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Most heat shock proteins (Hsp) function as molecular chaperones that help organisms to cope with stress of both an internal and external nature. Here, we review the recent evidence of the relationship between stress resistance and inducible Hsp expression, including a characterization of factors that induce the heat shock response and a discussion of the associated costs. We report on studies of stress resistance including mild stress, effects of high larval densities, inbreeding and age on Hsp expression, as well as on natural variation in the expression of Hsps. The relationship between Hsps and life history traits is discussed with special emphasis on the ecological and evolutionary relevance of Hsps. It is known that up-regulation of the Hsps is a common cellular response to increased levels of non-native proteins that facilitates correct protein folding/refolding or degradation of non-functional proteins. However, we also suggest that the expression level of Hsp in each species and population is a balance between benefits and costs, i.e. a negative impact on growth, development rate and fertility as a result of overexpression of Hsps. To date, investigations have focused primarily on the Hsp70 family. There is evidence that representatives of this Hsp family and other molecular chaperones play significant roles in relation to stress resistance. Future studies including genomic and proteonomic analyses will increase our understanding of molecular chaperones in stress research.

Upper thermal limits vary less than lower limits among related species of terrestrial ectotherms. This pattern may reflect weak or uniform selection on upper limits, or alternatively tight evolutionary constraints. We investigated this issue in 94 Drosophila species from diverse climates and reared in a common environment to control for plastic effects that may confound species comparisons. We found substantial variation in upper thermal limits among species, negatively correlated with annual precipitation at the central point of their distribution and also with the interaction between precipitation and maximum temperature, showing that heat resistance is an important determinant of Drosophila species distributions. Species from hot and relatively dry regions had higher resistance, whereas resistance was uncorrelated with temperature in wetter regions. Using a suite of analyses we showed that phylogenetic signal in heat resistance reflects phylogenetic inertia rather than common selection pressures. Current species distributions are therefore more likely to reflect environmental sorting of lineages rather than local adaptation. Similar to previous studies, thermal safety margins were small at low latitudes, with safety margins smallest for species occupying both humid and dry tropical environments. Thus, species from a range of environments are likely to be at risk owing to climate change. Together these findings suggest that this group of insects is unlikely to buffer global change effects through marked evolutionary changes, highlighting the importance of facilitating range shifts for maintaining biodiversity.niche conservatism | stress resistance | thermal adaptation | evolutionary history | space T emperatures are expected to rise across the globe over the coming decades and centuries (1), and many studies suggest the potential for range shifts/reductions in many species (2). When assessing the likely impact of temperature changes on species survival, an implicit assumption is that the thermal environment shapes resistance to temperature extremes and thus dictates species range limits. Nevertheless, few studies have directly tested for such links between physiological upper thermal limits of ectothermic species and temperature conditions within their geographic range (3, 4). A related, little-investigated key point for predicting species range responses to climate change is whether upper thermal limits are modifiable through plastic and/or evolutionary responses (5, 6). Ideally, evolutionary and plastic responses within and across generations, including short-term hardening and acclimation, should be separated, as through a common garden approach whereby species are kept under controlled laboratory conditions (7,8). Without controlling for plastic responses, it is not possible to distinguish adaptive evolutionary responses, and species may erroneously seem close to their upper thermal thresholds (9-12), biasing extinction risk estimates.Furthermore, by examining the evolution of upper thermal limits (heat re...

Laboratory studies on Drosophila have revealed that resistance to one environmental stress often correlates with resistance to other stresses. There is also evidence on genetic correlations between stress resistance, longevity and other fitness‐related traits. The present work investigates these associations using artificial selection in Drosophila melanogaster. Adult flies were selected for increased survival after severe cold, heat, desiccation and starvation stresses as well as increased heat‐knockdown time and lifespan (CS, HS, DS, SS, KS and LS line sets, respectively). The number of selection generations was 11 for LS, 27 for SS and 21 for other lines, with selection intensity being around 0.80. For each set of lines, the five stress‐resistance parameters mentioned above as well as longevity (in a nonstressful environment) were estimated. In addition, preadult developmental time, early age productivity and thorax length were examined in all lines reared under nonstressful conditions. Comparing the selection lines with unselected control revealed clear‐cut direct selection responses for the stress‐resistance traits. Starvation resistance increased as correlated response in all sets of selection lines, with the exception of HS. Positive correlated responses were also found for survival after cold shock (HS and DS) and heat shock (KS and DS). With regard to values of resistance across different stress assays, the HS and KS lines were most similar. The resistance values of the SS lines were close to those of the LS lines and tended to be the lowest among all selection lines. Developmental time was extended in the SS and KS lines, whereas the LS lines showed a reduction in thorax length. The results indicate a possibility of different multiple‐stress‐resistance mechanisms for the examined traits and fitness costs associated with stress resistance and longevity.

Species distributions are often constrained by climatic tolerances that are ultimately determined by evolutionary history and/or adaptive capacity, but these factors have rarely been partitioned. Here, we experimentally determined two key climatic niche traits (desiccation and cold resistance) for 92-95 Drosophila species and assessed their importance for geographic distributions, while controlling for acclimation, phylogeny, and spatial autocorrelation. Employing an array of phylogenetic analyses, we documented moderate-to-strong phylogenetic signal in both desiccation and cold resistance. Desiccation and cold resistance were clearly linked to species distributions because significant associations between traits and climatic variables persisted even after controlling for phylogeny. We used different methods to untangle whether phylogenetic signal reflected phylogenetically related species adapted to similar environments or alternatively phylogenetic inertia. For desiccation resistance, weak phylogenetic inertia was detected; ancestral trait reconstruction, however, revealed a deep divergence that could be traced back to the genus level. Despite drosophilids' high evolutionary potential related to short generation times and high population sizes, cold resistance was found to have a moderate-to-high level of phylogenetic inertia, suggesting that evolutionary responses are likely to be slow. Together these findings suggest species distributions are governed by evolutionarily conservative climate responses, with limited scope for rapid adaptive responses to future climate change. K E Y W O R D S :Ancestral trait reconstruction, evolutionary history, niche conservatism, phylogenetic signal, species distribution, stress resistance.

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Summary 1. Thermal tolerance may limit and therefore predict ectotherm geographic distributions. However, which of the many metrics of thermal tolerance best predict distribution is often unclear, even for drosophilids, which constitute a popular and well-described animal model. 2. Five metrics of cold tolerance were measured for 14 Drosophila species to determine which metrics most strongly correlate with geographic distribution. The species represent tropical to temperate regions but all were reared under similar (common garden) conditions (20°C). The traits measured were: chill coma temperature (CT min ), lethal temperature (LTe 50 ), lethal time at low temperature (LTi 50 ), chill coma recovery time (CCRT) and supercooling point (SCP). 3. Measures of CT min , LTe 50 and LTi 50 proved to be the best predictors to describe the variation in realized latitudinal distributions (R 2 = 0Á699, R 2 = 0Á741 and 0Á550, respectively) and estimated environmental cold exposure (R 2 = 0Á633, R 2 = 0Á641 and 0Á511, respectively).Measures of CCRT also correlated significantly with estimated minimum temperature (R 2 = 0Á373), while the SCP did not. These results remained consistent after phylogenetically independent analysis or when applying nonlinear regression. Moreover, our findings were supported by a similar analysis based on existing data compiled from the Drosophila cold tolerance literature. 4. Trait correlations were strong between LTe 50 , LTi 50 and CT min , respectively (0Á83 > R 2 > 0Á55). However, surprisingly, there was only a weak correlation between the entrance into coma (CT min ) and the recovery from chill coma (CCRT) (R 2 = 0Á256). 5.Considering the findings of the present study, data from previous studies and the logistical constraints of each measure of cold tolerance, we conclude that CT min and LTe 50 are superior measures when estimating the ecologically relevant cold tolerance of drosophilids. Of these two traits, CT min requires less equipment, time and animals and thereby presents a relatively fast, simple and dynamic measure of cold tolerance.

Summary1. Thermal adaptation was investigated in the fruitfly Drosophila buzzatii Patterson and Wheeler. Two natural populations originating from a high-and a low-temperature environment, respectively, were compared with respect to Hsp70 (heat shock protein) expression, knock-down resistance and heat shock resistance. 2. Three main hypotheses were tested: (i) The expression level of Hsp70 in flies from the high-temperature habitat should be down-regulated relative to flies from the colder habitat. (ii) Flies having higher Hsp70 expression levels should be weakened most by a hardening treatment and go faster into coma, as Hsp70 level reflects stress intensity, and therefore display reduced heat knock-down resistance. (iii) Heat shock resistance should be increased in the population with highest Hsp70 expression because the level of Hsp70 is positively associated with this trait. 3. The results generally matched the hypotheses. Hsp70 expression was reduced in the high-temperature population. Knock-down resistance was higher in the hightemperature population and survival after heat shock was lower in the high-temperature population. 4. This study showed genetic differences in thermal tolerance between populations, indicating that high temperature in nature may be an important selective factor. Moreover, knock-down resistance in this study seems to be a more relevant trait than standard heat shock resistance for identifying thermal adaptation in natural populations.