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Abstract

Meiotic recombination is a highly conserved process found across a wide range of plants and animals. It is necessary for the proper segregation of chromosomes in many species and also creating genetic variation by creating new haplotype combinations. Despite these important roles, fine-scale recombination rates can vary drastically across species and even individuals. Threespine stickleback fish (Gasterosteus aculeatus) provide an excellent model to study fine-scale recombination rate evolution across short evolutionary timescales as well as to study sexually dimorphic recombination landscapes. This dissertation shows the first example of rapidly evolving recombination hotspots outside of mammals. Interestingly, recombination hotspots are enriched around transcription start sites rather than being targeted away from genic regions as expected under the PRDM9 model. This suggests a novel hotspot targeting mechanism is functioning in threespine stickleback. However, this enrichment is dependent on accurate gene annotations. Long read RNA sequencing (Iso-Seq) was then utilized to refine the gene annotations in threespine stickleback. The enrichment of hotspots around transcription start sites increased when using the refined start sites. Transcription start sites are expected to be in accessible chromatin regions, suggesting that hotspots and regions with increased recombination rates are targeted to accessible regions. The role of chromatin accessibility was further examined using the chromatin accessibility hypothesis for sexually dimorphic recombination landscapes. Across many species, males tend to have more recombination towards the ends of chromosomes compared to females. Chromatin accessibility was different between males and females where male meiotic tissues showed an enrichment of accessible regions towards the ends of chromosomes, matching male-specific recombination distributions. This enrichment was not due to testis-specific gene expression, suggesting as with hotspots, some other mechanism is directing recombination machinery to accessible regions. Together, this work presents significant contributions to understanding recombination rate evolution across short evolutionary timescales and shows how chromatin accessibility is important for recombination rate variation across multiple scales.

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