<p>This thesis focuses on reproduction in a simultaneous hermaphrodite with internal fertilization; the nudibranch <i>Aeolidiella glauca</i>. Unlike most other nudibranchs, where copulation is the rule, <i>A</i>. <i>glauca</i> was found to transfer sperm via external spermatophores that were attached to the partner's back. Despite elaborate courtship the actual spermatophore transfer, which always involved two animals only, was of short duration. In most matings (88%) spermatophores were reciprocally exchanged. </p><p><i>A. glauca</i> was further found to be very promiscuous. During mating and sperm transfer the receiver exerts considerable control over sperm, and manipulative behaviours designed to increase the donor's reproductive success are thus likely to have evolved. An example of such manipulative behaviour may be <i>A. glauca's</i><b> </b>unique spermatophore avoidance behaviour. I found that slugs carrying a sign of previous mating activity, i.e. a spermatophore, were discriminated against in a situation where mate choice was possible. The presence of spermatophores was further found to reduce slugs' ability to interrupt matings, and displace other slugs. Body size, however, had no direct effect on displacement in <i>A. glauca</i> as small slugs interrupted matings as successfully as large ones. Furthermore, pair formation and mating were found to be random with respect to size. This was true also for pairs formed in the field. </p><p>In addition to mate choice hermaphrodites may increase their reproductive success by differential sex allocation. I tested whether differing mate encounter rates had any effects on allocation to male and female function in <i>A. glauca</i>. Slugs with more mating opportunities mated more, and had higher proportional spermatophore production that others. As predicted they also laid significantly fewer eggs than slugs presented with partners less often.</p>

Understanding the processes that govern mating behaviors is a fundamental goal of evolutionary biology and behavioral ecology. Population-level patterns of mate acquisition and offspring production, otherwise known as the genetic mating system, play a central role in the sexual selection on morphological and behavioral traits and may facilitate speciation. The central hypothesis of this research is that variation in environmental conditions, such as temperature, turbidity, and habitat, and demographic influences such as population density, sex ratios and temporal availability of mates, may limit mating and reproductive success in a predictive manner. Therefore the goal of this dissertation is to examine the contributions of geographic and temporal variation on the plasticity of the genetic mating system in two species of pipefish. The first study examined whether meaningful variation in the genetic mating system exists between two natural populations of the dusky pipefish, Syngnathus floridae. Results of this investigation provide evidence that the genetic mating system differs among different geographic locations. The second study considered the relative contributions of environmental conditions and population demographics on differences in the genetic mating system of dusky pipefish from five natural populations. The results of this investigation show strong trends for demographic and environmental factors to strongly influence the genetic mating system between populations. The third study considered how variation in the number of available mates predicts the outcome of sexual selection during the course of a breeding season in the broad-nosed pipefish, Sygnathus typhle. The results of this study indicate a strong influence of the operational sex ratio on the genetic mating system. In addition to these studies, a study was conducted to investigate whether phylogeographic relationships may be responsible for geographic variation in the genetic mating system of the dusky pipefish of pipefish. Mitochondrial DNA analysis does not substantiate subspecies designations for this species and microsatellite analysis show a clear pattern of isolation by distance. Taken together, these studies significantly enhance the understanding of how mating systems are organized over broad environmental gradients and temporal/spatial scales and to the evolution of sexual selection on the whole.

Ce travail s’est intéressé aux liens existant entre la stratégie d’exploitation développée par un crustacé ectoparasite et son comportement reproductif. Le crabe Pinnotheridae Dissodactylus primitivus exploite deux espèces Spatangidae vivant dans la Mer des Caraïbes, Meoma ventricosa et Plagiobrissus grandis. Des approches comportementales, démographiques et génétiques ont été adoptées afin de mettre en lumière le fonctionnement et la biologie de cette symbiose. Par son comportement alimentaire, le crabe occasionne des lésions tégumentaires sur ses hôtes. Celles-ci affectent la fitness de M. ventricosa, au travers de son développement gonadique. Dissodactylus primitivus exploite ses deux espèces hôtes de façon asymétrique. La reproduction des parasites se déroule sur les deux hôtes, alors que le recrutement ne s’effectue que sur M. ventricosa. Ce cycle vital asymétrique du crabe serait stabilisé par la qualité et la rareté de P. grandis. En outre, Le comportement sexuel du crabe sur M. ventricosa répondrait aux critères de la polygynandrie à femelles mobiles. Selon ce modèle, les mâles et les femelles se déplacent entre les hôtes à la recherche de partenaires multiples. Lors de ces déplacements, le crabe s’aiderait de son aptitude à localiser chimiquement ses hôtes. Néanmoins, ce mécanisme s’avère plastique et pourrait refléter l’asymétrie du cycle vital. En effet, cette différence n’a pas d’origine génétique, car les crabes vivant au sein du site d’étude constituent la même population quelle que soit l’espèce hôte considérée. Les marqueurs moléculaires microsatellites mis au point dans ce travail permettront lors de futurs travaux d’affiner les observations sur les modalités d’accouplement du crabe et d’estimer sa capacité de dispersion. This work aimed to highlight the relationships between the host exploitation strategy of an ectoparasite crustacean and its mating system. The pea crab Dissodactylus primitivus exploits two Spatangidae species living in the Caribbean Sea, Meoma ventricosa and Plagiobrissus grandis. Behavioural, demographic and genetic approaches have been conducted to examine the functioning and biology of this symbiosis. Owing to its feeding behaviour, the crab wounds the host tegument. The wounds negatively affect M. ventricosa's fitness through its gonadic development. Dissodactylus primitivus asymmetrically exploits its two host species. The reproduction of the parasites happens on each host, but the recruitment only takes place on M. ventricosa. The asymmetrical life cycle would be stabilised par the quality and the scarcity of P. grandis. The mating system of crabs living on M. ventricosa would correspond to the Pure-search polygynandry of mobile females criteria. According to this model, the males and the females practice the host switching behaviour to find several sexual partners. During these movements, the crab could use its chemodetection ability to locate its hosts. However, this mechanism is plastic and presumably reflects the asymmetrical life cycle of the crab. This difference has indeed not a genetic cause because the crabs living inside the investigated region belong to the same population, whatever the regarded host species. In future studies, the microsatellites markers developed for this work could be used to test the mating system of D. primitivus and to estimate its dispersion ability.

Background: Multidirectional interactions in social networks can have a profound effect on mate choice behavior; e.g., Poecilia mexicana males show weaker expression of mating preferences when being observed by a rival. This may be an adaptation to reduce sperm competition risk, which arises because commonly preferred female phenotypes will receive attention also from surrounding males, and/or because other males can copy the focal male's mate choice. Do P. mexicana males indeed respond to perceived sperm competition risk? We gave males a choice between two females and repeated the tests under one of the following conditions: (1) an empty transparent cylinder was presented (control); (2) another ("audience") male inside the cylinder observed the focal male throughout the 2nd part, or (3) the audience male was presented only before the tests, but could not eavesdrop during the actual choice tests (non-specific sperm competition risk treatments); (4) the focal male could see a rival male interact sexually with the previously preferred, or (5) with the non-preferred female before the 2nd part of the tests (specific sperm competition risk treatments). Results: The strength of individual male preferences declined slightly also during the control treatment (1). However, this decrease was more than two-fold stronger in audience treatment (2), i.e., with non-specific sperm competition risk including the possibility for visual eavesdropping by the audience male. No audience effect was found in treatments (3) and (5), but a weak effect was also observed when the focal male had seen the previously preferred female sexually interact with a rival male (treatment 4; specific sperm competition risk). Conclusion: When comparing the two 'non-specific sperm competition risk' treatments, a very strong effect was found only when the audience male could actually observe the focal male during mate choice [treatment (2)]. This suggests that focal males indeed attempt to conceal their mating preferences so as to prevent surrounding males from copying their mate choice. When there is no potential for eavesdropping [treatment (3)], non-specific specific sperm competition risk seems to play a minor or no role. Our results also show that P. mexicana males tend to share their mating effort more equally among females when the resource value of their previously preferred mate decreases after mating with a rival male (perceived specific sperm competition risk), but this effect is comparatively weak.

The aim of this thesis was to study the evolution of reproductive systems and reproductive traits in the fungal genus Neurospora. More specifically, I have investigated the evolutionary forces shaping the genes involved in sexual reproduction, focusing on mating-type (mat) and pheromone receptor (pre) genes. Neurospora contains species exhibiting three different mating systems, i.e., heterothallism (self-incompatibility), homothallism (self-compatibility) and pseudohomothallism (partial self-incompatibility). First, a robust phylogeny of Neurospora was established. The phylogenetic analyses revealed multiple independent transitions in reproductive life style during the evolutionary history of the genus. We argued for a heterothallic ancestor of the genus, although our subsequent ancestral reconstruction analyses favored a homothallic ancestor. To be able to settle the ancestral mating system, we zoomed in on the structural architecture of the mat-locus in four homothallic species of Neurospora, thought to have arisen from independent transitions. Our results led us to suggest two different genetic mechanisms (translocation and unequal crossover) to explain the transitions in mating system from heterothallism to homothallism. We pointed out that the mating-system transitions in Neurospora are unidirectional, and suggested that transposable elements might be driving the transitions. In conclusion, we suggest a heterothallic ancestor for Neurospora, and that at least six transitions to homothallism and two transitions to pseudohomothallism have occurred in its evolutionary history. Further, we used the phylogeny of Neurospora as a framework to test if the evolution of pre-genes (pre-1 and pre-2) in hetero- and homothallic Neurospora is dependent on mating systems and/or even the homothallic clades themselves (i.e., mating-system and/or switch-dependent). The molecular evolution results suggest that pre-1 and pre-2 are overall functional in both homothallic and heterothallic Neurospora. The molecular evolution of pre-1 seems to be independent of mating-system or homothallic clade, and we detected signs for positive selection in the C-terminal tail. For pre-2 we found no support for mating-system dependent evolution, but indications for switch-dependent evolution. In this study we also included expression analyses of both pre- as well as mat-genes, with the prospect to assess functionality and regulation. During this thesis work, we also performed a phylogenetic study were we found that reproductive genes might be more permeable to introgression than other genes, which is in contrast to theoretical expectations. In the last study, we confirmed the co-existence of two alternative splice variants of the pheromone receptor gene pre-1 in Neurospora crassa, and performed expression profiles studies using quantitative RT-PCR. I hope this thesis work will further strengthen Neurospora as a model for research in evolutionary genetics.

The thesis consists of a summary and four self-contained papers. Paper [I] examines the effects of interregional migration on gross earnings in married and cohabiting couples. In particular, we examine the link between education level and income gains. We find that pre-migration education level is a key determinant of migration and economic outcomes and is also a determinant of the effect of migration on income distribution within the household. The positive average effect on household earnings is largely explained by income gains among highly-educated males. Females generally experience no significant income gain from migration in absolute terms. Paper [II] analyzes the effect of the spouse’s education on individual earnings. In this study, we control for time-invariant heterogeneity that may be correlated with the spouse’s education level and use a rich data set that includes observations of individuals when they are single and when they are married. The results support the hypothesis of cross-productivity for both males and females. Furthermore, couples with education within the same field experience even larger effects. In Paper [III] we aim to study how the spouse’s productivity in the labor market affects one’s own individual earnings when married. Using longitudinal data on individuals as both single and married allows us to estimate the spouses’ productivity as single persons and thereby avoid problems of endogeneity between the two spouses’ labor market performances. Productivity is approximated with residuals from estimates of pre-marriage earnings equations. Results indicate that there are negative effects of the spouse’s productivity on individual earnings for both males and females, and that this effect appears to be enhanced by the duration of the marriage. Paper [IV] studies spousal matching on earnings for females in secondorder marriages. We aim to follow women who marry, divorce, and subsequently remarry compared with females who marry and stay married over the course of the study interval. Overall, we find significant positive correlations for all three of the marital partitions. The correlation tends to be smaller for the first of a sequence of marriages for women who divorce than for women who marry and stay so. For the second of the successive marriages, however, the correlation of the residuals is larger than that for women who marry but once.

This thesis focuses on reproduction in a simultaneous hermaphrodite with internal fertilization; the nudibranch Aeolidiella glauca. Unlike most other nudibranchs, where copulation is the rule, A. glauca was found to transfer sperm via external spermatophores that were attached to the partner's back. Despite elaborate courtship the actual spermatophore transfer, which always involved two animals only, was of short duration. In most matings (88%) spermatophores were reciprocally exchanged. A. glauca was further found to be very promiscuous. During mating and sperm transfer the receiver exerts considerable control over sperm, and manipulative behaviours designed to increase the donor's reproductive success are thus likely to have evolved. An example of such manipulative behaviour may be A. glauca's<b> </b>unique spermatophore avoidance behaviour. I found that slugs carrying a sign of previous mating activity, i.e. a spermatophore, were discriminated against in a situation where mate choice was possible. The presence of spermatophores was further found to reduce slugs' ability to interrupt matings, and displace other slugs. Body size, however, had no direct effect on displacement in A. glauca as small slugs interrupted matings as successfully as large ones. Furthermore, pair formation and mating were found to be random with respect to size. This was true also for pairs formed in the field. In addition to mate choice hermaphrodites may increase their reproductive success by differential sex allocation. I tested whether differing mate encounter rates had any effects on allocation to male and female function in A. glauca. Slugs with more mating opportunities mated more, and had higher proportional spermatophore production that others. As predicted they also laid significantly fewer eggs than slugs presented with partners less often.

To explore the similarities and differences of regulatory circuits among budding yeasts, we characterized the role of unscheduled meiotic gene expression 6 (UME6) and a novel mating type switching pathway in Kluyveromyces lactis. We found that Ume6 was required for transcriptional silencing of the cryptic mating-type loci HMLα and HMRa. Ume6 acted directly at these loci by binding to the cis-regulatory silencers. Ume6 also served as a block to polyploidy and was required for repression of three meiotic genes, independently of the Rpd3 and Sin3 corepressors. Mating type switching from MATα to MATa required the α3 protein. The α3 protein was similar to transposases of the mutator like elements (MULEs). Mutational analysis showed that the DDE-motif in α3, which is conserved in MULEs was necessary for switching. During switching α3 mobilizes from the genome in the form of a DNA circle. The sequences encompassing the α3 gene circle junctions in the MATα locus were essential for switching from MATα to MATa. Switching also required a DNA binding protein, Mating type switch 1 (Mts1), whose binding sites in MATα were important. Expression of Mts1 was repressed in MATa/MATα diploids and by nutrients, limiting switching to haploids in low nutrient conditions. In a genetic selection for strains with increased switching rates we found a mutation in the RAS1 gene. By measuring the levels of the MTS1 mRNA and switching rates in ras1, pde2 and msn2 mutant strains we show that mating type switching in K. lactis was regulated by the RAS/cAMP pathway and the transcription factor Msn2. ras1 mutants contained 20-fold higher levels of MTS1 mRNA compared to wild type whereas pde2 and msn2 expressed less MTS1 mRNA and had decreased switching rates. Furthermore we found that MTS1 contained several potential Msn2 binding sites upstream of its ORF. We suggest that these observations explain the nutrient regulation of switching. / At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: Manuscript.

Animals experience variation in their environment because of natural changes. However, due to anthropogenic disturbance, the speed and severity of these changes have recently increased. This thesis investigates how reproductive behaviours may be affected by human induced environmental change. In specific, I investigate how visual and chemical changes in the aquatic environment, caused by eutrophication, affect mating systems and sexual selection in fish. Broad-nosed- and straight-nosed pipefish, which both have been studied in detail for a long period, were used as model organisms. These two species are particularly suitable model organisms since they perform complex courtship behaviours, including the advertisement of ornaments and a nuptial dance. Further, two distinct populations were studied, one on the Swedish west coast and one in the Baltic Sea, as these two locations vary in the degree and extent of environmental disturbance, in particular turbidity. I found that changes in the visual environment had no impact on the development of female sexual ornaments in these sex-role reversed pipefishes, but it hampered adaptive mate choice. Turbidity also had a negative effect on reproductive success in the Baltic Sea population. Changes in the chemical environment in the form of increased pH reduced the probability to mate, while hypoxia did not alter mating propensity. However, hypoxic water delayed the onset of both courting and mating. Hence, human induced change in aquatic environments may alter the processes of sexual selection and population dynamics.

Flowering plants possess striking variation in reproductive traits and mating patterns, even among closely related species. In this thesis, I investigate morphological variation, mating and genetic diversity of five taxa of bird-pollinated Babiana (Iridaceae), including two species with specialized bird perches. Field observations in 12 populations demonstrated that sunbirds were the primary pollinators. Babiana ringens exhibited correlated geographic variation in flower and perch size. Controlled field pollinations revealed self-compatibility and low pollen limitation in B. ringens subspecies, and self-incompatibility and chronic pollen limitation in B. hirsuta. Allozyme markers demonstrated moderate to high selfing rates among populations and considerable variation in levels of genetic diversity. In B. ringens there was a positive relation between the geographic and genetic distance of populations. The results of a manipulative field experiment indicated position-dependent herbivory on inflorescences of B. hirsuta and this could play a role in the evolution of specialized bird perches in Babiana.