A number of hypotheses have been proposed about the association between developmental stability, phenotypic variability, heritability, and environmental stress. Stress is often considered to increase both the asymmetry and phenotypic variability of bilateral traits, although this may depend on trait heritability. Empirical studies of such associations often yield inconsistent results. This may reflect the diversity of traits and conditions used or a low repeatability of any associations. To test for repeatable associations between these variables, multiply replicated experiments were undertaken on Drosophila melanogaster using a combination stress at the egg, larval, and adult stages of reduced protein, ethanol in the medium, and a cold shock. Both metric and meristic traits were measured and levels of heritable variation for each trait estimated by maximum likelihood and parent-offspring regression over three generations. Trait means were reduced by stress, whereas among-individual variation increased. Fluctuating asymmetry (FA) was increased by stress in some cases, but few comparisons were significant. Only one trait, orbital bristle, showed consistent increases in FA. Changes in trait means, trait phenotypic variability, and developmental stability as a result of stress were not correlated. Extreme phenotypes tended to have higher levels of FA, but only the results for orbital bristles were significant. All traits had low to intermediate heritabilities except orbital bristle, which showed no heritable variation. Only traits with low heritability and high levels of phenotypic variability may show consistent increases in FA under stress. Overall, the independence of phenotypic variability, plasticity, and the developmental stability of traits extend to changes in these measures under stressful conditions.
In insects, the fluctuating asymmetry of bilaterally symmetrical traits has been suggested as an indicator of environmental stress because asymmetry is expected to increase when stressful conditions disturb the normal development of organisms. However, the extensive literature on asymmetry-stress associations is indeterminate. Here we contrast changes in asymmetry with changes in an alternate stress indicator, the shape of insect wings. The development of wing shape involves numerous genes that act throughout egg-to-adult development, so stresses that act at a specific time could alter shape in specific ways. Shape changes, as measured by the Procrustes technique, were considered in five data sets: exposure of Drosophila melanogaster (Meigen) to multiple stresses involving ethanol, low nutrition and cold shocks; exposure of a chironomid (Chironomus tepperi (Skuse)), a blowfly (Lucilia cuprina (Wiedemann)), and lightbrown apple moth (Epiphyas postvittana (Walker)) to pesticides; and development of C. tepperi under saline conditions. All these conditions influenced viability and development time. In none of these cases was a change in symmetry of wing size or wing shape detected. In contrast, in four of the five data sets there was a change in wing shape. These results suggest that wing shape may be altered more commonly by stress than trait asymmetry. Wing-shape monitoring may be useful in detecting stressful environmental conditions during development, at least under controlled conditions.
Some studies have found intermediate heritabilities for fluctuating asymmetry (FA) in traits, but almost all of these are flawed and/or based on laboratory experiments. We therefore tested if there was heritable variation for FA in bristle and wing traits in three field collections of Drosophila melanogaster by rearing F s from field flies under laboratory conditions. One of the collections was reared to the F generation in the laboratory to compare heritability estimates from the laboratory with those from the field-laboratory comparison. Trait means indicated an increase in size under laboratory rearing. FAs increased in one collection, decreased in another collection, and showed no changes in the third collection under laboratory rearing. FAs from the collections tended to converge under laboratory conditions. Morphological traits were heritable under field conditions. However, FA was not significantly heritable for any of the individual traits or when FA was determined by combining traits. Comparisons of the two laboratory generations showed that FA heritability was low under laboratory conditions, in contrast to the morphological traits themselves. These findings suggest a very low heritability for FA in field and laboratory Drosophila. FA in bristle and wing traits may therefore be a poor indicator of genetic quality in Drosophila.
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