Two divergent taxa in the marine mussel genus Mytilus are largely isolated geographically and are routinely exposed to distinctly different thermal environments. We tested the hypothesis that the two taxa are physiologically differentiated with respect to temperature and examined the evolved adaptations allowing one of the taxa to exploit habitats where warm-temperate conditions prevail for prolonged periods. We first analyzed the physiological response to high temperature of mussels collected from a hybrid population containing members of both pure taxa, F, hybrids, and a variety of introgressed genotypes. The experimental temperature of 23°C was chosen to be permissive to the taxon that occurs in warm-temperate regions (Mytilus galloprovincialis) and restrictive to the cold-water taxon (Mytilus edulis). The results show that the two taxa are physiologically differentiated. Under the experimental conditions, M. galloprovincialis exhibited a threefold higher feeding rate and a slightly elevated metabolic rate compared with M. edulis. These differences did not result in a significant difference in net energy balance between the two taxa, probably because of an interaction between physiological response and food availability. However, M. galloprovincialis grew significantly faster in the field, indicating that the physiological differences observed in the laboratory also occur in nature. Numerous introgressed genotypes provided the opportunity to test for cosegregation between the physiological differences and four highly differentiated genetic markers. Two of the markers (esterase and octopine dehydrogenase) cosegregate with variation in feeding rate and shell growth and explained most of the physiological differences observed between taxa. A strong concordance existed between these two loci, suggesting that they may be linked and may mark segregation of the same linkage group. The results suggest that the physiological differentiation between these taxa may be controlled by a few genes (perhaps only one) each with large effect.
Sl'ucrlla lapillus from 15 sites 50 m to 21 km apart in S. Devon, S. W. England were analysed for allozyme variation at eight soluble enzyme loci. Sites were classified as either Exposed or Sheltered and a hierarchical analysis of allozyme variation carried out using Wright's Fixation index, FSr. Whelks from Sheltered sites segregated for alleles at the Est-.?, Lap-2, Pep-.? and Mdh-1 loci which were virtually absent from exposed sites resulting in high FST values (0.289-0.506) when all samples were included. Consequently mean heterozygosity of the Sheltered samples was roughly twice that of the Exposed. Between Sheltered sites the frequencies of these 'additional' alleles varied substantially, even on a microgeographic scale and were found to be highly correlated with exposure. Within individuals the presence of the 'sheltered' alleles was correlated with the possession of the 'sheltered' shell shape as indicated by length divided by aperture height. This must necessarily he the case, however, if both are related independently to exposure. Although these correlations imply the action of selection there is evidence within the data for stochastic factors affecting the pattern observed. Complicating the picture further is the correspondence observed in the distribution of the 'sheltered' alleles with the known distribution of chromosomal translocations. Phenotypic associations within individuals also support the hypothesis that variation at the Est-3, Lap-2, Mdh-1 and Pet-2 loci is related to variation in chromosome number. However, the nature of the relations of karyotype and allozyme variation with shell shape and exposure remain speculative.
Triploid oysters were induced using cytochalasin B upon retention of either the first (meiosis I triploids) or the second (meiosis II triploids) polar body in embryos from a single cohort derived from mixed parentage. Allozyme and microsatellite assays enabled the confirmation of both parentage and triploidy status in each oyster. Comparison of meiosis I triploids, meiosis II triploids and diploid siblings established that improved physiological performance in triploids was associated with increased allelic variation, rather than with the quantitative dosage effects of ploidy status. An unidentified maternal influence also interacted with genotype. Among full sibs, allelic variation measured as multi-locus enzyme heterozygosity accounted for up to 42% of the variance in physiological performance; significant positive influences were identified upon feeding rate, absorption efficiency, net energy balance and growth efficiency (= net energy balance divided by energy absorbed). Whilst allelic variation was greater in both meiosis I and meiosis II triploids than in diploid siblings, both allelic variation and net energy balance were highest in triploids induced at meiosis I. This suggests that it may be preferable to induce triploidy by blocking meiosis I, rather than meiosis II as has traditionally been undertaken during commercial breeding programmes.
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