The fate of any lineage is contingent on the rate at which its genome changes over time. Genome dynamics are influenced by patterns of mutation and recombination. Mutations as the raw force of variation can be acted on independently during exchanges of homologous genetic regions via meiotic recombination. While molecular evolution in sexual lineages is impacted by both mutation and recombination, asexual lineage fate is primarily influenced by the mutation rate; recombination is often altered or absent in asexuals. Although multiple studies show accelerated mutation accumulation in asexual lineages that have lost recombination, virtually nothing is known about rate patterns when meiosis is retained. Here, I use parasitic wasps in genus Diachasma to investigate genome evolution in a recently-derived asexual lineage. I provide evidence that asexual Diachasma possess a canonical set of meiosis genes as well as high levels of genomic homozygosity. Taken together, these observations support an active, albeit modified, form of meiosis in this asexual lineage. In addition, I present the first documentation of accelerated mutation accumulation in the nuclear genome of a naturally-occurring, meiotically- reproducing organism. If harmful, these mutations could impede asexual lineage persistence and contribute strong support for the long-term benefits of sex.

No description available.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第20956号 / 理博第4408号 / 新制||理||1633(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)准教授 渡辺 勝敏, 教授 曽田 貞滋, 教授 中川 尚史 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

Esta tese aborda diversos temas em evolução molecular, usando extensivamente o formalismo de funções geratrizes para obter resultados analíticos sempre que possível. Em primeiro lugar, apresenta-se a solução exata para o comportamento dinâmico de uma população infinita de seqüências infinitamente longas (não há mutações reversas) evoluindo sob a ação de mutações deletérias em um relevo adaptativo multiplicativo ou truncado. Além disso, foi estudado o comportamento de uma população submetida a sucessivas diluições de intensidades arbitrárias, como ocorre em alguns protocolos de evolução experimental. Foram obtidas expressões matemáticas que, em princípio, podem ser úteis na caracterização de populações reais de microorganismos. Demonstrou-se também que um processo estocástico de ramificação multidimensional generalizado é uma excelente ferramenta para analisar numericamente os efeitos da degeneração mutacional (especificamente, de um fenômeno denominado catraca de Muller) em populações sob variadas condições de crescimento exponencial. Finalmente, simulações foram extensivamente utilizadas para analisar a história evolutiva de populações finitas e averiguar a possibilidade de certas grandezas, como certas medidas da topologia de árvores genealógicas, serem empregadas na elaboração de testes estatísticos capazes de detectar as marcas deixadas pela seleção natural. / This thesis discusses some topics of molecular evolution, extensively using generating function methods to find analytical results whenever possible. In first place, it gives the exact solution for the dynamics of an infinite population of infinitely long sequences (no back mutations) evolving under the action of deleterious mutations on either multiplicative or truncated fitness landscapes. In addition, the behavior of a population subject to successive dilutions of arbitrary intensity, just like some experimental evolution protocols, is found. The mathematical expressions, in principle, may prove useful in characterizing real populations of microor¬ganisms. It was also demonstrated that a generalized multidimensional branching process is a nice tool in numerically studying mutational degeneration effects (specifically a pheno¬menon called Muller\'s ratchet) in populations under a wide variety of exponential growth settings. Finally, the evolutionary history of finite populations was studied by simulations to probe the viability of certain statistic, like some topological measures in genealogical trees, being incorporated in statistical tests to detect the fingerprints of natural selection.

Most clonal plants exhibit a modular structure at multiple levels. At the level of the organs, they are characterized by functional modules, such as, internodes, leaves, branches. At the level of the genetic individual (clone or genet), they possess independent evolutionary and physiological units (ramets). These evolutionary units arise through the widespread phenomenon of clonal reproduction, achieved in a variety of ways including rhizomes, stolons, bulbils, or lateral roots. The focus of this study was Populus tremuloides, trembling aspen, a dioecious tree that reproduces sexually by seed and asexually through lateral roots. Local forest patches in western populations of Populus tremuloides consisted largely of multiple genotypes. Multi-clonal patches were dominated by a single genotype, and in one population (Riske Creek) we found several patches (five out of 17) consisting of a single genotype. A second consequence of modularity is that during the repeated cycle of ramet birth, development and death, somatic mutations have the opportunity to occur. Eventually, the clone becomes a mosaic of mutant and non-mutant cell lineages. We found that neutral somatic mutations accumulated across 14 microsatellite loci at a rate of between 10^-6 and 10^-5 per locus per year. We suggest that neutral genetic divergence, under a star phylogeny model of clonal growth, is an alternative way to estimate clone age. Previous estimates of clone age couple the mean growth rate per year of shoots with the area covered by the clone. This assumes a positive linear relationship between clone age and clone size. We found, however, no repeatable pattern across our populations in terms of the relationship of either shape or size to the number of somatic changes. A final consequence of modularity is that during clonal growth, natural selection is relaxed for traits involving sexual function. This means that mutations deleterious to sexual function can accumulate, reducing the overall sexual fitness of a clone. We coupled neutral genetic divergence within clones with pollen fitness data to infer the rate and effect of mildly deleterious mutations. Mutations reduced relative sexual fitness in clonal aspen populations by about 0.12x10^-3 to 1.01x10^-3 per year. Furthermore, the decline in sexual function with clone age is evidence that clonal organisms are vulnerable to the effects of senescence.

Most clonal plants exhibit a modular structure at multiple levels. At the level of the organs, they are characterized by functional modules, such as, internodes, leaves, branches. At the level of the genetic individual (clone or genet), they possess independent evolutionary and physiological units (ramets). These evolutionary units arise through the widespread phenomenon of clonal reproduction, achieved in a variety of ways including rhizomes, stolons, bulbils, or lateral roots. The focus of this study was Populus tremuloides, trembling aspen, a dioecious tree that reproduces sexually by seed and asexually through lateral roots. Local forest patches in western populations of Populus tremuloides consisted largely of multiple genotypes. Multi-clonal patches were dominated by a single genotype, and in one population (Riske Creek) we found several patches (five out of 17) consisting of a single genotype. A second consequence of modularity is that during the repeated cycle of ramet birth, development and death, somatic mutations have the opportunity to occur. Eventually, the clone becomes a mosaic of mutant and non-mutant cell lineages. We found that neutral somatic mutations accumulated across 14 microsatellite loci at a rate of between 10^-6 and 10^-5 per locus per year. We suggest that neutral genetic divergence, under a star phylogeny model of clonal growth, is an alternative way to estimate clone age. Previous estimates of clone age couple the mean growth rate per year of shoots with the area covered by the clone. This assumes a positive linear relationship between clone age and clone size. We found, however, no repeatable pattern across our populations in terms of the relationship of either shape or size to the number of somatic changes. A final consequence of modularity is that during clonal growth, natural selection is relaxed for traits involving sexual function. This means that mutations deleterious to sexual function can accumulate, reducing the overall sexual fitness of a clone. We coupled neutral genetic divergence within clones with pollen fitness data to infer the rate and effect of mildly deleterious mutations. Mutations reduced relative sexual fitness in clonal aspen populations by about 0.12x10^-3 to 1.01x10^-3 per year. Furthermore, the decline in sexual function with clone age is evidence that clonal organisms are vulnerable to the effects of senescence.

No description available.

Esta tese aborda diversos temas em evolução molecular, usando extensivamente o formalismo de funções geratrizes para obter resultados analíticos sempre que possível. Em primeiro lugar, apresenta-se a solução exata para o comportamento dinâmico de uma população infinita de seqüências infinitamente longas (não há mutações reversas) evoluindo sob a ação de mutações deletérias em um relevo adaptativo multiplicativo ou truncado. Além disso, foi estudado o comportamento de uma população submetida a sucessivas diluições de intensidades arbitrárias, como ocorre em alguns protocolos de evolução experimental. Foram obtidas expressões matemáticas que, em princípio, podem ser úteis na caracterização de populações reais de microorganismos. Demonstrou-se também que um processo estocástico de ramificação multidimensional generalizado é uma excelente ferramenta para analisar numericamente os efeitos da degeneração mutacional (especificamente, de um fenômeno denominado catraca de Muller) em populações sob variadas condições de crescimento exponencial. Finalmente, simulações foram extensivamente utilizadas para analisar a história evolutiva de populações finitas e averiguar a possibilidade de certas grandezas, como certas medidas da topologia de árvores genealógicas, serem empregadas na elaboração de testes estatísticos capazes de detectar as marcas deixadas pela seleção natural. / This thesis discusses some topics of molecular evolution, extensively using generating function methods to find analytical results whenever possible. In first place, it gives the exact solution for the dynamics of an infinite population of infinitely long sequences (no back mutations) evolving under the action of deleterious mutations on either multiplicative or truncated fitness landscapes. In addition, the behavior of a population subject to successive dilutions of arbitrary intensity, just like some experimental evolution protocols, is found. The mathematical expressions, in principle, may prove useful in characterizing real populations of microor¬ganisms. It was also demonstrated that a generalized multidimensional branching process is a nice tool in numerically studying mutational degeneration effects (specifically a pheno¬menon called Muller\'s ratchet) in populations under a wide variety of exponential growth settings. Finally, the evolutionary history of finite populations was studied by simulations to probe the viability of certain statistic, like some topological measures in genealogical trees, being incorporated in statistical tests to detect the fingerprints of natural selection.

Most clonal plants exhibit a modular structure at multiple levels. At the level of the organs, they are characterized by functional modules, such as, internodes, leaves, branches. At the level of the genetic individual (clone or genet), they possess independent evolutionary and physiological units (ramets). These evolutionary units arise through the widespread phenomenon of clonal reproduction, achieved in a variety of ways including rhizomes, stolons, bulbils, or lateral roots. The focus of this study was Populus tremuloides, trembling aspen, a dioecious tree that reproduces sexually by seed and asexually through lateral roots. Local forest patches in western populations of Populus tremuloides consisted largely of multiple genotypes. Multi-clonal patches were dominated by a single genotype, and in one population (Riske Creek) we found several patches (five out of 17) consisting of a single genotype. A second consequence of modularity is that during the repeated cycle of ramet birth, development and death, somatic mutations have the opportunity to occur. Eventually, the clone becomes a mosaic of mutant and non-mutant cell lineages. We found that neutral somatic mutations accumulated across 14 microsatellite loci at a rate of between 10^-6 and 10^-5 per locus per year. We suggest that neutral genetic divergence, under a star phylogeny model of clonal growth, is an alternative way to estimate clone age. Previous estimates of clone age couple the mean growth rate per year of shoots with the area covered by the clone. This assumes a positive linear relationship between clone age and clone size. We found, however, no repeatable pattern across our populations in terms of the relationship of either shape or size to the number of somatic changes. A final consequence of modularity is that during clonal growth, natural selection is relaxed for traits involving sexual function. This means that mutations deleterious to sexual function can accumulate, reducing the overall sexual fitness of a clone. We coupled neutral genetic divergence within clones with pollen fitness data to infer the rate and effect of mildly deleterious mutations. Mutations reduced relative sexual fitness in clonal aspen populations by about 0.12x10^-3 to 1.01x10^-3 per year. Furthermore, the decline in sexual function with clone age is evidence that clonal organisms are vulnerable to the effects of senescence. / Medicine, Faculty of / Medical Genetics, Department of / Graduate

No description available.