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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.
To assess the effect of organic compounds on the aquatic environment, organisms are typically exposed to toxicant solutions and the adverse effects observed are linked to the concentration in the surrounding media. As compounds generally need to be taken up into the organism and distributed to the respective target sites for the induction of effects, the internal exposure is postulated to best represent the observed effects.
The aim of this work is to contribute to an improved effect assessment of organic compounds by describing experimental and modelling methods to obtain information on the internal exposure of contaminants in organisms.
Chapter 2 details a protocol for the determination of bioconcentration parameter for uptake (k1) and elimination (k2) of organic compounds in zebrafish (Danio rerio) eggs. This enables the simulation of the internal exposure in zebrafish eggs from an ambient exposure concentration over time. The accumulated contaminant amount in zebrafish eggs was also determined, using a biomimetic extraction method. Different bioconc-entration estimation models for the determination of internal steady-state concentrat-ion of pharmaceutical compounds in fish to an environmental exposure are presented in Chapter 3. Bioconcentration factors were estimated from the compounds octanol: water partition coefficient (KOW) to determine the internal exposure to an ambient concentration.
To assess the integral bioavailable fraction from the water and sediment phase of environmental contaminants for rooted aquatic plants, the internal exposure in river-living Myriophyllum aquaticum plants were determined over time, presented in Chapter 4. The plants were collected at different time points, with the accumulated organic contaminants determined using a liquid extraction method.
In Chapter 5 a protocol was established to enable the non-invasive observation of effects in M. aquaticum plants exposed to contaminated sediments over time. Since the toxicant effects are a result of all uptake and distribution processes to the target site and the toxico-dynamic process leading to an observed effect during static exposure, information on the internal exposure could thus be gained from the temporal effect expression.rn
Wild boars belong to the most wide spread ungulates in the world. They are characterized by a well performed adaption to their environment mainly due to their omnivorous dietary. The wild boar population in Germany increased during the past three decades. Nowadays their high density leads to problems in agricultural areas due to damage of crops and plays a significant role as disease vector as the classical swine fever. For an effective population management population size information is of crucial importance. Different traditional methods exist to estimate population sizes as direct sightnings, faecal drop counts or hunting harvest which provide only relative estimates and population trends. Absolute population sizes could be yielded by a Capture-Mark-Recapture (CMR) approach. However, capturing of wild boars is difficult to realize and costly in terms of personnel and field effort.
Furthermore the capture probabilities are heterogeneous due to the variable behaviour of individuals influenced by age, sex, and experience of the animals. Non-invasive genetic methods are a promising complement to the traditional methods for population size estimation particularly for wild boar. These methods reduce stress and capture bias and increase the number of re-captures. Faeces proved to be a suitable DNA source for wild boar genotyping, due to almost equal capture probability. However working with faeces implicates difficulties such as low DNA rnquality and quantity, genotyping errors as dropout and false alleles.
The main aim of the present study was to develop a reliable, cost-efficient, reproducible and practicable method for wild boar genotyping. This method should provide a reliable dataset of genotypes obtained from the collected faeces samples. Individual identification forms the basis for an improved mark-recapture approach. As there is no sound method for absolute population counts in free living wild boar, reference values for the validation of this new approach are missing. Therefore, different routines to reduce and to assess genotyping errors were compared within this thesis. For maximum amplification rate, the storage, the extraction methods and the PCR-procedure were optimised. A step by step procedure was evaluated in order to determine the minimum required microsatellite (MS) number for reliable individual identification including a test with family groups (female and embryo tissue) to distinguish even between close relatives. A multiple-tubes approach, post-amplification checking and different correction procedures were applied to reduce genotyping errors. In order to quantify real genotyping error rates (GER) of datasets derived from sampling in the Palatinate Forest in western Germany, different methods for GER determination were compared with each other, obtaining GERs between 0% and 57.5%. As a consequence, more strict criteria for the multi-tube approach and increased repetition number of homozygous samples were used. An additional method validation was the implementation of a blind test to achieve the reliability of the genotyping and error checking procedure. Finally a strict and practicable proposal for the lab procedure was developed, by beginning with faecal sample collection and ending with a reliable dataset with genotypes of each sample.
The results of the presented method were derived from two sampling periods in a 4000 ha area in the Palatinate Forest in Rhineland-Palatinate in December 2006 and 2007. Both provided high confidence intervals (CI) applying inaccurate estimates (eg. for 2006 population size amounted to 215 with CI 95% of 156-314 and for 2007 population size amounted to 415 with CI 95% of 318-561) due to low sampling sizes (for 2006 n = 141 and for 2007 n = 326), successfully analysed samples (for 2006 n = 89 and for 2007 n = 156) and recapture numbers (for 2006 n = 12 and for 2007 n = 24). Furthermore, the population estimates even for the lowest values were considerably higher than previously assumed by hunting statistics, which implicates an ineffective hunting regime in the study area. For the future prospect, to obtain more precise population size estimations the increase of sampling sizes is inevitable, because absolute and reliable estimates are highly desirable for wildlife management and the control of diseases transmission. Nevertheless, the method for individual genotyping of wild boars evaluated in this thesis could be successfully established resulting in reliable datasets for population estimation modelling with sufficiently low GER.
New media are continually gaining importance in society. This process also has an increasing influence on developments in the field of education. Due to the use of computers as an integral part of schooling, new possibilities with regard to the organisation of learning processes arise. In this context, it is of great significance that appropriate computer applications for the respective learning group be prepared, so that justifiable use of computers in lessons can take place. Furthermore, efficient integration of computers requires changes in spatial organisation, in teaching methodology and in the role of the teacher. Such reflection and re-orientation are the essential basis for meaningful usage of new media in teaching and learning processes. An initial aim of this thesis is an empirical analysis of the situation regarding the usage of computers in geometry lessons in primary schools, based on a regional survey. The evaluation gives information as to how intensively the computer is used in the learning process and shows us which factors determine the use of computers in geometry lessons.
The results are an empirical foundation for the development of a computer-based learning environment called "Geolizi" (the second aim of my study). Within this learning environment, the pupils should work independently on the topics "mirror-imaged figures" and "the construction of rectangles and squares", with the help of the computer. During this process, hands-on media, traditional drawing instruments and interactive worksheets are available to the pupils. The computer (with its appropriate applications) takes over different functions in this learning process. Testing of this learning environment ("Geolizi") took place in several primary school classes, within the scope of formative and summative evaluation. With the help of questionnaires filled in by the pupils, the usability of the individual elements was tested. Based on a pre-post-investigation design, an attempt has been made to discover possible changes in the attitude of teachers regarding the usage of computers in the teaching of elementary geometry.
The results of this test phase, together with the evaluation of the questionnaires, lead to the founded presumption that usage of the multimedia-based learning environment " Geolizi " could result in greater use of computers in geometry lessons. All in all, the developed learning environment demonstrates an interesting possibility of how to use computers in the teaching of geometry at primary schools, thus making an important contribution to an independent, individualised learning process.