Patterns of longevity and fecundity at two temperatures in a set of heat-selected recombinant inbred lines of Drosophila melanogaster.

Quantitative trait loci (QTL) were mapped for longevity and fecundity at two temperatures, 20 and 30 °C, in two sets of recombinant inbred lines (RIL) highly differing in thermotolerance. Early fecundity (EF) and longevity showed a negative association between temperatures. For instance, longevity was higher and fecundity was lower in the RIL panel showing higher life span at 30 °C. One X-linked QTL (7B3-12E) co-localized for longevity and EF at 20 °C, with one QTL allele showing a positive additive effect on longevity and a negative effect on EF. The across-RIL genetic correlation between longevity and EF was not significant within each temperature, and most QTL that affect life span have no effect on EF at each temperature. EF and longevity can mostly be genetically uncoupled in the thermotolerance-divergent RIL within each temperature as opposed to between temperatures. QTL were mostly temperature specific, although some trait-specific QTL showed possible antagonistic effects between temperatures.

Heat-stress survival in the pre-adult stage of the life cycle in an intercontinental set of recombinant inbred lines of Drosophila melanogaster.

In insects, pre-adult stages of the life cycle are exposed to variation in temperature that may differ from that in adults. However, the genetic basis for adaptation to environmental temperature could be similar between the pre-adult and the adult stages of the life cycle. Here, we tested quantitative trait loci (QTL) for heat-stress survival in larvae of Drosophila melanogaster, with and without a mild-heat-stress pre-treatment. Two sets of recombinant inbred lines derived from lines artificially selected for high and low levels of knockdown resistance to high temperature in young flies were used as the mapping population. There was no apparent increase in heat-shock survival between heat-pretreated and non-pretreated larvae. There was a positive correlation between the two experimental conditions of heat-shock survival (with and without a heat pre-treatment) except for males from one set of lines. Several QTL were identified involving all three major chromosomes. Most QTL for larval thermotolerance overlapped with thermotolerance QTL identified in previous studies for adults, indicating that heat-stress resistance is not genetically independent between life cycle stages because of either linkage or pleiotropy. The sign of the effects of some QTL alleles differed both between the sexes and between life stages.

X-linked QTL for knockdown resistance to high temperature in Drosophila melanogaster.

Knockdown Resistance to High Temperature (KRHT) is an adaptive trait of thermotolerance in insects. An interval mapping was performed on chromosome X of Drosophila melanogaster to search for quantitative trait loci (QTL) affecting KRHT. A backcross population was obtained from two lines that dramatically differ for KRHT. Microsatellites were used as markers. Composite interval mapping identified a large-effect QTL in the region of band 10 where putative candidate genes map. To further test for this QTL a set of recombinant (but non-inbred) lines was obtained from backcrosses between the parental lines used for the interval mapping. Recombinant line analysis confirmed that one QTL is targeted by band 10. We identify and discuss candidate loci contained within our QTL region.

Altitudinal patterns for longevity, fecundity and senescence in Drosophila buzzatii.

We tested for variation in longevity, senescence rate and early fecundity of Drosophila buzzatii along an elevational transect in Argentina, using laboratory-reared flies in laboratory tests performed to avoid extrinsic mortality. At 25 degrees C, females from lowland populations lived longer and had a lower demographic rate of senescence than females from highland populations. Minimal instead of maximal temperature at the sites of origin of population best predicted this cline. A very different pattern was found at higher test temperature. At 29.5 degrees C, longevity of males increased with altitude of origin of population. No clinal trend was apparent for longevity of females at 29.5 degrees C. There was evidence for a trade-off between early fecundity and longevity at non-stressful temperature (25 degrees C) along the altitudinal gradient. This trait association is consistent with evolutionary theories of aging. Population-by-temperature and sex-by-temperature interactions indicate that senescence patterns are expressed in environment specific ways.

Altitudinal variation for stress resistance traits and thermal adaptation in adult Drosophila buzzatii from the New World.

Multiple stress resistance traits were investigated in the cactophilic fly Drosophila buzzatii. Adults from seven populations derived from North-Western Argentina were compared with respect to traits relevant for thermal stress resistance and for resistance to other forms of environmental stress. The populations were collected along an altitudinal gradient spanning more than 2000 m in height, showing large climatic differences. The results suggest that knock-down resistance to heat stress, desiccation resistance and Hsp70 expression at a relatively severe stressful temperature best reflect thermal adaptation in this species. Furthermore, cold resistance seemed to be of less importance than heat resistance, at least for the adult life stage, in these populations. Clinal variation in thermal resistance traits over short geographical distances suggests relatively strong adaptive differentiation of the populations. This study provides the first evidence for altitudinal differentiation in stress-related traits, and suggests that Hsp70 expression level can be related to altitudinal clines of heat-stress resistance.

Behavioral differentiation in oviposition activity in Drosophila buzzatii from highland and lowland populations in Argentina: plasticity or thermal adaptation?

Highland populations of several Drosophila species in Argentina were active early in the afternoon in the field as opposed to populations from a much warmer lowland site, where flies were mainly active in the early evening prior to sunset. For one of these species, Drosophila buzzatii, we tested for a genetic component of activity differences by carrying out crosses within and between populations and measuring oviposition activity of the progeny in the laboratory. We found that activity in the highland population exceeded that in the lowland one during the midafternoon, whereas activity in the lowland population exceeded that in the highland one prior to the beginning of the dark period. Oviposition activity for the period corresponding to the field observations was regressed on the proportion of the genome derived from the highland population. This variable significantly predicted oviposition activity between 1400 and 1600 and between 2000 and 2200 h. Activity of both reciprocal crosses was intermediate and not significantly different from each other, suggesting that nuclear genetic, rather than cytoplasmic factors contribute to differences in oviposition activity between the populations. Two morphological, one genetic, and one stress resistance trait were also scored to examine whether temperature differences between environments were associated with other differences between populations. Wing length of wild-caught and laboratory-reared flies from the highland population significantly exceeded that in the lowland. Thorax length of laboratory-reared flies from the highland population also significantly exceeded that from the lowland. Chromosomal inversion frequencies differed significantly between the two populations with a fivefold reduction in the frequency of arrangement 2st in the highland as compared to the lowland population. This arrangement is known for its negative dose effect on size, and thus, the highland population has experienced a genetic change, perhaps as a result of adaptation to the colder environment, where body size and the frequency of arrangement 2st have changed in concert. Finally, a heat knockdown test revealed that the lowland population was significantly more resistant to high temperature than the highland one. In conclusion, we suggest that temperature has been an important selective agent causing adaptive differentiation between these two populations. We also suggest that the activity rhythms of the two populations have diverged as a consequence of behavioral evolution, that is, through avoidance of stressful temperatures as a mean of thermal adaptation.

Developmental time, body size and wing loading in Drosophila buzzatii from lowland and highland populations in Argentina.

Genotype-by-temperature interaction is a necessary condition for adaptive evolution of fitness traits as a response to temperature. Several fitness-related traits (developmental time, pre-adult survival, thorax and wing lengths, and wing loading) were measured in laboratory-reared D. buzzatii from four populations sampled at different altitudes in north-western Argentina: a lowland population (407 m a.s.l.), two populations from intermediate altitude (780 to 950 m a.s.l.), and a highland population (2380 m a.s.l.). Temperature is the main climatic difference between the collection sites: lowland but not highland populations are exposed to physiologically high temperatures during both spring and summer in nature. Three growth temperatures (20, 25 and 30 degrees C) were used to test for population-by-temperature interactions. Both developmental time and pre-adult survival exhibit highly significant population-by-temperature interaction. Pre-adult survival at 30 degrees C is significantly higher in lowland than in highland populations, but not so at lower growth temperatures (20 and 25 degrees C). Both wing length and wing loading show no population-by-temperature interaction, indicating that these traits are not the direct targets of thermal adaptation in nature. Wing loading is higher in highland than in lowland populations, suggesting that flight performance is subject to stronger selection in the highland population. This hypothesis is consistent with ecological observations in both types of populations. There is no obvious among-population relationship between developmental time and body size, even though both traits are related within populations in a well-known trade-off. Overall, thermal adaptation is evident for developmental time and pre-adult survival but not for size-related traits.