This thesis integrates the fields of sexual selection, parental investment and sex role theory by investigating mutual mate choice and mate competition in the sex role reversed deep snouted pipefish Syngnathus typhle (Pisces: Syngnathidae) through a series of laboratory experiments. In S. typhle, the female transfers her eggs to the male's brood pouch where they are nourished and oxygenated for about a month, when the male gives birth to the independent fry. Mate choice was found to be adaptive. Both sexes benefited from mating with preferred partners in terms of increased offspring viability and got larger, or faster growing, offspring when mating with large fish. Females were also shown to prefer males with thicker brood pouches. Thus, females, the more competitive sex, had multiple preferences. Both male and female choice behaviour was found to be flexible and influenced by available information on partner quality. In addition, males, but not females, copied the mate choice of consexuals. Both sexes were found to take their own quality in relation to surrounding competitors into account when deciding whether to display to potential partners. Male-male competition was found to influence both the mate choice of males and, potentially, overrule the mate choice of females. Males did not compete as intensely as females, nor did they use their sexual ornament in this context as females do. Rather, the ornament was used in interactions with females, and males that displayed more received more eggs. The findings in this thesis emphasise the importance of not viewing mate choice and competition as opposite behaviours, but rather to apply a dynamic approach in mate choice studies, integrating choice and competition in both sex

A central goal of evolutionary biology is to understand the processes responsible for morphological, genetic and behavioural differentiation between sexes and among geographically distinct populations. Perhaps the most significant processes are genetic drift, natural selection, phenotypic plasticity and sexual selection. The main aim of this thesis was to investigate differentiation among individuals and populations of the sex-role reversed pipefish (Syngnathus typhle) and, consequently, determine which processes may be responsible for emerging patterns. This unique species is characterised by males predominately choosing amongst displaying females. In this thesis I revealed, on a microgeographic scale, morphological differentiation without genetic divergence among populations. Interestingly, females differed in size whereas the males did not. For females in this sex-role reversed species, the costs of expressing a plastic phenotype may be outweighed by the potential gains from greater survivorship, higher fecundity or increased mating success. Thus, females gain the ability to make themselves as conspicuous and attractive to males as possible in the specific environment they are living. Moreover, behavioural experiments, which focussed on describing “personalities”, reproductive investment strategies, and mate-sampling tactics, also indicated that males as well as females had the behavioural plasticity required to adjust to the environment in which they live. To this end, using video playbacks as experimental stimuli may be especially rewarding in this species. Overall, the studies in this thesis acknowledge the ability of species to fine-tune their phenotype to maximise fitness and, therefore, highlight the importance of considering patterns of differentiation in an environment-specific context.

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.

Canada is the fourth largest producer of farmed salmon in the world, with Atlantic salmon being the major species cultivated. Paradoxically British Columbia (BC), which borders the Pacific Ocean, is the major producer province where Atlantic salmon was introduced in the mid-80’s. Escaped salmon may constitute a threat to natural populations of Pacific salmon as they compete for the same resources such as food and spawning territory. A potential solution to the aquaculture industry would be to further develop the aquaculture of native species in the region. The work presented here used semi-natural spawning channels to evaluate the effects of breeding strategies and early-rearing environments on the immune performance of Chinook salmon. Breeding strategy was tested analyzing artificial hatchery practices versus semi-natural propagation in spawning channels. Early-rearing environmental assessment contrasted indoor plastic hatchery tanks with outdoor gravelled-bottom spawning channels. A disease challenge involving over 1400 fish showed interaction effects between breeding strategy and rearing environment. Fish artificially mated presented a disease susceptibility influenced by the rearing environment. The contrary occurred in the offspring of self-breeding brood stock in the spawning channels, as no differences were observed in their susceptibility to the disease regardless of rearing environment. Monitoring of anti-Vibrio anguillarum antibodies during the disease challenge and a follow up of the survivors in sea net pens further confirmed the interaction between breeding strategy and rearing environment. Gene expression in pre- and post-infected artificially propagated fish showed differential gene expression when analyzed with a 695-gene cDNA microarray for Chinook salmon. Genotyping of major histocompatibility (MH) class II β1 alleles showed a tendency of a higher heterozygosity in survivors as expected, as well as a general tendency of a higher heterozygosity in semi-naturally propagated fish. The latter is likely a direct consequence of MH-linked mate choice, which was recently described in Chinook salmon (Neff et al., 2008). To further characterize the mating system of Chinook salmon in the spawning channels, brood stock were genotyped at 12 microsatellite loci. Females and males were found to mate randomly with regards to genetic pairwise relatedness, but they tended to mate with fish of similar condition as revealed by their pairwise differences in Fulton’s condition factor. This work demonstrated that genotype-by-environment interactions can modify the disease resistance of Chinook salmon. More importantly, these effects were seen after just one round of semi-natural spawning of domesticated hatchery fish, suggesting that further studies on spawning channels may highlight other hidden benefits. Therefore, breeding strategy and early-rearing environment should be considered when propagating cultured stocks. The use of more natural propagation methods such as spawning channels could improve the immune performance of Chinook salmon and help to expand the aquaculture of this native species in BC.

Choosing with whom to mate is one of the most important decisions a female makes in her lifetime and inter-individual variation of these preferences can have important evolutionary consequences. In order to get a complete understanding of why and how females choose a mate, we must identify factors that can contribute to variation of female mate choice. Many decades of research sought to understand ultimate mechanisms of female mate choice with proximate mechanisms receiving a lot more attention in recent years. For my thesis, I identify intrinsic and extrinsic factors that correlate with individual variation of female Xiphophorus nigrensis mate preference. I provide evidence that a female’s size (e.g. age and sexual experience) as well as male behavioral displays can predict female mate preference. Using genes associated with female mate preference (neuroserpin, neurologin-3), I identify four brain regions (Dl, Dm, HV, POA) that show significant differences in gene expression between females exhibiting high preference for males relative to females displaying little mate preference. Neuroserpin and neuroligin-3 gene expression within these brain regions are also positively correlated with female mate preference behavior. Two of these brain regions (Dm and Dl) integrate multisensory information and are found in the putative teleost mesolimbic reward circuitry; the other two regions (HV and POA) are involved in sexual behaviors. With the implication of the reward circuitry, I assess whether there are changes in dopamine synthesis (via tyrosine hydroxylase, TH) in dopaminergic brain regions associated with the degree of mate preference. I do not find evidence of rapid changes (within 30 minutes) of TH expression (i.e. dopamine synthesis) in dopaminergic brain regions related to variation in female mate preference. Collectively my results suggest that mate preference behavior in the brain may be coordinated not just through regions associated with sexual response but also through forebrain areas that may integrate primary sensory information, with no associated changes of a proxy for dopamine synthesis in dopaminergic brain regions. / text

<p>Sexual selection has long been accepted as a widespread force of evolution shaping male traits across taxa. In recent years, biologists have begun to investigate the extent to which sexual selection may also shape traits among females. However, current models of sexual selection have largely been developed using assumptions that--while generally met in males--often do not apply to females. Thus, attempts to apply these contemporary models to the study of sexual selection in females reveal weaknesses in the theoretical framework for sexual selection research. One consequence of this for empirical research is that researchers often infer the action of sexual selection upon evidence of male mate choice. Although male mate choice is increasingly common, it is much less likely to exert selection pressure than its female counterpart. I begin by proposing a conceptual framework that explicitly accounts for ...Next, I investigate a female trait that has recently become an iconic example of sexual selection in females; that is the exaggerated estrous swellings of cercopithecine primates. By combining morphological data collected with a non-invasive photographic method and observational behavioral data with longitudinal ecological and demographic data from the ongoing Amboseli Baboon Research Project, I examine the sources of variance in this exaggerated signal of fertility. Finally, I test the hypothesis that male baboons prefer females with larger sexual swellings because those females have higher fitness. I find no evidence to support this hypothesis. Instead, my results suggest that mate choice among male baboons has evolved to detect, not the intrinsic quality of the female as has typically been proposed, but the quality of a reproductive opportunity.</p> / Dissertation

This thesis is an investigation of the stress response of salmonid fish and the regulation of the endocrine mechanisms of growth during changes in physiological conditions. Controlled by the HPI axis, the stress response incurs extensive catabolic demand on endogenous metabolite stores at the expense of growth through catabolic actions under the assumed direction of cortisol. It is suspected that the stress response also suppresses the growth-promoting actions of the GH/IGF-I/IGFBP axis. The central theme of this thesis was to characterize the influence the stress response on the endocrine regulation of growth during conventional (e.g. emersion, salinity transfer) and unconventional stresses (e.g. competition, social interaction, parasite infection), using rainbow trout (Oncorhynchus mykiss) and Chinook salmon (Oncorhynchus tshawytscha) as models. Findings corroborate the inhibitive impact of chronic stress on growth through catabolism of endogenous metabolites in the presence of cortisol levels representative of moderate stress in salmonids. Trout in infected with Cryptobia salmositica, demonstrated similar evidence of pathogen-induced growth suppression via changes in catabolic elements within the GH/IGF-I/IGFBP axis in a similar manner to hypercorticoidic fish, but without elevated cortisol despite clear physical duress. Accompanying reduction in food intake and change to nutritional status influenced much of the growth-suppressing impacts observed on the endocrine axis during disease incidence. Moreover, Cryptobia infection inhibited cortisol signaling and production the pituitary and in the interrenals, respectively. In Chinook salmon, the endocrine response to stress was altered by parental breeding strategy and early rearing environment; traditional hatchery breeding and rearing methods impacted growth performance during physiological challenge when contrasted to mate choice cohorts. Rearing Chinook in a semi-natural channel environment revealed clear differences in performance between these fish stocks that were not observable in the hatchery environment. Moreover, social interaction and competition between mate choice and hatchery-bred salmon influenced the inhibitive effects of the stress response on growth performance, as well as the physiological responses to endocrine-regulated changes during smoltification. This thesis characterizes the novel regulatory actions of the stress response on the endocrine growth axis via regulation of both central and peripheral elements of the GH/IGF-I/IGFBP axis. / NSERC, OGS

Mate choice among males is relatively understudied, despite recent evidence supporting its ubiquity. Theory predicts male mate choice in response to variation in female quality, and male mating strategies that limit polygyny. However empirical research investigating these connections, particularly under natural conditions, is generally lacking. Using the Western black widow spider, Latrodectus hesperus, I investigated male mate choice, male investment, and the advantage of mating with females that differ in their potential to deliver fitness benefits to males via higher fecundity and/or reduced risk of sperm competition. Males were found to be preferentially attracted to larger, high-diet virgin females over females that were smaller due to a lower diet, or having been previously mated, or both. Through a three-year field study on females I found that males likely benefit from selectivity, as smaller, low-diet females often failed to deposit any egg sacs throughout the breeding season, likely due to their shortened web-site tenure and/or lack of bodily resources. I investigated the costs and benefits to male genital breakage in L. hesperus, a strategy typically assumed to result in male sterility while providing paternity protection. I found that genital mutilation in L. hesperus did not cause sterility and that males were capable of inseminating multiple females, likely because of the comparatively low amount of genital damage and the likelihood of the efficient placement of genital fragments. Evidence from double mating trials supported the efficacy of broken genital fragments as plugs, but first male sperm precedence was often maintained in cases where male genital breakage failed or fragments were positioned incorrectly, which occurred frequently. Together, these laboratory and field experiments contribute to a more complete view of mate choice.

Mate choice among males is relatively understudied, despite recent evidence supporting its ubiquity. Theory predicts male mate choice in response to variation in female quality, and male mating strategies that limit polygyny. However empirical research investigating these connections, particularly under natural conditions, is generally lacking. Using the Western black widow spider, Latrodectus hesperus, I investigated male mate choice, male investment, and the advantage of mating with females that differ in their potential to deliver fitness benefits to males via higher fecundity and/or reduced risk of sperm competition. Males were found to be preferentially attracted to larger, high-diet virgin females over females that were smaller due to a lower diet, or having been previously mated, or both. Through a three-year field study on females I found that males likely benefit from selectivity, as smaller, low-diet females often failed to deposit any egg sacs throughout the breeding season, likely due to their shortened web-site tenure and/or lack of bodily resources. I investigated the costs and benefits to male genital breakage in L. hesperus, a strategy typically assumed to result in male sterility while providing paternity protection. I found that genital mutilation in L. hesperus did not cause sterility and that males were capable of inseminating multiple females, likely because of the comparatively low amount of genital damage and the likelihood of the efficient placement of genital fragments. Evidence from double mating trials supported the efficacy of broken genital fragments as plugs, but first male sperm precedence was often maintained in cases where male genital breakage failed or fragments were positioned incorrectly, which occurred frequently. Together, these laboratory and field experiments contribute to a more complete view of mate choice.

Canada is the fourth largest producer of farmed salmon in the world, with Atlantic salmon being the major species cultivated. Paradoxically British Columbia (BC), which borders the Pacific Ocean, is the major producer province where Atlantic salmon was introduced in the mid-80’s. Escaped salmon may constitute a threat to natural populations of Pacific salmon as they compete for the same resources such as food and spawning territory. A potential solution to the aquaculture industry would be to further develop the aquaculture of native species in the region. The work presented here used semi-natural spawning channels to evaluate the effects of breeding strategies and early-rearing environments on the immune performance of Chinook salmon. Breeding strategy was tested analyzing artificial hatchery practices versus semi-natural propagation in spawning channels. Early-rearing environmental assessment contrasted indoor plastic hatchery tanks with outdoor gravelled-bottom spawning channels. A disease challenge involving over 1400 fish showed interaction effects between breeding strategy and rearing environment. Fish artificially mated presented a disease susceptibility influenced by the rearing environment. The contrary occurred in the offspring of self-breeding brood stock in the spawning channels, as no differences were observed in their susceptibility to the disease regardless of rearing environment. Monitoring of anti-Vibrio anguillarum antibodies during the disease challenge and a follow up of the survivors in sea net pens further confirmed the interaction between breeding strategy and rearing environment. Gene expression in pre- and post-infected artificially propagated fish showed differential gene expression when analyzed with a 695-gene cDNA microarray for Chinook salmon. Genotyping of major histocompatibility (MH) class II β1 alleles showed a tendency of a higher heterozygosity in survivors as expected, as well as a general tendency of a higher heterozygosity in semi-naturally propagated fish. The latter is likely a direct consequence of MH-linked mate choice, which was recently described in Chinook salmon (Neff et al., 2008). To further characterize the mating system of Chinook salmon in the spawning channels, brood stock were genotyped at 12 microsatellite loci. Females and males were found to mate randomly with regards to genetic pairwise relatedness, but they tended to mate with fish of similar condition as revealed by their pairwise differences in Fulton’s condition factor. This work demonstrated that genotype-by-environment interactions can modify the disease resistance of Chinook salmon. More importantly, these effects were seen after just one round of semi-natural spawning of domesticated hatchery fish, suggesting that further studies on spawning channels may highlight other hidden benefits. Therefore, breeding strategy and early-rearing environment should be considered when propagating cultured stocks. The use of more natural propagation methods such as spawning channels could improve the immune performance of Chinook salmon and help to expand the aquaculture of this native species in BC.