Thesis (M.S.)--Missouri State University, 2009. / "May 2009." Includes bibliographical references (leaves 27-34). Also available online.

Phenotypic plasticity is an environmentally based change in phenotype and can be adaptive. Often, the change in an organism's phenotype is induced by the presence of a predator and serves as a defense against that predator. Defensive phenotypes are induced in freshwater physid snails in response to both crayfish and molluscivorous fish. Alternative morphologies are produced depending on which of these two predators snails are raised with, thus protecting them from each of these predators' unique mode of predation. Snails and other mollusks have been shown to produce thicker, differently shaped shells when found with predators relative to those found without predators. This production of thicker, differently shaped shells offers better protection against predators because of increased predator resistance. The first study in this thesis explores costs and limits to plasticity using the snailfish- crayfish system. I exposed juvenile physid snails (using a family structure) to either early or late shifts in predation regimes to assess whether developmental flexibility is equally possible early and late in development. Physid snails were observed to produce alternative defensive morphologies when raised in the presence of each of the two predators. All families responded similarly to the environment in which they were raised. Morphology was found to be heritable, but plasticity itself was not heritable. Morphology was found to become less flexible as snails progressed along their respective developmental pathways. In the second study, I raised physid snails with and without shell-crushing sunfish and examined the differences in shell thickness, shell mass, shell size and shell microstructural properties between the two treatment groups. Shells of snails raised with predators were found to be larger, thicker and more massive than those raised without predators, but differences in microstructure were found to be insignificant. I conclude that the observed shell thickening is accomplished by the snails' depositing more of the same material into their shells and not by producing a more complex shell composition.

La théorie de la sélection sexuelle a été largement élaborée à partir du constat de dimorphisme sexuel chez les espèces à sexes séparés. Une de ses caractéristiques générales est une sélection plus forte pour l'augmentation du nombre de partenaires sexuels chez les mâles que chez les femelles qui résulterait d'un investissement différentiel dans les descendants entre les deux sexes (anisogamie). Si hermaphrodisme et sélection sexuelle sont considérés comme compatibles depuis les travaux de Charnov (1979), les études sur le sujet restent rares que ce soit chez des animaux ou des plantes. Une raison importante est que la méthodologie disponible pour quantifier la sélection sexuelle ne prend pas en compte les particularités des hermaphrodites (par ex., corrélations ou effets croisés entre les deux fonctions sexuelles d'un même individu, autofécondation). Le premier objectif de cette thèse est de combler cette lacune méthodologique en proposant un cadre travail adapté aux hermaphrodites, que nous appliquons dans une étude empirique chez Physa acuta, un gastéropode hermaphrodite d'eau douce. Nous observons que la sélection sexuelle est plus intense sur la fonction mâle, comme généralement chez les espèces gonochoriques. Par ailleurs, nous ne détectons aucun effet des particularités des hermaphrodites dans cette expérience. Dans un deuxième temps, nous nous intéressons de manière plus détaillée aux composantes du succès reproducteur mâle (RSm). Nous montrons que chez P. acuta il existe une priorité spermatique au premier partenaire mâle lorsque plusieurs individus sont en compétition. Enfin, nous proposons une décomposition de la variance de RSm en ses composantes pré- et post-copulatoires, qui représentent respectivement 60 et 40% de la variance. Dans la troisième partie, nous intégrons la sélection sexuelle à l'étude de l'évolution de l'allocation sexuelle d'un hermaphrodite, via une approche d'évolution expérimentale chez P. acuta. Menée sur plus de 10 générations, elle vise à faire évoluer l'allocation sexuelle de manière disruptive (lignées mâle ou femelle) en sélectionnant les composantes mâle et femelle du succès reproducteur. Les résultats préliminaires suggèrent qu'il est possible de manipuler l'allocation sexuelle chez un hermaphrodite simultané en sélectionnant sur son régime d'appariement. Nous concluons que l'anisogamie suffit à justifier l'existence de la sélection sexuelle sans avoir à supposer un dimorphisme sexuel. Son étude chez les hermaphrodites simultanés ouvre des perspectives pour la compréhension du rôle de l'allocation sexuelle dans l'évolution des systèmes de reproduction. / A cornerstone of the theory of sexual selection in gonochoric species is sexual dimorphism. A very general result is stronger selection on males than on females for increasing mating success, and this fundamentally relies on differential investment in offspring between the sexes (anisogamy). Although sexual selection does operate in hermaphroditic species as well, few empirical studies have been performed whether in animals or in plants. The main reason is that the current framework for studying sexual selection does not incorporate the particularities of hermaphrodites, including correlations or cross-sex effects between sex functions and self-fertilization. The first goal of this thesis is to fill this gap by proposing an appropriate framework for hermaphrodites (generalizing that available for gonochoric species). It was applied to approach sexual selection in the hermaphroditic freshwater gastropod Physa acuta. Sexual selection turns out to be stronger on the male than on the female function, as classically observed in gonochorists. Moreover, we do not detect any effect in relation to hermaphrodites' particularities. We then focus on the components of male reproductive success (RSm) in more details. We detect a pattern of sperm precedence in conditions of sperm competition. We develop a new method for decomposing the variance in RSm into pre- and post-copulatory components (representing 60 and 40% of the variance respectively). The third section aims at integrating sexual selection in studies of sex allocation and its evolution. It relies on a protocol of experimental evolution in P. acuta. Conducted over more than 10 generations. Its aim is to observe the evolution of sex allocation by disruptively selecting male and female components of reproductive success. Preliminary results indeed suggest that it is possible to manipulate sex allocation in a simultaneous hermaphrodite by manipulating its mating system. We conclude that anisogamy alone is a sufficient condition for sexual selection to proceed, and that sexual dimorphism is not required. Study sexual selection in simultaneous hermaphrodites gives insights for understanding the role of sex allocation in the evolution of mating systems.

Freshwater ecosystems will likely become sinks for future silver loadings as a result of increased nanosilver (n-Ag) use in industrial and commercial applications. A series of bioassays was performed to assess how n-Ag toxicity may be influenced by abiotic factors associated with natural freshwater ecosystems. Additionally, these bioassays provide insight into how environmentally relevant concentrations of n-Ag may sublethaly affect the freshwater benthic gastropod, Physa acuta, that plays pivotal roles in maintaining the structure and function of freshwater ecosystems. In sediment with no benthic organic carbon (BOC), gastropod vital rates decreased in treatments containing any n-Ag, gastropods in sediment with relatively low BOC appeared to trade off growth for reproduction at high n-Ag treatments, while gastropod vital rates in high BOC sediment remained unaffected at all nanosilver treatments. Sediment type may abate nanosilver toxicity as a result of organic carbon content. Effects of n-Ag on gastropod vital rates were not dependant on pH, suggesting aqueous pH does not directly influence n-Ag toxicity. Nanosilver (0.2 μg/L) stressed gastropods, altering their growth and reproduction tradeoff dynamics. Nanosilver concentrations modeled to exist in natural freshwaters, disrupted gastropod ability to detect and respond to a natural predator, while greater n-Ag concentrations stimulated gastropods to exhibit contaminant avoidance behavior and thereby attempted to flee their habitat. This study provides direction in understanding how adverse n-Ag effects may be influenced by abiotic parameters, while assessing sublethal effects of n-Ag on freshwater gastropods that are likely to occur in natural freshwater ecosystems, given current estimates of environmental n-Ag concentrations. / Department of Biology