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We have studied fluctuating asymmetry (FA), as indicator of developmental stability, and between-individual variation, as surrogate of developmental canalization (DC), in long bones (humerus, ulna, radius, femur, tibia) of 72 wild-living adult-sized brown haresLepus europaeus Pallas, 1778 with variable individual heterozygosity (H).H was calculated from 13 polymorphic allozyme loci. According to the “over-dominance hypothesis”, we expected increased developmental stability and canalization at higherH-levels. But at the individual level we did not find any significant correlation between overall FA (FAI) andH. Also, standard deviations (SD) of mean length (over both body sides) of bones did not differ between individuals from two intentionally created groups of hares, namely one with high and one with lowH. FA-indices and variances of FA-indices of bone lengths did not differ significantly when compared between two intentionally created groups of hares with high and low SD of bone lengths, respectively. These latter findings suggest that developmental stability and DC are two separate or partly separate mechanisms of developmental homeostasis in the studied appendicular skeleton, and thatH has no traceable effect on develop-mental homeostasis. If there is still such an effect, it should be clearly smaller than a possibly combined effect of (presently uncontrolled) environmental stressors.
Regular development of morphology is challenged by any environmental influence that draines energy from developing individuals. For half a century biologists have recognized that developmental stability, measured as the capability to regulate development of normal morphological structures, is influenced by genetic variation. This review considers the influence of enzyme polymorphism upon developmental stability. Empirical studies in a wide variety of animals have reported morphological variability and bilateral asymmetry to decrase with the heterozygosity of enzyme polymorphisms. A controversy focuses on the question of whether enzyme polymorphisms directly influence energy budgets and metabolism, or whether the enzymes are neutral markers that either reveal variation in levels of inbreeding or are in linkage disequilibrium with genes directly influencing development. Another controversy focuses on whether the relationships between enzyme heterozygosity and development stability, most commonly reported in poikilotherms, will also be found in homeotherms. These controversies are addressed by considering recent empirical studies of enzyme polymorphism and developmental stability. Kinetic and physiological studies have now established that enzyme polymorphisms can have a major impact upon flux through metabolic pathways and physiological variation. Enzyme heterozygosity is associated with resistance to parasites, which may decrease developmental stability. Enzyme heterozygosity is also related to secondary sexual characters, such as the size and symmetry of horns in white-tailed deer and the size of horns in bighorn sheep. Because symmetry is so important in sexual selection, the development of secondary sexual characters may yield important insights to the relationship between heterozygosity and developmental stability. Many empirical observations are consistent with hypothesis that enzymes have a direct influence on developmental stability. It is not likely that the relationship between heterozygosity and developmental stability will differ substantially between poikilotherms and homeotherms.
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