Genomic and Fitness Consequences of Hybridization between Cutthroat and Rainbow Trout
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Hybridization is an important and common evolutionary process that can contribute to diversification, adaptation, and speciation. When species hybridize, divergent genomes are combined through recombination and may result in phenotypic changes. Such phenotypic changes may be the result of differences in chromosomal structure or adaptive divergence between the parental species and may ultimately affect fitness. Understanding how phenotypes change following hybridization, as well as the genetic mechanisms responsible for changes is critical for understanding divergent selection, speciation, and identifying populations that may be at risk from hybridization. Here, the effect of hybridization between cutthroat (Oncorhynchus clarkii) and rainbow trout (O. mykiss) on fitness was investigated using three techniques. In the first chapter, fitness influencing traits were compared among individuals at various hybridization levels. In the second chapter, genomic changes that could affect fitness were identified in the hybrid relative to rainbow trout. And, in the third chapter, correlations between reproductive success and hybridization were investigated in a wild population, as well as the genomic and ecological mechanisms responsible for those changes. The first chapter of this dissertation aimed to identify how traits potentially involved in fitness (embryonic survival, ova size, ova energy concentration, sperm motility, burst swimming performance, juvenile survival, and juvenile growth) changed with hybridization between cutthroat and rainbow trout and whether those changes could explain previously observed reductions in reproductive success of individuals with increased rainbow trout ancestry. Using progeny from wild caught fish, differences in phenotypes based on hybridization were observed for embryonic survival, ova energy concentration, juvenile weight, and burst swimming based on ancestry. However, the correlations differed from previously observed patterns of reproductive success and likely do not explain declines in reproductive success associated with hybridization. The second chapter of this dissertation aimed to identify how hybridization affects the genome by identifying genomic regions with changes in recombination rates in the hybrid relative to rainbow trout as well as genomic areas with excess species-specific ancestry in the hybrid. Previous studies of hybridization have observed recombination suppression in genomic regions where structural differences, such as inversions or karyotype differences, exist between parental species. Such regions may retain groups of adaptive alleles. Additionally, adaptive divergence between the parental species may result in alleles that are preferentially selected in the hybrid progeny. Identification of regions with suppressed recombination or excess species-specific ancestry would provide insight into markers that may be important to fitness and that have differentially evolved in each of the parental species. In total, eight and seven chromosomes were identified to have changes in recombination rates in the hybrid female and male relative to O. mykiss. Recombination was suppressed in the hybrids on two chromosomes with known structural differences between the parental species. In addition, changes in recombination rates were observed on five chromosomes with high proportions of duplicated markers and may be due to increased homeologous chromosome pairing. Recombination patterns were similar between the sexes which suggests that hybridization affects recombination in the same way in females and males. Regions of excess species-specific ancestry covered 11 and 10% of the mapped genome in the female and male and regions of excess were evenly split between cutthroat trout and O. mykiss. Genetic drift may be responsible for much of the observed patterns of excess species-specific ancestry, but selection may also play a role. The aim of the third chapter of this dissertation was to identify the fitness consequences of hybridization, mechanisms responsible for the retention of hybridization, and genomic regions correlated with changes in reproductive success in a wild population of westslope cutthroat trout hybridized with non-native rainbow trout. Adult samples from a previous study, collected over a five year period, were sequenced at 3027 loci. Increased admixture from non-native rainbow trout had a strong, negative effect on reproductive success. A decline of 53% was observed for individuals with an increased genetic contribution of 0.20 from rainbow trout. Despite apparent strong selection against rainbow trout ancestry, hybridization appears to be maintained largely by the invasion of rainbow trout from outside populations as well as the relatively high fitness of few hybrid individuals. Ten loci correlated with reproductive success were identified in females. Seven of the ten loci were linked to chromosomes and three were positioned on chromosomes. Loci linked to reproductive success were identified on chromosomes with excess species-specific ancestry in hybrid progeny (RYHyb14 and RYHyb18) as well as chromosomes with a high proportion of duplicated markers (RYHyb02) and known Robertsonian polymorphism (RYHyb20). The research presented in this dissertation will elucidate our understanding of the phenotypic and genetic changes correlated with hybridization between rainbow and cutthroat trout as well as identify genetic and ecological mechanisms that may be responsible for those changes. In addition, results from this study provide insight into differences in adaptive divergence and markers that may be involved in the early stages of speciation in the wild. Results could be used by managers to identify populations that are at risk from hybridization.
- Fisheries