Sexual reproduction is expected to facilitate the removal of deleterious mutations from populations because biparental inheritance (i.e., segregation) and recombination during meiosis break down linkage disequilibria (LD), allowing mutations to be selected independently from their genetic background. Accordingly, the absence of recombination and segregation is expected to increase selective interference between loci, translating into reduced efficacy of natural selection. While there now exist multiple lines of evidence demonstrating that asexual lineages do experience accelerated accumulation of putatively harmful mutations, whether these mutations influence phenotype in a manner that could contribute to the maintenance of sex remains almost entirely unevaluated. Here, I use the New Zealand freshwater snail, Potamopyrgus antipodarum, to address these questions. In particular, I take advantage of the fact that the mitochondrial genome is expected to suffer from these mutational effects and interacts extensively with the nuclear genome to evaluate potential harmful effects of mutation accumulation in asexuals on a genome-wide scale. I present evidence that harmful mutations remain extant longer in asexual populations than in sexual populations, that the degree of functional constraint determines the extent of mutation accumulation in asexuals, that there is genetic variation for mitochondrial function in asexual lineages of P. antipodarum, and that phenotypic variation for mitochondrial function is mediated by both genetic and environmental variation. Together, these analyses provide strong evidence that asexual lineages are accumulating deleterious mutations, and that there is genetic variation, structured by lake, for mitochondrial function.

Pagrindinis šio darbo tikslas buvo ištirti hidromorfologinių upės savybių ir invasinės rūšies, Dreissena polymorpha poveikį vandens moliuskų bendrijų struktūrai. Pasirinktos penkios tyrimų upės: Babrungas, Lakaja, Peršokšna, Verknė ir Žeimena. Taipogi atliktas laboratorinis eksperimentas, įvertinti dreisenos apaugimo poveikį Unio pictorum somatiniam augimui. Tyrimo metu, iš viso rasta 31 vandens moliuskų rūšis. Didžiausia rūšinė įvairovė nustatyta šaltavandenėjeVerknės upėje (22 rūšys), o mažiausia, 7 rūšys – šiltavandenėje Lakajos upėje. Pirmą kartą Lietuvos gėluosiuose vandenyse, Vilkokšnio ežere ir Verknės upėje aptikta Potamopyrgus antipodarum rūšis. Buvo tikrinta hipotezė, kad didžiausias dreisenų gausumas yra upės ištakose, o upės kontinuume jis mažėja. Ryškiausias dreisenų poveikis vietinėms vandens moliuskų bendrijoms nustatytas ties ištakomis, kur šios invazinės rūšies gausumas ir biomasė didžiausi. Šis poveikis gradientiškai mažėja nuo ištakų link žiočių. Be to, buvo tikrintas hidromorfologinių upės savybių, tokių kaip gruntas, srovės greitis, poveikis moliuskų gausumui, biomasei ir bioįvairovei. Nustatyta, kad substratas, srovės greitis ir pati upė turi reikšmingos įtakos moliuskų gausumui ir biomasei. Bioįvairovę reikšmingai įtakojo tik grunto tipas. Taip pat, tikrinta hipotezė, kad nedidelis dreisenų apaugimas neturi reikšmingo poveikio Unio pictorum individų somatiniam augimui. Nenustatyta reikšmingų skirtumų tarp eksperimentinių grupių sausosios biomasės... [toliau žr. visą tekstą] / The main objective of this work was to investigate the effect of hydromorphological features of the rivers and invasive species to the molluscs community structure. The selected rivers were: Babrungas, Lakaja, Peršokšna, Verknė and Žeimena. In these rivers mollusc abundance, biomass, biodiversity and dominance were examined. What is more, the laboratory experiment was made to check the influence of infestation of Dreissena polymorpha on Unio pictorum mussel somatic growth. Durind the study, 31 species of freshwater molluscs were found in 5 examined rivers. The bigest species richness was found in a coldwater river Verknė (22 species) and the lowest – in warmwater river Lakaja (7 species). First time in a freshwaters of Lithuania, invasive species Potamopyrgus antipodarum was found in a lake Vilkokšnis and Verknė river. The hypothesis, that dreissena is most abundant near the outfall and decreases in the continuum of the rivers was tested. The most significant effect of Dreissena polymorpha to the mollusc communities was found to be near the outfall of the rivers, where its abundance and biomass are biggest. This effect declines, because zebra mussel abundance and biomass decreases in the continuum of the river. What is more, the effect of hydromorphological features, such as substrate and current speed, of the rivers to mollusc abundance, biomass and biodiversity was studied. Substrate, current speed and river were found to have significant effect to the abundance and biomass... [to full text]

The trajectory of evolutionary adaptation can be influenced both by the interactions of organisms with their environments as well as by the biological characteristics of the organisms themselves. My dissertation research uses the New Zealand freshwater snail Potamopyrgus antipodarum to 1) gain important insight into how coevolutionary interactions between hosts and parasites influence patterns of gene expression and genetic differentiation of hosts and, 2) evaluate how reproductive mode, and ploidy level affect patterns of adaptive molecular evolution. Coevolutionary interactions between hosts and parasites are a primary source of strong natural selection that can lead to rapid evolutionary change. Here, I used evaluation of patterns of gene expression and genetic differentiation to take critical steps towards characterizing the genomic basis of coevolutionary interactions between P. antipodarum and Microphallus livelyi. I found that M. livelyi-infected P. antipodarum exhibit systematic downregulation of genes relative to uninfected P. antipodarum. The specific genes involved in response to parasites differ markedly across lakes, consistent with population-specific host-parasite interactions leading to population-specific evolutionary trajectories. I also identified a set of rapidly evolving loci that represent promising candidates for targets of parasite-mediated selection across lakes as well as within each lake population. These results constitute the first genomic evidence for population-specific responses to coevolving infection in the P. antipodarum-M. livelyi interaction and provide new insights into the genomic basis of coevolutionary interactions in nature. I also generated and characterized the first transcriptomic resources for Microphallus parasites collected from two species of Potamopyrgus snails (P. antipodarum and P. estuarinus). These data both revealed that these parasites appear to represent distinct genetic lineages, which is interesting in light of the tight coevolutionary interactions between P. antipodarum and M. livelyi, and lay the groundwork for future research. Polyploidy has the potential to facilitate adaptive evolution by providing redundant genome copies that are free to evolve new functions. By contrast, asexuality, with which polyploidy is often associated, is expected to restrict adaptive evolution by decreasing the efficacy of natural selection and access to new genetic variation. I evaluated whether and how ploidy level and reproductive mode influence patterns of adaptive molecular evolution in P. antipodarum to assess 1) the potential evolutionary genomic benefits of recent polyploidy, and 2) how patterns of adaptive molecular evolution in asexuals are influenced by polyploidy. I compared patterns of positive selection in 60 genes across 27 P. antipodarum lineages (10 diploid sexuals, 12 triploid asexuals, 5 tetraploid asexuals) and a diploid sexual outgroup, Potamopyrgus estuarinus. I found little evidence that ploidy level and/or reproductive mode influence patterns of positive selection in P. antipodarum. Even so, this study provides initial steps in evaluating whether ploidy level and reproductive mode influence patterns of adaptive molecular evolution. Taken together, my dissertation work contributes new insights to the field of host-parasite coevolutionary interactions and will inform future studies into how ploidy level and reproductive mode influence patterns of adaptive molecular evolution.

Although genomes are perhaps the single most important element of living systems, why they feature such striking variation and how this variation is maintained within and across natural populations remains unclear. One of the most common and important means by which genomic variation is generated is ploidy elevation. While polyploidy has been implicated in the remarkably successful radiations of angiosperms, teleost fish, and amphibians, the phenotypic consequences of changes in ploidy level are poorly understood, especially in animals. I use a large, multi-year common garden experiment to identify potential life history costs and benefits of polyploidy and asexual reproduction, a trait often associated with polyploidy, in Potamopyrgus antipodarum. This snail is well suited for studying ploidy variation and sex because diploid sexuals and triploid and tetraploid asexuals frequently coexist, allowing us to use comparisons of sexuals to asexuals and triploid to tetraploid asexuals to study both the effects of ploidy elevation and sex. I detected a strong negative correlation between growth rate and time to maturity and found that sexual P. antipodarum grew and matured significantly more slowly than the polyploid asexuals. Sexual P. antipodarum were also more likely to die before achieving reproductive maturity than their asexual counterparts. By contrast, there were no apparent life history differences between triploid and tetraploid asexuals, indicating that direct phenotypic benefits of ploidy elevation are unlikely to explain the relatively rapid growth and maturation of asexuals. My results suggest that ploidy elevation does not inevitably confer phenotypic consequences, that reproductive mode influences life history trait expression, and that sexual P. antipodarum persist in many natural populations in spite of substantial life history disadvantages.