cuticle (either due to melanisation or cuticular lipids) leads to lesser cuticular water loss and higher desiccation resistance across seasons as well as according to altitude. Based on within and between population analyses, body melanisation was found to be positively correlated with desiccation resistance but negatively with cuticular water loss. Interestingly, there were no changes in the amount of cuticular lipids per fly across seasons as well as along altitude; and therefore cuticular lipids did not account for desiccation resistance. Cuticular water loss exhibited negative correlation with body melanisation but not with cuticular lipids as well as with changes in body size across seasons. Thus, our data suggest that seasonal changes in body melanisation confer desiccation resistance in montane populations of D. melanogaster.     

The involvement of lipids in light-induced charge separation in the reaction center of photosystem II

Extraction of PS II particles with 50 mM cholate and 1 M NaCl releases several proteins (33-, 23-, 17- and 13 kDa) and lipids from the thylakoid membrane which are essential for O₂ evolution, dichlorophenolindophenol (DCIP) reduction and for stable charge separation between P680⁺ and Q_A ^-. This work correlates the results on the loss of steady-state rates for O₂ evolution and PS II mediated DCIP photo-reduction with flash absorption changes directly monitoring the reaction center charge separation at 830 nm due to P680⁺, the chlorophyll a donor. Reconstitution of the extracted lipids to the depleted membrane restores the ability to photo-oxidize P680 reversibly and to reduce DCIP, while stimulating O₂ evolution minimally. Addition of the extracted proteins of masses 33-, 23- and 17- kDa produces no further stimulation of DCIP reduction in the presence of an exogenous donor like DPC, but does enhance this rate in the absence of exogenous donors while also stimulating O₂ evolution. The proteins alone in the absence of lipids have little influence on charge separation in the reaction center. Thus lipids are essential for stable charge separation within the reaction center, involving formation of P680⁺ and Q_A ^-.Abbreviations A₈₃₀ Absorption change at 830 nm - Chl Chlorophyll - D₁ primary electron donor to P680 - DCIP 2,6-dichlorophenolindophenol - DPC 1,5-diphenylcarbazide - MOPS 3-(N-morpholino)propanesulfonic acid - P680 reaction center chlorophyll a molecule of photosystem II - PPBQ Phenyl-p-benzoquinone - PS II Photosystem II - Q_A, Q_B first and second quinone acceptors in PS II - V-DCIP rate of DCIP reduction - V-O₂ rate of oxygen evolution - Y water-oxidizing enzyme system - CHAPS 3-Cyclohexylamino-propanesulfonic acid     

An inversion supergene in Drosophila underpins latitudinal clines in survival traits

Chromosomal inversions often contribute to local adaptation across latitudinal clines, but the underlying selective mechanisms remain poorly understood. We and others have previously shown that a clinal inversion polymorphism in Drosophila melanogaster, In(3R)Payne, underpins body size clines along the North American and Australian east coasts. Here, we ask whether this polymorphism also contributes to clinal variation in other fitness‐related traits, namely survival traits (lifespan, survival upon starvation and survival upon cold shock). We generated homokaryon lines, either carrying the inverted or standard chromosomal arrangement, isolated from populations approximating the endpoints of the North American cline (Florida, Maine) and phenotyped the flies at two growth temperatures (18 °C, 25 °C). Across both temperatures, high‐latitude flies from Maine lived longer and were more stress resistant than low‐latitude flies from Florida, as previously observed. Interestingly, we find that this latitudinal pattern is partly explained by the clinal distribution of the In(3R)P polymorphism, which is at ~ 50% frequency in Florida but absent in Maine: inverted karyotypes tended to be shorter‐lived and less stress resistant than uninverted karyotypes. We also detected an interaction between karyotype and temperature on survival traits. As In(3R)P influences body size and multiple survival traits, it can be viewed as a ‘supergene’, a cluster of tightly linked loci affecting multiple complex phenotypes. We conjecture that the inversion cline is maintained by fitness trade‐offs and balancing selection across geography; elucidating the mechanisms whereby this inversion affects alternative, locally adapted phenotypes across the cline is an important task for future work.