The colonization of novel environments requires organisms to shift their trait means in response to differing abiotic and biotic conditions in order to survive and persist. This response can be done via phenotypic plasticity (a trait shift in response to the environment), adaptation (a trait shift due to genetic change), or both strategies can be used together, with plasticity "buying time" for adaptation to occur. The colonization of novel environments is especially important to the establishment of agricultural weeds worldwide, which thrive in these extreme environments of intense competition and frequent disturbance. In this dissertation, I address the establishment and evolution of a harmful agricultural weed, weedy radish (Raphanus raphanistrum), as well as its divergence from a wild relative of the same species, the native radish ecotype. I first investigated the hypothesis of phenotypic plasticity "buying time" for adaptation to agricultural fields in weedy radish. Using growth chambers to simulate the ancestral (native) and derived (weedy) environments of weedy radish, I performed a reciprocal transplant with the weedy and native radish ecotypes. I found phenotypic plasticity between environments and genetic divergence between ecotypes to be equally common among traits, suggesting similar importance of plasticity and adaptation in weedy radish establishment. Further, in the majority of traits that were both plastic and differentiated between ecotypes, the direction of change matched, with the weedy environment producing phenotypic shifts in the direction of the weedy ecotype mean. This suggests plasticity in these traits may have enabled the subsequent adaptation and ecotype differentiation, supporting the buying-time hypothesis.Next, I explored the role of the plant hormone Gibberellic Acid (GA) in the evolution of weedy radish. Using exogenous application of GA both in the greenhouse and in weedy and native growth chamber environments, I found evidence that there has been an evolutionary change in the role of GA in trait expression between the two ecotypes. Namely, weedy radish is less responsive to GA application than native radish, suggesting either upregulation in GA production in weeds, or a lower level of GA required to enable gene expression in the weedy ecotype. This change in gene regulation by GA may have been important in the evolution of weedy radish in the agricultural field.Finally, I assessed the likelihood of weedy radish diverging from a native ancestor via adaptive evolution. I found that adaptive evolution was likely in the establishment of weedy radish due to increased fitness of the weedy ecotype compared to the native ecotype in the agricultural field. I also found traits under directional selection in the native ecotype, with the key takeaway that faster flowering is adaptive in the agricultural fields. I finally looked at the ability of weedy radish to evolve advanced flowering in the agricultural field via standing genetic variance by artificially selecting for early flowering in native radish. I found that in only two generations of selection, native populations significantly advanced their flowering time, supporting the notion of weedy radish rapidly adapting to agricultural conditions via standing genetic variation alone. Taken together, these findings work to piece together the evolutionary history of weedy radish, providing insight into its mechanisms of establishment. This work also contributes to our overall understanding of rapid evolution and phenotypic plasticity in the colonization of novel environments, in agricultural weeds and beyond.
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