Apomixis is a clonal propagation method that produces offspring identical to the mother plant. With this feature, superior traits could be maintained over generations. However, our knowledge about apomixis is limited. In this study, we analyzed several apomictic Boechera embryologically to learn the details of apomixis. Meanwhile, we designed chemical treatments to successfully induce sex in apomictic plants and apomixis in sexual plants. Our experiments suggest that sex and apomixis coexist in plants and that sexual and apomictic reproduction are switchable by treating with specific chemicals.

In flowering plants, apomixis is asexual reproduction by seeds. Apomixis allows the production of offspring with the same genetic characteristics as the mother plant. Fertilization is not required. Apomixis could become a tool for naturally cloning high-yielding crop hybrids through their own seed. However, apomixis does not occur in major crop plants, except for citrus. In the present study, genes that might cause apomixis in naturally occurring apomictic plants were investigated. Sexual and apomictic species of the genus Boechera were exposed to stressed and non-stressed conditions. Effects of these treatments on the expression of apomixis was then measured. Stress triggered an increase in the frequency of sexual development in apomictic plants, but continuation of sexual development to form sexual seeds did not occur. Stress also triggered alterations in the expression of stress-related genes.

<p>What evolutionary processes and ecological patterns underlie the maintenance of asexual reproduction in natural populations? Although a vast body of literature offers theory to explain the existence of sexual and asexual reproduction, there has been little study of these forms of reproduction in the natural environment. In this dissertation I use a combination of field experiments, greenhouse studies, and genetic techniques to answer this question in the model plant system Boechera. </p><p>In Chapter 1, I review the utility of this system for studying ecological and evolutionary questions in general. Boechera offers an array of genetic and genomic tools, facilitated in part by a close evolutionary relationship with the model plant Arabidopsis thaliana, as well as undisturbed ecology and habitat that extends across much of North America. Additionally, the presence of apomixis (asexual reproduction via parthenogenetic formation of seeds) at the diploid level makes Boechera an ideal system for studying sex without the often-confounding factor of polyploidy; Boechera is one of very few plant groups in which this is possible. </p><p>In Chapter 2, I use a combination of microsatellite markers, flow cytometry, chromosome squashes, and morphological work to characterize apomixis, polyploidy, and species diversity in over 100 natural populations collected from central Idaho and western Montana. As in many other apomictic systems, I find that apomixis in Boechera is strongly linked to hybridization between species or between genetically divergent intraspecific lineages. I then explore associations between apomixis and ecological and topographical variables, as well as variables underlying differentiation between apomictic and sexual lineages. I find that ecological variables associated with apomixis are largely in congruence with the hypothesis of geographic parthenogenesis, and that geographic parthenogenesis is likely driven by the consequences of interspecific hybridization. We also find that apomixis is linked with disturbance and slope, with apomicts occurring in flatter locations than sexuals. </p><p>In Chapter 3, I use a large-scale field experiment comprising three years of data from two cohorts of sexual and asexual lineages to compare fitness between these two groups. I find that, despite herbivory levels that are much higher in apomicts than sexuals, apomictic fitness is consistently higher than sexual. Viability selection strongly favors apomicts, which results in a total fitness advantage for apomicts, despite variable fecundity selection. Selection varies in intensity between cohorts and among gardens. The results of a complementary greenhouse experiment show that the effects of herbivory differ by reproductive mode. Together, these experiments suggest that Red Queen dynamics may contribute to the coexistence of sex and asex in this group.</p><p>In Chapter 4, I use inter- and intraspecific F2 crosses to conduct a greenhouse study and a field experiment to explore the effects of hybridization and heterozygosity on fitness. I find that heterozygosity is favored in the field, with viability selection strongly favoring outcrossed over inbred lineages. However, hybridization results in lower survival, reproduction, and total fitness of interspecific F2 crosses, although hybrids that do reproduce produce more fruits than selfed parental lineages of both species, resulting in fecundity selection for hybrids. It is clear that the benefits of apomixis are due to hybridization, as hybrids are less fit overall. Evidence for both heterosis and outbreeding depression, dependent on lineage and on trait, are found in the field; these phenomena are not clearly associated with geographic distance between parental populations. Ongoing SNP genotyping will facilitate assessment of heterozygosity-fitness correlations as well as correlation of fitness and heterozygosity.</p> / Dissertation

Boechera is a large genus of flowering plants whose taxa are found primarily in North America. Boechera vivariensis (S.L. Welsh) W.A. Weber (the Park rockcress) is restricted to the Uintah Basin on Weber sandstone substrates in the vicinity of Dinosaur National Monument and Blue Mountain. The nomenclature of Park rockcress is significantly impacted by the discovery that the type collections of the taxon represent a rare, apomictic diploid resulting from the hybridization between B. thompsonii and an undescribed sexual diploid (to be called Boechera caeruleamontana sp. nov. Allphin and Windham). As a result, greater information is needed regarding how B. vivariensis and B. caeruleamontana. are distributed geographically in the region of Dinosaur National Monument and surrounding areas. Thus, we performed genetic analyses on leaf samples taken from over 50 individuals at known sites of B. vivariensis throughout its geographic range. Individuals from each site were also compared morphologically. We also compared associated plant communities at each site and characterized the soils. In our thorough sampling, we did not pick up B. vivariensis. All individuals sampled belonged to B. caeruleamontana, suggesting that most individuals previously assigned to B. vivariensis, are actually representative of B. caeruleamonanta. Populations of B. caeruleamontana were genetically diverse compared to other Boechera species, most likely indicative of its insect pollination strategy. However, all populations had lower heterozygosity than expected based upon Hardy-Weinberg expectations. Reproductive and genetic data indicated that populations are showing signs of inbreeding. The population at Jones Hole Fish Hatchery was most unique genetically, morphologically, and reproductively.

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