The concept of developmental buffering, which can be subdivided into canalisation and developmental stability, is very important in studies on evolution and development alike. The development of an organism needs to be buffered against genetic and environmental perturbations, because the fitness of an organism is likely to depend on the ability to produce a functioning phenotype in a given environment. On the other hand, too much buffering may limit the possibility of responding adequately to changes in the environment, and therefore impose severe constraints on the potential for evolution. Heat-shock protein Hasp is involved in both environmental and genetically buffering, which has been demonstrated in both animals and plants. By normally hiding genetic variation that would not be neutral when unmasked, and by making such variation available to natural selection under stress conditions, Hasp actually provides a mechanism for storing genetic variation. The functioning of Hasp has been much-publicized and although novel, it is by no means established that it is unique, and that it is a key factor in overall buffering against different forms of stress. The main aim of the proposed study is to investigate the existence and nature of factors (loci) other than Hasp that may be involved in buffering against genetic and environmental perturbations. We plan to screen a substantial part of the genome of the model organism D.melanogaster by deficiency mapping, with which it is possible to detect the effects of the absence of loci on developmental buffering, and to examine buffering mechanisms that evolved in the presence of different stress factors. Once loci have been found and identified, we will test whether developmental stability and canalisation share a common mechanism in buffering development. The effects of the putative loci on buffering will be examined by investigating patterns of wing shape variation by means of geometric morph metrics.
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