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Population genetic structure in European Hyalodaphnia species: Monopolization versus gene flow
(2012)
Cyclic parthenogens displays an alternation of asexual and sexual reproduction which has consequences for the genetic structure of these organisms. The clonal diversity of cyclic parthenogenetic zooplankton populations is influenced by the size of the dormant egg bank, i.e., the amount of sexually produced dormant eggs that assembled in the sediment, as these dormant eggs contribute new genetic variants to the populations. Further, the clonal diversity is impacted by clonal erosion over time, which reduces the number of different clones through stochastic and selective processes. Although freshwater invertebrates are good dispersers through their dormant stages, the influence of gene flow is assumed to be negligible, as the local population successfully monopolizes the available resources. As these populations reach carrying capacity fast due to the asexual reproduction, the first colonizing individuals are able to successfully establish in the habitat, resulting in a priority effect which hinders the invasion of new genotypes. Due to clonal selection and sexual reproduction a population will locally adapt over time and will establish a dormant egg bank which facilitates the fast re-colonization after a hostile period. This thesis evaluates the processes altering the population genetic structure of cyclic parthenogenetic zooplankton with a special focus on the concepts of monopolization as well as the counteracting effects of gene flow, using large-lake Daphnia species. Thirty-two variable microsatellite DNA markers were developed and a subset of twelve markers was evaluated regarding their suitability for species assignment and hybrid class detection. With this marker set and an additional mitochondrial DNA marker forty-four natural European populations of the species D. cucullata, D. galeata and D. longispina were studied. In D. galeata, most populations were characterized by low clonal diversities which suggest high influence from clonal erosion over the growing season and a low contribution from the dormant egg bank. Further, recent expansions as well as gene flow were detected, probably caused by the anthropogenic alteration of freshwater habitats, in particular eutrophication of many European lakes. D. longispina and D. cucullata revealed a different genetic structure compared to D. galeata, with high genetic differentiation among populations. This indicates low levels of effective gene flow which is in line with the predictions of monopolization. Further, high clonal diversities were found in populations of the two taxa, suggesting a high contribution from the dormant egg bank while clonal erosion was often not detectable. In D. longispina, mitochondrial data revealed an ancient expansion which was probably initiated by the formation of glacial lakes after the last ice age.
In addition, in D. longispina not only clonal diversity but also genetic diversity was high, indicating that during the build-up of the studied populations the influence from gene flow was probably high. To better understand the processes that act on early populations the population build-up in regard to the temporal advantage of clones during invasion succession was experimentally studied and revealed that priority effects shape population structure of Daphnia species. However, in certain cases the highly superior clones resulted in the extinction of inferior clones independent of the temporal advantage the single clones had.
This clearly shows that not only the time of succession is important but also the competitive strength. rnIn conclusion, the results obtained show that the population genetic structure in cyclic parthenogenetic zooplankton species is impacted by various processes. In addition to earlier studies, which mainly focus on local adaptation, clonal erosion and the size of the dormant egg bank to understand population genetic structure, this thesis could show that gene flow may be effective as well. During population build-up the advantage of early arriving individuals does not necessarily predict the outcome of population assembly, as additional genotypes may contribute to the population. Finally, the genetic structure of established populations may be severely impacted by effective gene flow, if severe environmental changes alter the habitat of the locally adapted population.