Evolutionary Consequences of Desiccation Resistance in the Male Ejaculate

Avoiding water loss for insects is critical for survival. Selection for reduced water loss will depend on trade-offs between resources allocated for reproduction and those allocated for resisting desiccation. However, we lack knowledge on how selection for desiccation resistance can affect the male ejaculate. Furthermore, as male ejaculate composition is complex, desiccation resistant females could evolve traits that enable them to derive longevity benefits from mating. Here, we assessed how selection for desiccation resistance impacts male testes and accessory gland size, protein content of these organs, female sperm storage and male ability to inhibit female remating behavior, in the Mexican fruit fly Anastrepha ludens. Additionally, we tested if mating increased longevity and fecundity in desiccation resistant females. Males selected for resistance to desiccation stress had smaller accessory glands and seminal vesicles and females mating with these males stored less sperm compared to control males. Females mating with resistant males had lower fecundity compared to females mating with control males. Desiccation resistant females lived longer than control females, yet this was irrespective of mating. Rapid evolutionary responses to hydric stress can have correlated effects in reproductive capabilities, which are not restricted to pre-copulatory traits. Trade-offs between resistance to desiccation stress are reflected in decreased allocation of resources to reproductive organs. Thus, production of the ejaculate may be costly for A. ludens males. Knowledge on the evolution of ejaculate traits and reproductive organ size in response to directional selection for desiccation resistance, will aid our understanding of differential sex-specific responses to environmental stress.     

Temperature and photoperiod affect stress resistance traits in Drosophila melanogaster

The long‐term survival of species and populations depends on their ability to adjust phenotypic values to environmental conditions. In particular, the capability of dealing with environmental stress to buffer detrimental effects on fitness is considered to be of pivotal importance. Resistance traits are readily modulated by a wide range of environmental factors. In the present study, Drosophila melanogaster Meigen is used to investigate plastic responses to temperature and photoperiod in stress resistance traits. The results reveal that stress resistance traits (cold, heat, starvation and desiccation resistance) are affected by the factors temperature and sex predominantly. Cooler temperatures compared with warmer temperatures increase cold tolerance, desiccation and starvation resistance, whereas they reduce heat tolerance. Except for heat resistance, females are more stress‐resistant than males. Stress resistance traits are also affected by photoperiod. Shorter photoperiods decrease cold tolerance, whereas longer photoperiods enhance desiccation resistance. Overall, thermal effects are pervasive throughout all measured resistance traits, whereas photoperiodic effects are of limited importance in the directly developing (i.e. nondiapausing) flies used here, suggesting that pronounced photoperiodic effects on stress resistance traits may be largely limited to, and triggered by, diapause‐inducing effects.     

Size specific demography of three species of Anastrepha fruit flies

Individuals of three Anastrepha species: A. obliqua, A. ludens, and A. serpentina (Diptera: Tephritidae), were sorted according to pupal weight in cohorts of large and small flies. Demographic parameters and reproductive patterns and heterogeneity were determined for each cohort. Large flies of the three species presented greater expectation of life and gross fecundity rates. A. ludens was the species with the longest life span (expectation of life of large adults was 110 days) and the greatest gross fecundity rates (1597 eggs/female for the large flies). While, A. obliqua had the shortest mean age of reproduction (33 days), and the greatest daily egg production (14 eggs/female/day). Net fecundity was similar in these two. A. serpentina had lower fecundity rates.Reproductive information for each size and each species include: age-by-parity relations, fraction of sexually mature life in which females lay eggs, and frequency distribution of individual egg production. Results demonstrate that even under constant laboratory conditions and using standard artificial hosts, there is a great deal of life history variation among these Anastrepha species and among other tephritid fruit flies.     

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.     

Reproductive and post‐reproductive life history of wild‐caught Drosophila melanogaster under laboratory conditions

The life history of the fruit fly (Drosophila melanogaster) is well understood, but fitness components are rarely measured by following single individuals over their lifetime, thereby limiting insights into lifetime reproductive success, reproductive senescence and post‐reproductive lifespan. Moreover, most studies have examined long‐established laboratory strains rather than freshly caught individuals and may thus be confounded by adaptation to laboratory culture, inbreeding or mutation accumulation. Here, we have followed the life histories of individual females from three recently caught, non‐laboratory‐adapted wild populations of D. melanogaster. Populations varied in a number of life‐history traits, including ovariole number, fecundity, hatchability and lifespan. To describe individual patterns of age‐specific fecundity, we developed a new model that allowed us to distinguish four phases during a female's life: a phase of reproductive maturation, followed by a period of linear and then exponential decline in fecundity and, finally, a post‐ovipository period. Individual females exhibited clear‐cut fecundity peaks, which contrasts with previous analyses, and post‐peak levels of fecundity declined independently of how long females lived. Notably, females had a pronounced post‐reproductive lifespan, which on average made up 40% of total lifespan. Post‐reproductive lifespan did not differ among populations and was not correlated with reproductive fitness components, supporting the hypothesis that this period is a highly variable, random ‘add‐on’ at the end of reproductive life rather than a correlate of selection on reproductive fitness. Most life‐history traits were positively correlated, a pattern that might be due to genotype by environment interactions when wild flies are brought into a novel laboratory environment but that is unlikely explained by inbreeding or positive mutational covariance caused by mutation accumulation.     

Evolution of pre- and post-copulatory traits in female Drosophila melanogaster as a correlated response to selection for resistance to cold stress

Exposure to low temperatures reduces gamete viability and fecundity in females of insect species like Drosophila. Hence, adaptation to cold stress can in principle involve modifications in reproductive traits in females. Studies on resistance to cold stress have mostly addressed the evolution of adult survivorship post cold shock. Very few studies have addressed the evolution of reproductive traits in females in response to cold stress. We have successfully selected replicate populations of Drosophila melanogaster for resistance to cold shock. After 50 generations of selection, we investigated pre- and post-copulatory traits i.e. mating latency, copulation duration, mating frequency and progeny production in female flies exposed to cold shock or control conditions. Post cold shock, females from the selected populations were better at recovery in terms of mating latency, mating success, and progeny production relative to females from the control populations. Performance of the two types of females was not different under control conditions. These findings clearly indicate that adaptation to cold stress involves rapid modification of the reproductive traits.     

Demography of a genetic sexing strain of Anastrepha ludens (Diptera: Tephritidae): effects of selection based on mating performance

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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.     

Fecundity in Drosophila following desiccation is dependent on nutrition and selection regime

Life-history trade-offs of populations of Drosophila melanogaster selected for enhanced desiccation resistance (D populations) include increased pupal mortality, retarded development, increased longevity, and decreased fecundity. We tested the effects of acute desiccation on fecundity and the effects of various nutritional regimes on fecundity following a bout of desiccation in the D populations and their control populations (C populations). Female flies were desiccated for 8 h (C populations) or 24 h (D populations) and then recovered on one of four treatments: distilled water, a saline solution, a saline and sucrose solution, or food. Each treatment was tested in the presence and absence of a yeast supplement. Fecundity was then measured on an oviposition medium over a 12-h period. Fecundity was also measured in flies that did not undergo a desiccation stress. Yeast increased fecundity under all conditions examined in the C populations but not in the D populations. There was an overall effect of recovery treatment on fecundity but no effect of desiccation stress.     

Selection for adult desiccation resistance in Drosophila melanogaster: fitness components, larval resistance and stress correlations

In previous experiments we found that Drosophila melanogaster lines selected for increased adult desiccation resistance had increased resistance to other environmental stresses at the adult stage including starvation, intense ⁶⁰Co-γ radiation and a toxic ethanol level. In further studies on these lines, we now show that selection did not alter resistance to desiccation and ethanol at the larval stage. As well as having a lower early fecundity, selected lines showed increased adult male longevity and increased viability at high larval densities compared with control lines. There were no changes in development time or mating success. The increased male longevity is consistent with the reduced metabolic rate of the selected lines.
A genetic correlation between resistance to different stresses was confirmed by an analysis of isofemale lines derived from a population founded by flies from a stress-resistant line and an unselected line. The results are consistent with the existence of genes segregating in natural populations conferring increased general stress resistance.     

Resistance to environmental stress in Drosophila ananassae: latitudinal variation and adaptation among populations

Geographical variation in traits related to fitness is often the result of adaptive evolution. Stress resistance traits in Drosophila often show clinal variation, suggesting that selection affects resistance traits either directly or indirectly. Multiple stress resistance traits were investigated in 45 natural populations of Drosophila ananassae collected from all over India. There was significant positive correlation between starvation resistance and lipid content. Significant negative correlations between desiccation and lipid content and between desiccation and heat resistance were also found. Flies from lower latitudes had higher starvation resistance, heat resistance and lipid content but the pattern was reversed for desiccation resistance. These results suggest that flies from different localities varied in their susceptibility to starvation because of difference in their propensity to store body lipid. Multiple regression analysis provided evidence of climatic selection driven by latitudinal variation in the seasonal amplitude of temperature and humidity changes within the Indian. Finally, our results suggest a high degree of variation in stress resistance at the population level in D. ananassae.     

Comparative response to post-production process of two Anastrepha ludens strains: Application in the sterile insect technique

The new desiccation-resistant (DR) strain of Anastrepha ludens Loew differs in its life-history traits from the non-selected strain (NS). Given the innate resistance of DR flies to stressors, it is necessary to determine the packing and shipment conditions for their use in the sterile insect technique (SIT). First, we used pupae to evaluate the interaction of hypoxia period (12, 24 and 36 hr) and temperature (15.3 (±0.9), 19.5 (±1.0), 25.5 (±0.6) and 30.1 (±0.5) ^oC). Second, adults were exposed to densities of 1.0, 1.3 and 1.5 flies/cm² and chilling periods of 0, 3 and 5 hr. We observed a negative relationship of emergence and flight ability with hypoxia period and temperature in both strains especially when flies were exposed to 36 hr of hypoxia and 30℃. Interestingly, knock-down time decreased as male density increased, but when chilling time increased, recovery time also increased. Both chilling time and male density had negative effects on male weight. Chilling time reduced male survival when exposed to desiccation, while both chilling time and male density decreased male survival under starvation conditions. Negative effects on number of matings were observed only on the first day after chilling but not on the following days. We found that DR flies survive approximately twice as long as flies from the NS strain when exposed to starvation or desiccation stressors. We discuss our results on the basis of the potential use of DR flies in SIT programmes.     

Desiccation resistance along an aridity gradient in the cactophilic fly Drosophila buzzatii: sex-specific responses to stress

Stress resistance characters are valuable tools for the study of acclimation potential, adaptive strategies and biogeographic patterns in species exposed to environmental variability. Water stress is a challenge to terrestrial arthropods because of their small size and relatively high area: volume ratio. Fruit flies have been investigated to record adaptive morphological and physiological traits, as well as to test their responses to stressful factors. In this study, we investigate the ability to cope with water stress, by examining variation in desiccation resistance in a species that lives mainly in desert lands. Specifically, we explored the genetic and ecological basis of desiccation resistance in populations of Drosophila buzzatii from Northern Argentina. We used a common garden experiment with desiccation treatments on a number of isofemale lines from four populations along an aridity gradient. Our results revealed significant among-population differentiation and substantial amounts of genetic variation for desiccation resistance. We also detected significant genotype-by-environment and genotype-by-sex interactions indicative that desiccation resistance responses of the lines assayed were environment- and sex-specific. In addition, we observed clinal variation in female desiccation resistance along gradients of altitude, temperature and humidity; that desiccation resistance is a sexually dimorphic trait, and that sexual dimorphism increased along the aridity and altitudinal gradients. Based on current evidence, we propose that the observed sex-specific responses may reflect different life history traits, and survival and reproductive strategies in different ecological scenarios.     

Genotype-by-environment interactions and adaptation to local temperature affect immunity and fecundity in Drosophila melanogaster

Natural populations of most organisms harbor substantial genetic variation for resistance to infection. The continued existence of such variation is unexpected under simple evolutionary models that either posit direct and continuous natural selection on the immune system or an evolved life history "balance" between immunity and other fitness traits in a constant environment. However, both local adaptation to heterogeneous environments and genotype-by-environment interactions can maintain genetic variation in a species. In this study, we test Drosophila melanogaster genotypes sampled from tropical Africa, temperate northeastern North America, and semi-tropical southeastern North America for resistance to bacterial infection and fecundity at three different environmental temperatures. Environmental temperature had absolute effects on all traits, but there were also marked genotype-by-environment interactions that may limit the global efficiency of natural selection on both traits. African flies performed more poorly than North American flies in both immunity and fecundity at the lowest temperature, but not at the higher temperatures, suggesting that the African population is maladapted to low temperature. In contrast, there was no evidence for clinal variation driven by thermal adaptation within North America for either trait. Resistance to infection and reproductive success were generally uncorrelated across genotypes, so this study finds no evidence for a fitness tradeoff between immunity and fecundity under the conditions tested. Both local adaptation to geographically heterogeneous environments and genotype-by-environment interactions may explain the persistence of genetic variation for resistance to infection in natural populations.     

Experimental Evidence for Nutrition Regulated Stress Resistance in Drosophila ananassae

Background

The amount and quality of nutrients consumed by organisms have a strong impact on stress resistance, life-history traits and reproduction. The balance between energy acquisition and expenditure is crucial to the survival and reproductive success of animals. The ability of organisms to adjust their development, physiology or behavior in response to environmental conditions, called phenotypic plasticity, is a defining property of life. One of the most familiar and important examples of phenotypic plasticity is the response of stress tolerance and reproduction to changes in developmental nutrition. Larval nutrition may affect a range of different life-history traits as well as responses to environmental stress in adult.

Principal Findings

Here we investigate the effect of larval nutrition on desiccation, starvation, chill-coma recovery, heat resistance as well as egg to adult viability, egg production and ovariole number in Drosophila ananassae. We raised larvae on either protein rich diet or carbohydrate rich diet. We found that flies consuming protein rich diet have higher desiccation and heat shock resistance whereas flies developed on carbohydrate rich diet have higher starvation and cold resistance. Egg production was higher in females developed on protein rich diet and we also found trade-off between egg production and Egg to adult viability of the flies. Viability was higher in carbohydrate rich diet. However, sex specific viability was found in different nutritional regimes. Higher Egg production might be due to higher ovariole number in females of protein rich diet.

Conclusion

Thus, Drosophila ananassae adapts different stress tolerance and life-history strategies according to the quality of the available diet, which are correlated with phenotypic adjustment at anatomical and physiological levels.     

Effect of sodium channel abundance on Drosophila development,reproductive capacity and aging

The voltage-gated Na⁺ channels (VGSC) are complex membrane proteins responsible for generation and propagation of the electrical signals through the brain, the skeletal muscle and the heart. The levels of sodium channels affect behavior and physical activity. This is illustrated by the maleless mutant allele (mle^napts) in Drosophila, where the decreased levels of voltage-gated Na⁺ channels cause temperature-sensitive paralysis.

Here, we report that mle^napts mutant flies exhibit developmental lethality, decreased fecundity and increased neurodegeneration. The negative effect of decreased levels of Na⁺ channels on development and ts-paralysis was more pronounced at 18 and 29°C than at 25°C, suggesting particular sensitivity of the mle^napts flies to temperatures above and below normal environmental conditions. Similarly, longevity of mle^napts flies was unexpectedly short at 18 and 29°C compared with flies heterozygous for the mle^napts mutation. Developmental lethality and neurodegeneration of mle^napts flies was partially rescued by increasing the dosage of para, confirming a vital role of Na⁺ channels in development, longevity and neurodegeneration of flies and their adaptation to temperatures.     

LONG-TERM LABORATORY EVOLUTION OF A GENETIC LIFE-HISTORY TRADE-OFF IN DROSOPHILA MELANOGASTER. 1. THE ROLE OF GENOTYPE-BY-ENVIRONMENT INTERACTION

Trade-offs among life-history traits are often thought to constrain the evolution of populations. Here we report the disappearance of a trade-off between early fecundity on the one hand, and late-life fecundity, starvation resistance, and longevity on the other, over 10 yr of laboratory selection for late-life reproduction. Whereas the selected populations showed an initial depression in early-life fecundity, they later converged upon the controls and then surpassed them. The evolutionary loss of the trade-off among life-history traits is considered attributable to the following factors: (1) the existence of differences in the culture regimes of the short- and long-generation populations other than the demographic differences deliberately imposed; (2) adaptation of one or both of these sets of populations to the unique aspects of their culture regimes; (3) the existence of an among-environment trade-off in the expression of early fecundity in the two culture regimes, as reflected in assays that mimic those regimes. The trade-off between early and late-life reproductive success, as manifest among divergently selected populations, is apparent or not depending on the assay environment. This demonstration that strong genotype-by-environment interactions can obscure a fundamental trade-off points to the importance of controlling all aspects of the culture regime of experimental populations and the difficulty of doing so even in the laboratory.     

Evolution of increased adult longevity in Drosophila melanogaster populations selected for adaptation to larval crowding

In holometabolous animals such as Drosophila melanogaster, larval crowding can affect a wide range of larval and adult traits. Adults emerging from high larval density cultures have smaller body size and increased mean life span compared to flies emerging from low larval density cultures. Therefore, adaptation to larval crowding could potentially affect adult longevity as a correlated response. We addressed this issue by studying a set of large, outbred populations of D. melanogaster, experimentally evolved for adaptation to larval crowding for 83 generations. We assayed longevity of adult flies from both selected (MCUs) and control populations (MBs) after growing them at different larval densities. We found that MCUs have evolved increased mean longevity compared to MBs at all larval densities. The interaction between selection regime and larval density was not significant, indicating that the density dependence of mean longevity had not evolved in the MCU populations. The increase in longevity in MCUs can be partially attributed to their lower rates of ageing. It is also noteworthy that reaction norm of dry body weight, a trait probably under direct selection in our populations, has indeed evolved in MCU populations. To the best of our knowledge, this is the first report of the evolution of adult longevity as a correlated response of adaptation to larval crowding.     

Intraspecific variability in egg maturation patterns and associated life‐history trade‐offs in a polyembryonic parasitoid wasp

1. Life‐history theory predicts a trade‐off between the resources allocated to reproduction and those allocated to survival. Early maturation of eggs (pro‐ovigeny) is correlated with small body size and low adult longevity in interspecific comparisons among parasitoids, demonstrating this trade‐off. The handful of studies that have tested for similar correlations within species produced conflicting results. 2. Egg maturation patterns and related life‐history traits were studied in the polyembryonic parasitoid wasp, Copidosoma koehleri (Hymenoptera: Encyrtidae). Although the genus Copidosoma was previously reported to be fully pro‐ovigenic, mean egg loads of host‐deprived females almost doubled within their first 6 days of adulthood. 3. The initial egg‐loads of newly emerged females were determined and age‐specific realised fecundity curves were constructed for their clone‐mate twins. The females' initial egg loads increased with body size, but neither body size nor initial egg load was correlated with longevity and fecundity. 4. The variation in initial egg loads was lowest among clone‐mates, intermediate among non‐clone sisters and highest among non‐sister females. The within‐clone variability indicates environmental influences on egg maturation, while the between‐clone variation may be genetically based. 5. Ovaries of host‐deprived females contained fewer eggs at death (at ～29 days) than on day 6. Their egg loads at death were negatively correlated with life span, consistent with reduced egg production and/or egg resorption. Host deprivation prolonged the wasps' life span, suggesting a survival cost to egg maturation and oviposition. 6. It is concluded that adult fecundity and longevity were not traded off with pre‐adult egg maturation.     

Reproductive biology of the isopod Excirolana braziliensis at the southern edge of its geographical range

A full analysis of the reproductive biology of the isopod Excirolana braziliensis Richardson 1912 was conducted in a sandy beach of Uruguay, located at the southernmost edge of its distributional range in the Atlantic Ocean. Reproductive and recruitment periods of E. braziliensis were concentrated in austral summer. Females with oostegites appeared in November, whereas total biomass, individual sizes and fecundity of ovigerous females peaked between December and January. These concurrent traits were responsible for the significant peak of juveniles in January. The size at maturity was 9.88 mm. Four embryonic developmental stages were described and identified: mean length linearly increased from stages I to III, whereas dry weight exponentially decreased from stages I to IV. The high reproductive output (0.41–0.58), reported for the first time in this isopod, exceeds the rates documented for other isopods. Reproduction of E. braziliensis at the southern edge of its range is semelparous: females produce one brood during the reproductive season, exhaust their energy reserves during incubation, and probably die at the end of the reproductive season. A macroscale comparison suggests that E. braziliensis at the southern edge of its range counteracts its narrow reproductive period by a short incubation period with larger individual mature female and embryo sizes, higher fecundity and a higher percentage of ovigerous females than in subtropical and tropical populations. These extreme reproductive indicators could be attributed to the internal retention of embryos that assures offspring survival, coupled with a high adaptation capability to environmental variations across its range.