In order to maximize fitness, individuals must partition their limited resources among competing physiological processes, creating negative statistical associations between processes known as “life-history trade-offs”. Evidence indicates that individuals tend to decrease their reproductive investment when confronted with a significant immunological challenge in order to increase investment in immune defense. This trade-off is often accompanied by a significant decrease in the sexual signal, which provides an honest signal of the male’s infection status to potential mates. However, if individual residual reproductive value is low, they may instead increase their reproductive investment to maximize reproductive success before the end of their life (a.k.a. terminal investment). Here, we investigate the potential for terminal investment in the ground cricket Allonemobius socius by inoculating males with varying dosages of an immune challenge. We predicted that both high dose and advanced male age would induce terminal investment. Furthermore, we predicted that terminally investing males would produce a dishonest signal by increasing their signaling effort. We found that upon infection We found that upon infection, young males and old males differentially alter their reproductive strategy. Young males exhibited the classic deceleration of reproductive effort. However, old males increased their calling song energetics and decreased their parental investment (nuptial gift size), suggesting that old males are dishonestly signaling their condition to the female.

Male reproductive success in many mammals depends on their ability to allocate sufficient energetic resources to mating competition. Such costs are particularly pronounced in species with high levels of sexual body dimorphism, intense polygyny and distinct breeding seasons. I tested the hypothesis that male reproductive effort incurs significant energetic costs in wild chimpanzees (Pan troglodytes), a species with moderate sexual dimorphism, promiscuous mating and lack of breeding seasonality. My field studies combined behavioral observations on male chimpanzee behavior with non-invasive sampling of urinary C-peptide (UCP). UCP is a biomarker of insulin production that indexes individual energy balance. This dissertation contributes to the understanding of UCP as an energy assay by (1) validating the application of UCP for assessing dietary quality in bonobos (Pan paniscus) at Kokolopori, DRC and (2) providing a detailed assessment of diurnal variation in UCP levels in relation to short-term changes in food intake in chimpanzees at Kanyawara, Kibale NP, Uganda. I used UCP measurements in conjunction with full-day focal observations of male chimpanzees to assess the energetic costs of male-male competition for status and mating opportunities. Data on feeding time and rates of aggression suggested that males experience a reduction in energy intake and an increase in energy expenditure when highly attractive parous females were in estrus. UCP data supported these conclusions because males had lower UCP levels on mating days, and rates of aggression were negatively associated with UCP levels. Mean daily party size was also associated with low UCP levels, controlling for the presence of estrous females. Habitat-wide availability of preferred fruits was positively associated with male rates of aggression suggesting that energy availability mediates male investment towards energetically costly competitive behaviors. Contrary to expectations males who were most successful in obtaining copulations (high-ranking males) did not suffer higher energetic costs than lower-ranking males during periods of mating competition. Costs or reproductive effort include both direct competition for matings and long-term competition over social status. Maintenance of social rank over long periods appears to be particularly important in this slow-reproducing, long-lived and nonseasonally breeding primate. / Human Evolutionary Biology

Abstract Both the length of the season that is favourable for insect growth and reproduction and the number of generations emerging per season (voltinism) increase with decreasing latitude. Thus, time constraints on reproduction and juvenile development decrease with decreasing latitude, except where voltinism changes and time constraints suddenly increase as the season must be shared with one more generation. I studied the evolution of insect life histories in relation to time constraints from two perspectives: polymorphism and clinal variation. Life history polymorphism in seasonal environments was studied with the butterfly Pieris napi that has discrete life history strategies, and polymorphic natural populations. Experimental studies showed that asymmetric intraspecific larval competition and divergent timing of reproduction between the strategies may promote the maintenance of polymorphism. A simulation model showed that the divergent timing of reproduction between the strategies is sufficient to maintain polymorphism even in the absence of intraspecific competition. Clinal variation was studied empirically with four geometrid moths (Cabera exanthemata, Cabera pusaria, Chiasmia clathrata and Lomaspilis marginata) and generally in theory. Due to latitudinal saw-tooth variation in time constraints, traditional theory predicts a saw-tooth cline in body size and development time. A common garden experiment with the four geometrid moths did not support the traditional theory even when a saw-tooth cline in body size was found, suggesting that the theory is based on unrealistic assumptions. A theoretical analysis showed that reproductive effort should be high in populations under intense time constraints and low in populations experiencing no time constraints, resulting in a saw-tooth cline. In the four geometrid moths, support for these predictions was found, although the observed clinal variation deviated from the predicted pattern. The results imply that clinal variation is expected in almost any continuous life history trait, whereas polymorphism of different strategies may emerge when life histories fall into discrete categories. When inferring the evolution of a single trait, complex interdependencies among several traits should be considered, as well as the possibility that the time constraints are not similar for each generation in multivoltine populations. / Tiivistelmä Hyönteisten kasvu ja kehitys ovat pääsääntöisesti mahdollisia vain kesän aikana. Etelään päin mentäessä kesä pitenee, ja saman kesän aikana kehittyvien hyönteissukupolvien määrä kasvaa. Kesän pituus aiheuttaa lisääntymiseen ja toukkien kasvuun kohdistuvan aikarajoitteen, joka heikkenee etelään päin siirryttäessä. Aikarajoite kuitenkin tiukkenee siellä, missä yksi uusi sukupolvi ehtii juuri kehittymään saman kesän aikana, sillä kesä on nyt jaettava useamman sukupolven kesken. Väitöstyössä tarkastelin hyönteisten elinkierto-ominaisuuksien evoluutiota suhteessa aikarajoitteisiin sekä diskreetin että jatkuvan muuntelun näkökulmista. Diskreettiä muuntelua tutkin lanttuperhosella (Pieris napi), jolla esiintyy diskreettejä elinkiertostrategioita. Kokeellisesti osoitin, että toukkien välinen kilpailu on epäsymmetristä, mikä yhdessä eri elinkiertostrategioiden erilaisen lisääntymisen ajoittumisen kanssa voi ylläpitää diskreettiä muuntelua. Simulaatiomalli osoitti, että erilaiset elinkiertostrategiat voivat säilyä populaatiossa pelkästään niiden erilaisen lisääntymisen ajoittumisen ansiosta. Elinkierto-ominaisuuksien jatkuvaa muuntelua tutkin neljän mittariperhosen (Cabera exanthemata, Cabera pusaria, Chiasmia clathrata ja Lomaspilis marginata) avulla ja teoreettisesti yleisellä tasolla. Aikaisempi teoria ennustaa ruumiinkoon ja kehitysajan muuntelevan sahalaitakuvion mukaisesti siirryttäessä pohjoisesta etelään, koska aikarajoitteet muuntelevat samalla tavalla. Tämä teoria perustunee epärealistisiin oletuksiin, koska kokeelliset tulokset eivät tukeneet teoriaa silloinkaan, kun mittariperhosten ruumiinkoko muunteli ennustetulla tavalla. Teoreettinen tutkimus osoitti, että myös lisääntymispanostuksen tulisi muunnella sahalaitakuvion mukaisesti suhteessa kesän pituuteen siten, että se on korkeimmillaan siellä, missä aikarajoitteet ovat tiukat. Mittariperhosten tutkiminen antoi jossain määrin tukea tälle ennusteelle. Tulosten perusteella jatkuvaa maantieteellistä muuntelua ennustetaan elinkierto-ominaisuuksille, jotka muuntelevat jatkuvalla asteikolla. Erilaiset elinkiertostrategiat voivat sen sijaan säilyä populaatiossa, jos elinkierto-ominaisuuksien muuntelu on diskreettiä. Eri ominaisuuksien monimutkaiset vuorovaikutukset sekä eri sukupolvien mahdollisesti kokemat erilaiset aikarajoitteet olisi syytä tuntea, kun tarkastelun kohteena on yksittäisen ominaisuuden evoluutio.

Altitudinal gradients offer attractive opportunities for studies of population differentiation in response to environmental heterogeneity. In this thesis, I examined population differentiation along altitudinal gradients by combining common-garden experiments with field studies and experiments in alpine, subalpine and boreal populations of the perennial herb Solidago virgaurea. More specifically, I determined whether leaf physiology in terms of nitrogen concentration and resorption, flowering phenology, flower production and reproductive effort vary along altitudinal gradients. Nitrogen concentration in green leaves were higher in alpine than in subalpine and boreal populations. These differences persisted when plants were grown from seeds in a common-garden experiment at two sites, suggesting that the differences have a genetic component. There was mixed support for a trade-off between maximized carbon gain through the maintenance of high nitrogen concentration, and minimized nitrogen loss through high resorption. In their natural habitats alpine populations began flowering later than subalpine populations, but this difference was reversed when plants were grown in a common environment. This suggests that genetic differences among populations counteract environmental effects and reduce phenotypic variation in flowering time among populations. Flowering time thus shows countergradient genetic variation in S. virgaurea. In a common-garden experiment, boreal populations produced more flowers and had a higher reproductive effort than subalpine and alpine populations indicating habitat-specific genetic differences in reproductive allocation. In a field study, which included three populations, seed set was close to zero in the alpine population, intermediate in the subalpine population, and high in the boreal population. Experimental flower removal showed that seed production was associated with a considerable cost in terms of reduced flowering propensity the following year, but did not support the hypothesis that a large floral display is important for pollination success.

Sexually selected ornaments are among the most spectacular traits in nature. Indeed, the extreme costs associated with producing sexual traits seem to play a crucial role in their evolution by enforcing honest levels of advertisement: only males with high levels of acquired resources (or high ‘condition’, as it is known in the literature) can afford to produce extravagant signals, a phenomenon which maintains signal reliability in a constant environment. In my thesis I examine many implications of this condition-dependent model of ornament and preference evolution for variation in age-dependent allocation to sexual signals and other life history traits. In Chapter 1, I review theoretical implications of condition-dependent signalling for life history and sexual selection theory. I note that a universal cost of expenditure in sexual advertisement is metabolic in nature: metabolites used to fund ornament expression are by definition unavailable to other life history traits that compete for a limited resource pool. This universal constraint on expenditure does more than maintain honesty (as noted above), however: the reliance of sexual displays on high levels of nutrient acquisition may help maintain genetic variation in sexual signals that would otherwise be eroded by strong mate choice, and without which the selective basis for good-genes choice would disappear. Three mechanisms in particular probably help to maintain genetic variation in acquisition. 1) Because acquiring resources and converting them efficiently to useful forms depends on the high function of many biochemical pathways, condition is undoubtedly highly polygenic, which slows the erosion of genetic variation under strong directional selection by females (especially in the presence of epistatic interactions). 2) The highly polygenic nature of condition also presents a large target for mutation, which continually restores variation at the loci under selection. 3) The many loci underlying condition may also be particularly sensitive to environmental heterogeneity in time or space. By favouring the most ornate males, females acquire high performing genes for their offspring, regardless of the precise allele combinations that have conferred the ability to acquire resources. Selection on specific alleles is liable to fluctuate over time or space whenever allelic performance is strongly context-specific. I close by noting the considerable challenges in advancing research on sexual selection and life history allocation, including the fact that two key processes central to life history (acquisition and allocation) are latent variables that interact in complexways and are intrinsically difficult to measure empirically. In the remainder of my thesis I conduct a series of experiments involving decorated crickets, Gryllodes sigillatus, which are useful models for studying life history because they enable precise measurement of male reproductive effort. Male G. sigillatus face important allocation decisions owing to the highly polyandrous nature of females, and the substantial costs involved in signalling and mating. Chapter 2 examines sex differences in age-dependent reproductive effort as a function of diet and development stage. I reared outbred crickets using four combinations of diet nutritional quality, and studied the effects of these combinations on male and female reproductive effort (calling effort in males and fecundity in females) and longevity. While I expected males to be more sensitive than females to variation in diet and developmental changes in its quality, I actually observed the opposite: males in all treatments increased calling effort over time, exhibiting consistently positive covariance between calling effort and longevity across treatments. By contrast, the relationships between female reproductive effort and longevity changed dramatically across treatments, and females who lived to intermediate ages had the highest fecundity. Although my results support sex-specific selection on life history allocation over time, a compelling additional explanation for my findings relates to the strategic role of calling for achieving male fitness. In the absence of positive feedback from potential mates, perhaps male allocation to sexual advertisement is careful and only increases gradually as a function of accumulating metabolic resources and increasing risk of intrinsic mortality. Alleles underlying condition are expected to be particularly sensitive to environmental heterogeneity. While this sensitivity may help maintain additive variation in male quality (which is essential for the sustenance of adaptive good-genes mate choice, as noted in Chapter 1), too much environmental sensitivity could also underiii mine the signal value of the male trait. For example, if there are strong genotypeby- environment interactions (GEIs) for sexual advertisement, in a rapidly changing environment females risk favouring a male whose alleles are no longer best suited to current conditions. This problem is particularly pressing for animals like crickets where males exhibit a behaviourally plastic sexual display (such as calling), and so may dynamically adjust signalling effort over time. In Chapter 3, I used inbred lines of decorated crickets to quantify age and diet dependent genetic variation in male signalling. I demonstrate that while genetic correlations across diets were quite strong for morphological traits, correlations between measures of the male sexual trait rapidly approached zero as I increased the distance in time (i.e., across widely spaced ages) or diet (i.e., comparing more dissimilar dietary histories) between samples. While extrapolating from my laboratory experiments to nature is difficult, my findings nevertheless cast doubt on the value of behaviourally dynamic signals (such as cricket calls) for reliably indicating genetic quality in realistically complex environments. In Chapter 4 I used physiological assays to evaluate factors affecting metabolite storage and use over time in decorated crickets. I manipulated the acquisition ability of all males using artificial diets that varied linearly in nutrient quality, and manipulated access to female mates over the course of the second week of adult life. By sacrificing crickets at key stages before and after manipulating the diet and social environment, I was able to estimate changes in stored metabolites, and relate these changes to calling effort and longevity. During the first week of adulthood (in the absence of females), higher diet quality significantly increased calling effort and storage of lipid, glycogen, and carbohydrate (but not protein). The presence of females increased both the probability of calling and the amount of calling during the second week, whereas diet quality only improved calling effort. By the end of the second week, calling effort had decreased, even by high quality males in the presence of females, suggesting a depletion of resources. Furthermore, the loss of condition during week 2 covaried with calling effort during the previous week irrespective of diet. Males who started the second week in high condition lost more glycogen and carbohydrate than rivals; meanwhile, lipid accumulation covaried positively with calling effort during week 2. The contrasting patterns of storage and use for lipids compared to the ‘quick-release’ metabolites (glycogen and carbohydrates) affirms starkly distinct functions for the different storage components, and underlines the importance of specific physiological measures in life history research. Finally, in the general discussion, I attempt to synthesise my thesis’s contributions to the study of life history trade-offs involving behavioural sexual displays.

In this thesis, I combined comparative and experimental approaches to examine selection on reproductive traits and population differentiation in the insect-pollinated, self-incompatible, perennial herb Primula farinosa. More specifically, I (1) determined whether the effects of floral display and interactions with pollinators and seed predators, and plant reproductive success were frequency-dependent and affected by surrounding vegetation context, (2) examined the consequences of intermittent drought years on population dynamics using numerical simulations based on demographic data collected over seven years, (3) analyzed among-population differentiation in flowering phenology and reproductive allocation, and its relationship to soil-depth at the site of origin. A field experiment suggested that conspicuous plants facilitate inconspicuous plants in terms of pollinator attraction, and that the facilitation effect is contingent on the height of the surrounding vegetation. Further experiments revealed that both mutualistic and antagonistic interactions can result in frequency-dependent selection on floral display. Among inconspicuous plants, both fruit initiation, and damage from seed predators increased with the proportion of the conspicuous morph. The relative strength of these effects, and therefore their net outcome on the relationship between morph ratio and seed production varied among years. I combined information on vital rates and their relation to environmental conditions in simulations to predict future population viability in changing environments. Simulated stochastic population growth rate decreased with increasing frequency of drought years. Reproductive allocation varied significantly among populations both in the field and in a common-garden experiment, but was correlated with soil depth at the site of origin only in the field. The results suggest that among-population variation in reproductive effort in the field mainly reflects plastic responses to environmental conditions, and that this plasticity may be adaptive. The common-garden experiment suggested that the study populations have diverged genetically in flowering time.

<p>In this thesis, I combined comparative and experimental approaches to examine selection on reproductive traits and population differentiation in the insect-pollinated, self-incompatible, perennial herb <i>Primula farinosa</i>. More specifically, I (1) determined whether the effects of floral display and interactions with pollinators and seed predators, and plant reproductive success were frequency-dependent and affected by surrounding vegetation context, (2) examined the consequences of intermittent drought years on population dynamics using numerical simulations based on demographic data collected over seven years, (3) analyzed among-population differentiation in flowering phenology and reproductive allocation, and its relationship to soil-depth at the site of origin. </p><p>A field experiment suggested that conspicuous plants facilitate inconspicuous plants in terms of pollinator attraction, and that the facilitation effect is contingent on the height of the surrounding vegetation. Further experiments revealed that both mutualistic and antagonistic interactions can result in frequency-dependent selection on floral display. Among inconspicuous plants, both fruit initiation, and damage from seed predators increased with the proportion of the conspicuous morph. The relative strength of these effects, and therefore their net outcome on the relationship between morph ratio and seed production varied among years. </p><p>I combined information on vital rates and their relation to environmental conditions in simulations to predict future population viability in changing environments. Simulated stochastic population growth rate decreased with increasing frequency of drought years. </p><p>Reproductive allocation varied significantly among populations both in the field and in a common-garden experiment, but was correlated with soil depth at the site of origin only in the field. The results suggest that among-population variation in reproductive effort in the field mainly reflects plastic responses to environmental conditions, and that this plasticity may be adaptive. The common-garden experiment suggested that the study populations have diverged genetically in flowering time.</p>

Age is often a neglected factor in ecological studies. However, age-related changes in reproduction and survival of organisms may strongly influence population dynamics. The Gotlandic population of collared flycatchers is a perfect system for studying age-related changes in the wild, as the exact age and reproductive history of most individuals is known. Collared flycatchers (Ficedula albicollis) on Gotland show the typical pattern of age-related changes in survival and reproductive success; both factors show an increase early in life and a decrease late in life. This thesis presents a broad study not only of age-related patterns of reproduction and immunity, but also proposes the mechanisms driving these patterns. My results show that in addition to survival probability and reproductive performance, reproductive costs and life-history trade-offs also change with progressing age. There is a significant increase in reproductive performance at the population level during first years of life which result from selection against low quality phenotypes. On the individual level this pattern is best explained by an optimization of reproductive effort. However, high quality individuals have higher reproductive success as early as their first breeding event and are long-lived. Thus, they seem to adopt a different strategy than lower quality, short-lived individuals. Differences in individual quality seem to be shaped by the developmental conditions experienced as nestlings. Fledglings with longer tarsi, but lower body mass become long-lived, high quality adults. Young individuals breeding for first time pay higher costs of reproduction. They also express a limited ability to reduce these costs by breeding in high quality territories when compared to middle-aged individuals. Young individuals seem to invest more into self-maintenance, whereas old individuals reduce the level of self-maintenance (measured as immune response) and redistribute their investment towards reproduction. Thus, old individuals are limited in their ability to reduce reproductive costs under favorable conditions, especially as they also senesce, which pattern is also shaped by individual quality. Variation in individual quality appears to have an strong effect on age-related survival probability, reproductive performance, reproductive costs, and even life-history decisions. Therefore, taking this factor into account in studies of life-history patterns is necessary to obtain reliable results.

Rising archipelagos provide unique settings for the study of the temporal and spatial dynamics of their biota. This offers the possibility to study the ecology and genetics of early successional processes; both between islands that differ in age and within islands when already established organisms have to keep pace with the changing environment. I have worked in the Skeppsvik Archipelago housing about 100 islands that due to land uplift vary in age, thus representing various stages of primary succession. I have utilized a naturally created metapopulation of Silene dioica, which in this archipelago is a dominant plant of the deciduous border, offering the possibility to study subpopulations on islands of different ages and in different phases of primary succession. Many plant species exist as metapopulations, which consists of many local populations which may differ in size and degree of connectivity. Metapopulations are further characterized by recurrent colorizations and extinctions of local populations, meaning that a species continually must disperse and relocate to allow for persistence in this system. For a dioecious plant species, gene flow is in the shape of seeds and pollen and to allow for the persistence of populations, it is necessary that levels of seed dispersal and pollen gene flow are enough to ensure both colonisation, establishment and subsequent population growth. Levels of seed dispersal and pollen gene flow is in turn influenced by how the two sexes partition resources between reproduction, growth and survival. In paper I, I combined a field survey, a common garden experiment and a nine-year demographic study to assess the demographic consequences of sex-specific resource allocation and to investigate if differential costs of reproduction may be a driver in the evolution of sexual dimorphism in dioecious Silene dioica. Significant somatic intersexual dimorphism was found with females being the larger sex, both in terms of above – and belowground biomass. Furthermore, the reproductive effort of females exceeds that of males across a growing season which largely confirms what has been observed earlier in dioecious, herbaceous plant species. According to the cost of reproduction hypothesis, high reproductive investment should result in trade-offs with somatic and/or life-history traits. Somatic trade-offs were not observed, and instead I found strong, positive associations between reproductive investment and vegetative growth in both males and females. Compensation mechanisms were found in both sexes although females are generally more efficient at compensating their reproductive costs. At the end of a flowering season, after having paid the current costs of reproduction, females are better than males at provisioning perennial roots and rosettes potentially influencing the ability to set future flower buds and winter survival. Trade-offs were found between current and future reproduction and survival, but this is condition dependent and compensation through frequency of flowering plays an important role. The cost of reproduction hypothesis appears to play some role in driving the somatic and demographic sexual dimorphisms observed in this system but sexual selection acting on males will be a fruitful avenue for future research. In paper II, I investigated the population genetic consequences of metapopulation dynamics in Silene dioica. The occurrence of islands in different phases of primary succession together with successional gradients across islands, makes it possible to investigate the genetic dynamics occurring in an age-structured metapopulation across several hierarchical levels. Genetic diversity and differentiation were estimated in eight young, recently colonised populations and in ten populations of an intermediate successional stage. Young populations were less genetically diverse compared to older populations, indicating that bottlenecks, created by small founding groups derived from a limited number of source populations, reduce the genetic diversity within newly founded populations. The observation of strong genetic structure both between islands and between patches with islands, indicates that gene flow is restricted across several spatial levels in this system. However, the lack of statistically significant differences in genetic differentiation between young and intermediate populations, indicates that levels of gene flow may not be high enough to reduce the genetic differentiation that arise from the initial founder event. The patterns of sexual dimorphism and the roles of males and females in Silene dioica have evolved to allow persistence in an ecological and population context of this species. The nature of this habitat, where islands rise up from the sea creating new environments for colonisation while at the same time, autogenic primary succession processes eventually leads to extinction, means that S. dioica continuously must relocate within successional phases for its persistence. The obvious success of this dioecious plant is apparent as it is one of the few dominant species in the deciduous border. This suggests that levels of seed dispersal and gene flow are sufficient enough to allow for establishment and persistence of island populations and that the sexual dimorphisms that have evolved in this metapopulation system act to increase levels of gene flow. The "live hard – die young" strategy, with extensive flowering bouts, which we find in the males may have evolved as a way of maintaining sufficient levels of genetic diversity in the metapopulation but will only be a possible strategy if there are continuous opportunities for re-establishments. Thus, the continuous land uplift that is occurring in the northern part of the Gulf of Bothnia may very well be a prerequisite for the long-term persistence of this dioecious, perennial plant species. / Landhöjningsprocesser i skärgårdsmiljöer skapar nya habitat som gör det möjligt att studera naturliga populationer i ett rumsligt och tidsmässigt sammanhang. Detta möjliggör studier av ekologi och genetik i tidiga successionsprocesser, både mellan öar som skiljer sig åt åldersmässigt och inom öar, där redan etablerade organismer måste anpassa sig till en föränderlig miljö. Jag har utfört studier i Skeppsviks skärgård som rymmer cirka 100 öar. På grund av landhöjningen så varierar dessa öar i ålder och de representerar således olika stadier i primärsuccession. Jag har använt mig av en naturlig Silene dioica metapopulation lokaliserad i Skeppsviks skärgård. Många växtarter existerar i metapopulationer, vilket består av ett antal lokala populationer som kan skilja sig åt i storlek och grad av anknytning. Metapopulationer kännetecknas även av återkommande koloniseringar och utrotningar av lokala populationer, vilket innebär att en art kontinuerligt måste sprida sig för att garantera sin fortlevnad i detta system. Genflöde inom dioika växtarter är i form av pollen och frön, och för att populationer skall kunna överleva så är det nödvändigt att nivåerna av fröspridning och pollen-genflöde är tillräckliga för att säkerhetsställa både kolonisering, etablering och efterföljande populationstillväxt. Nivåer av fröspridning och pollen-genflöde påverkas i sin tur av hur de två könen partitionerar resurser mellan reproduktion, tillväxt och överlevnad. I den första studien har jag kombinerat en fältundersökning, ett frilandsexperiment och en nioårig demografisk studie för att undersöka de demografiska konsekvenserna av könsspecifik resursallokering och för att utreda om könsspecifika skillnader i reproduktiv kostnad kan vara en drivkraft för evolutionen av sexuell dimorfism hos den dioika växten Silene dioica. Jag upptäckte signifikant somatisk intersexuell dimorfism där honor hade betydligt mer ovanjordisk och underjordisk biomassa jämfört med hanar. Över en växtsäsong så investerar honorna mer resurser i reproduktion, vilket i stor utsträckning bekräftar vad som tidigare har observerats i örtartade, dioika växter. Enligt hypotesen för reproduktiv kostnad så bör en hög investering i reproduktion leda till trade-offs med somatiska egenskaper, t.ex. tillväxt. Jag observerade inga somatiska trade-offs och istället fann jag positiva associationer mellan reproduktion och tillväxt hos både honor och hanar. Båda könen verkar ha utvecklat kompensationsmekanismer, även om honorna generellt är mer effektiva i hur de kompenserar för sina reproduktiva kostnader. Vid slutet av en växtsäsong, efter att ha betalat för de nuvarande reproduktiva investeringarna, så är honor bättre än hanar på att allokera resurser till fleråriga strukturer, såsom bladrosetter och rötter. Detta kan potentiellt påverka hur de anlägger sina knoppanlag för nästkommande år och hur väl de överlever vintern. Trade-offs hittades mellan nuvarande reproduktion och framtida reproduktionsmöjligheter och överlevnad men detta var habitat-specifikt och kompensation med hjälp av hur ofta en växt blommar under sin livstid spelar en viktig roll. Hypotesen för reproduktiv kostnad verkar vara en del av förklaringen till den somatiska och demografiska könsdimorfism som observerats i detta system men sexuell selektion, som verkar på hanar, kan vara ett möjligt område för framtida studier. I den andra studien undersökte jag populationsgenetiska konsekvenser av metapopulationsdynamik i Silene dioica. Förekomsten av öar i olika faser av primär succession tillsammans med olika grader av succession inom öar gör det möjligt att undersöka den genetiska dynamiken som uppträder i en åldersstrukturerad metapopulation över flera hierarkiska nivåer. Genetisk mångfald och differentiering uppskattades i åtta unga, nyligen koloniserade populationer och i tio populationer av ett intermediärt successionsstadium. Unga populationer hade lägre genetisk diversitet jämfört med äldre populationer, vilket indikerar att genetiska flaskhalsar, skapade av fåtal antal koloniserande individer, s.k. founders, som härrör från ett begränsat antal källpopulationer, minskar den genetiska diversiteten inom nybildade populationer. Observationen av stark genetisk strukturering, mellan och inom öar, indikerar att genflödet är begränsat över flera rumsliga nivåer i detta system. Bristen på statistiskt signifikanta skillnader i genetisk differentiering mellan unga och intermediära populationer indikerar emellertid att nivåer av genflöde kanske inte är tillräckligt höga för att minska den genetiska differentieringen som uppstår från den ursprungliga founder-händelsen. Mönstren av sexuell dimorfism och hanarnas och honornas roll har utvecklats för att möjliggöra fortlevnad i ett ekologisk och populationsmässigt sammanhang hos Silene dioica. I denna livsmiljö, där öar stiger upp ur havet och skapar nya miljöer för kolonisering samtidigt som autogena primära successionsprocesser leder till utrotning, måste S. dioica kontinuerligt sprida sig mellan olika successionsfaser för att överleva. Den uppenbara framgången för den här dioika växten är uppenbar eftersom den är en av de få dominerande arterna i lövkanten. Detta tyder på att nivåer av fröspridning och genflöde är tillräckliga för att möjliggöra etablering och beständighet av ö-populationer och att de sexuella dimorfismer som har utvecklats i detta metapopulationssystem verkar för att öka nivåerna av genflöde. "Lev hårt – dö ung" -strategin med omfattande blomningar som vi finner hos hanarna kan ha utvecklats som ett sätt att upprätthålla tillräckliga nivåer av genetisk diversitet i metapopulationen men den kommer endast att vara en möjlig strategi om det finns kontinuerliga möjligheter för re-etableringar. Således kan den kontinuerliga landupphöjningen som förekommer i norra delen av Bottniska viken mycket väl vara en förutsättning för den långsiktiga beständigheten av denna dioika, fleråriga växtart. / <p>Felaktigt angivet "Dissertation for PhD" i kolofon.</p>

L'idée que les souches de rongeurs de laboratoire ne soient pas des modèles idéaux pour la recherche sur le vieillissement n’est pas nouvelle. Pourtant, l’attitude des chercheurs face à l’introduction d’animaux de type sauvage dans leurs travaux demeure frileuse bien que ces derniers apporteraient une solution adéquate pour l’étude d’un processus aussi complexe et multifactoriel que le vieillissement et permettraient d’intégrer les traits d’histoire de vie des individus afin de rendre compte de façon pertinente des changements liés à l’âge. Chez la souris glaneuse, le report de l’âge de première reproduction des animaux juvéniles hivernants conduit à la création de deux cohortes. Ces deux groupes d’animaux devront, à un âge plus ou moins avancé, explorer des environnements anxiogènes et faire face à des compétiteurs lors de leur dispersion, se reproduire et élever leur progéniture qui devra à son tour disperser. L’objectif de cette thèse est d’étudier les conséquences émotionnelles et sociales du vieillissement qui pourraient influencer les stratégies comportementales de ce rongeur de type sauvage. Ce travail s’ouvre également sur des thématiques d’actualité dans le domaine de la biogérontologie. Notre étude montre qu’à l’instar de celles réalisées chez l’Homme, le niveau d’anxiété d’un individu peut être déterminé par son âge mais aussi par l’âge de ses parents. L’âge de mise en couple va quant à lui influencer certaines stratégies liées à la reproduction comme la latence d’accouplement ou l’effort parental fourni par les mères et les pères et va conditionner l’apparition des premiers signes de sénescence reproductive. Enfin, cette étude dévoile que la durée de vie reproductive d’un couple monogame pourrait être un facteur renforçateur des liens qui unissent un mâle à sa partenaire. / The idea that standard laboratory rodents may not be an ideal model for aging research is not new. Nonetheless, the researcher's attitude toward using wild-type species remains cautious although these animals would make a suitable solution to study a process as complex and multifaceted as aging, and would allow to incorporate the life history traits of individuals to reflect appropriately age-related changes. In the mound-building mouse, the delay of the age of first reproduction of over wintering juvenile animals leads to the establishment of two cohorts. These two groups of animals have to, at more or less advanced ages, explore anxiogenic environnements and face competitors during dispersal, reproduce and raise their offspring wich will in turn disperse. The aim of this thesis is to study the emotional and social consequences of aging that could influence the behavioral strategies in this wild-type rodent focusing on current issues in the biogerontogical field. Our study shows that, like in humans, the anxiety level of an individual can be determined by its own age but also by the age of its parents at conception. For its part, the age at pairing influences reproduction-related strategies as the latency of first reproduction or parental effort provided by mothers and fathers and affects the appearence of the first signs of reproductive senescence. Finally, this work reveals that the duration of pairing could strengthen the social bond between a male and its partner in a monogamous species.