Temperature has dramatic evolutionary fitness consequences and is therefore a major factor determining the geographic distribution and abundance of ectotherms. However, the role that age might have on insect thermal tolerance is often overlooked in studies of behaviour, ecology, physiology and evolutionary biology. Here, we review the evidence for ontogenetic and ageing effects on traits of high-and low-temperature tolerance in insects and show that these effects are typically pronounced for most taxa in which data are available. We therefore argue that basal thermal tolerance and acclimation responses (i.e. phenotypic plasticity) are strongly influenced by age and/or ontogeny and may confound studies of temperature responses if unaccounted for. We outline three alternative hypotheses which can be distinguished to propose why development affects thermal tolerance in insects. At present no studies have been undertaken to directly address these options. The implications of these age-related changes in thermal biology are discussed and, most significantly, suggest that the temperature tolerance of insects should be defined within the age-demographics of a particular population or species. Although we conclude that age is a source of variation that should be carefully controlled for in thermal biology, we also suggest that it can be used as a valuable tool for testing evolutionary theories of ageing and the cellular and genetic basis of thermal tolerance.
SUMMARYMany endogenous factors influence thermal tolerance of insects. Among these, age contributes an important source of variation. Heat tolerance is typically high in newly eclosed insects, before declining dramatically. It is not known whether this phenomenon relates to cold tolerance also. In addition, the underlying mechanisms of this variation are unresolved. In this study, we tested whether cold tolerance declines in Drosophila melanogaster females aged from 0 to 5days. We also assessed whether expression (basal and induced) of eight stress genes (hsp22, hsp23, hsp40, hsp68, hsp70Aa, hsp83, Starvin and Frost) varied post-eclosion in correspondence with changes found in cold tolerance. We report that cold tolerance was very high at eclosion and then it rapidly declined in young flies. hsp23 and hsp68 showed a dramatic age-related variation of basal expression that was associated with cold tolerance proxies. Significant age-related plasticity of cold-induced expression was also found for hsp22, hsp23, hsp68, hsp70Aa, Frost and Starvin. Induced expression of hsp22 and hsp70Aa was high in newly enclosed phenotypes before declining dramatically, whilst opposite age-related patterns were found for hsp23, hsp68, Starvin and Frost. This study shows a marked within-stage variation in cold tolerance. The involvement of the stress genes in setting basal thermal tolerance is discussed.Supplementary material available online at http://jeb.biologists.org/lookup/suppl
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