xvi, 112 p. : ill. (some col.) / In seasonal environments, the ability to take advantage of the favorable seasons and avoid or mitigate the effects of the unfavorable ones is essential for organismal fitness. Many polar and temperate organisms use photoperiod (length of day) to time seasonal life history events because photoperiod's regular annual cycle makes it a very reliable indicator of seasonality. This reliability allows organisms to anticipate and properly prepare for seasonal change. Although photoperiodism is widespread in polar and temperate vertebrates, little is known relative to invertebrates regarding how its use varies with environment and this method's underlying genetic and physiological basis. This dissertation is focused on demonstrating the proper methodology for the study of photoperiodism and establishing the threespine stickleback as a model of vertebrate photoperiodism. Chapter I is an introduction to photoperiodism, how it is influenced by environment, the physiological basis of its output, and a summary of the chapters that follow. Chapter II explains an analytical method to test for causality and applies this method to data that have been interpreted as evidence that the circadian clock is causally involved in photoperiodism. Chapter III describes the photoperiodic response of threespine stickleback Gasterosteus aculeatus populations from two latitudes. These results are used to inform an empirical examination of the previously described assertion that the circadian clock is causally involved in photoperiodism. Chapter IV examines the physiological basis of early photoperiodic response using the threespine stickleback as a model teleost fish. Chapter V summarizes the previous chapters, describes their significance, and suggests future research directions. This dissertation includes both previously published and co-authored material. Supplementary Excel files demonstrating the analyses used in Chapter III are also included in this dissertation. / Committee in charge: Eric Johnson, Chairperson; William Cresko, Advisor; William Bradshaw, Member; Judith Eisen, Member; Patricia McDowell, Outside Member

Evolution shapes ecosystems but the processes by which this occurs are not well understood. Adaptive change in resource expensive traits may underlie one such process, as evolution altering a species’ resource needs may effect how that species interacts with ecosystem resources. For this, Ecological Stoichiometry (ES) may be a tractable framework, as it simplifies organisms into elemental ratios and then applies mass-balance to predict changes in diet and waste interactions. ES detects variation in resource expensive traits as variation in elemental ratios, and predicts compensation via parallel changes in diet (e.g. high phosphorous individuals consume high phosphorus diets) and/or offsetting changes in waste (e.g. high phosphorous individuals release low phosphorus waste). To test the utility of this framework and improve our understanding of eco-evolutionary dynamics, I studied variation in phenotypic traits, genetics, elemental content and resource interactions within and across natural populations of highly regarded eco-evolutionary model species threespine stickleback (Gasterosteus aculeatus). First, I related heritable variation in phosphorus rich bony traits and genetics commonly under natural selection with variation in elemental content (N:P) to determine the magnitude and basis of intraspecific variation in N:P. Second, I investigated the ecosystem consequences of variation in elemental content by determining whether stickleback compensate through changes in diet choice and excretion rates. I found stickleback vary widely in elemental composition (3.0 – 9.4:1 N:P) which models explained well with four bone related traits: bone mineralization, body size, lateral plating and pelvis size (R2 > 0.52). Additional genetic models linked variation in Eda alleles (which underlie lateral plating) with a 12% shift in stickleback N:P. Stickleback compensated for this variation in N:P demand by altering diet choice rather than excretion rates, and by maximizing dietary inputs through changes in gut morphology. Within and across populations, high phosphorus stickleback consumed a larger proportion of high phosphorus prey and contained longer gastrointestinal tracts that more efficiency process diet resources. These results demonstrate that heritable variation in elemental composition is ecologically relevant with individual traits and genetics having large effects. As individuals compensated by altering resource acquisition rather than release, the direct ecological consequences of evolutionary change in these resource expensive traits is likely larger for food web structure and abundance than nutrient dynamics. / Graduate / 2018-12-19

As individuals and populations diverge ecologically, they become exposed to new parasites and pathogens with potentially harmful fitness consequences. Populations are therefore expected to evolve resistance, possibly at a cost of less resistance to parasites rarely encountered parasites. This trade-off in resistance should generate local adaptation to parasites in different habitats. In chapter one, I show how local adaptation can potentially evolve in response to variation in parasite exposure among eighteen ecologically variable populations of threespine stickleback (Gasterosteus aculeatus). Within populations infection appeared to reflect morphology/diet based exposure differences among individuals. Among populations, however, these patterns were absent or reversed, consistent with the evolution of local adaptation. In chapters two and three I set out to test whether variation major histocompatibility (MHC) genes can underly such local adaptation in stickleback. MHC genes are important components of vertebrate immunity; however, there is little direct empirical support for spatially divergent selection driving local adaptation on MHC loci in the wild. In chapter two I tested for the action of parasite mediated balancing and divergent selection on on MHC loci using naturally infected stickleback in three replicate lake-stream pairs. Despite consistent divergence in parasites and MHC alleles, lakes tended to show decreased parasite burdens with increased allelic richness (consistent with balancing selection), while streams showed some support for divergent selection between lake and stream types. In chapter three I use the same lake-stream pairs to investigate how divergent selection could instead be reflected in variation in the effects of individual MHC alleles among populations. When comparing parapatric populations experiencing gene flow, MHC alleles maintained at relatively high frequency in one population were more likely to be associated with reduced, rather than increased, parasite abundances in that population. Allopatric populations experiencing no gene flow showed no such general relationship between allele frequency and resistance. These results are only consistent with spatially divergent selection, and imply that gene flow and environmental heterogeneity can be important for maintaining MHC diversity. / text

The seasonal and diel feeding habits of the goosander, Mergus merganser, the red-breasted merganser, M. serrator, gulls (Larus canus, L. argentatus and L. fuscusj and terns, Sterna hirundo/paradisaea were studied at 64V05'N. Birds' activity patterns were influenced by the nocturnal spawning of the river lamprey (Lampetra fluviatilis). Food selection and food consumption by hand-raised mergansers together with aquarium studies of the diel activity patterns of their most important prey supplemented the field data. River lamprey dominated the diet of the goosander by weight and the sculpin Cottus gobio by number. The fish consumption of the goosanders was found to be about 12% of the available river lamprey biomass and about 17% of the sculpin biomass during the breeding season. In experimental situations the river lamprey was a low- pritority species compared with salmon , Salmo salar, brown trout, 53. trutta, and minnow, Phoxinus phoxinus, when presented to satiated birds of both Mergus species. When hungry, however, the birds caught the available prey irrespective of species, but they selected larger prey when two size classes were present. Experimental results were compared with field data on availability, consumption and the escape behaviour of the various fish species. The rivers emptying in the Bothnian Bay are regarded as important feeding areas for birds breeding along the coast. In early spring they get access to abundant and reliable food resources, e.g the river lamprey. Later on the three-spined stickleback, Gasterosteus aculeatus, supply both Mergus species with food in the river and also along the coast. From the middle of June potential food supply available to birds decreases in the rivers and becomes more abundant in the coastal area. / <p>S. 1-41: sammanfattning, s. 43-227: 7 uppsatser</p> / digitalisering@umu

Reproduction in vertebrates, including fishes, is under control of the brain-pituitary-gonad (BPG) axis. The female three-spined stickleback, Gasterosteus aculeatus, produces egg clutches at intervals of a few days and spawns them in a nest built by male. Following ovulation, eggs are stored in the ovarian cavity surrounded by the ovarian fluid (OF). If spawning or spontaneous release do not occur, the eggs can undergo overripening, a phenomenon occurring both in nature and captivity. In this PhD thesis, the changes of reproductive hormones and vitellogenesis were studied at overripening of eggs and over the natural spawning cycle. OF properties were also examined at overripening of eggs and after treatment with sex steroids. Plasma levels of steroids: testosterone (T), estradiol (E2), 17,20β-dihydroxypregn-4-en-3-one (17,20β-P) and 17,20β,21-trihydroxypregn-4-en-3-one (17,20β,21-P) were measured by radioimmunoassay, and relative mRNA levels of the pituitary gonadotropins (fsh-β/lh-β), brain gonadotropin-releasing hormones (gnrh2/gnrh3) and kisspeptin and its receptor (kiss2/gpr54) by qPCR. Overripening of eggs was accompanied with a significant reduction in most of endocrine parameters of BPG axis (T, E2, 17,20β-P; lh-β; kiss2, gpr54). Low level of hormones could be advantageous for the overripe egg-bound females, since this would reduce further ovulations giving higher chances to survive and reproduce again. Over the 3-day spawning cycle, T and E2 were highly correlated, showed cyclicity with low levels at ovulation and increasing from 24 and 6 hours post-spawning (hps), respectively. Spawning may give rise to this increase as these rises did not occur if release of the eggs does not happen (overripe females). A peak at pituitary lh-β mRNA levels appeared 48 hps, a day before the next ovulation. No significant changes were found for the other studied hormones. Vitellogenesis was studied by measurement of the vitellogenin mRNA levels in the liver by qPCR. The levels were highest at 24 and 48 hps and were positively correlated to both E2 and T over the cycle. However, changes were small suggesting a rather continuous vitellogenesis over the stickleback spawning cycle which could be an advantage for a multiple spawner with a limited spawning season. Overripening reduced vitellogenin mRNA levels but did not abolish it. OF amount was diminished in overripe females and had a lower viscocity but higher dry weight and protein levels than in non-overripe ovulated females, suggesting that changes in OF properties are related to the egg overripening. The effects of steroids were studied using Silastic capsules. T and 17,20β-P induced an increase of OF amount, but protein levels were only increased in 17,20β-P-treated females, proposing a role of this steroid in the control of OF secretion. 1-D SDS-PAGE showed that OF contained several proteins, some of them came from eggs, but no consistent differences between groups. Concluding, the knowledge of the reproductive physiological changes is important for understanding their essential roles in the production of viable eggs in this species but also in the reproductive physiology of female fishes in general. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: Manuscript. Paper 4: Manuscript.</p>

The populations of three–spined sticklebacks (Gasterosteus aculeatus) in the Baltic Sea have increased tenfold over the last decade. A large increase in abundance can alter the offshore and coastal food webs. Despite of these facts, there are large gaps in the knowledge about the stickleback ecology in the Baltic and the possible effect they might have on their environment. Earlier investigations state that stickleback mainly occupy the deeper areas offshore, and only migrate to the shallow areas during May–July to spawn. Observations by recreational fishermen indicate that this may be incorrect, and that some adult sticklebacks are present in the shallow areas even during the winter. One aim of this study was to investigate the timing of stickleback migration to a shallow coastal area in Kalmar Sound. The study also aimed to examine the relative abundance in two adjacent shallow areas in the archipelago south of Kalmar, where one of the areas is a pike spawning ground. A one month long test fishing with fyke nets was started on the first day after ice break. Results show that the sticklebacks are present in the bays immediately after the ice break, and that high abundances coincide with the pike spawning period. Stomach analyses showed that sticklebacks consumed a large proportion of crustaceans, but also fish eggs were found. These results shed new light on the management actions for many of the coastal spring spawning fish species that have shown decreasing abundances during the last decades. / Bestånden av storspigg (Gasterosteus aculeatus) i Östersjön har ökat markant, data visar på en tiofaldig ökning under det senaste decenniet. Ökningen kan medföra att bl.a. näringsväven, till havs och längs kusterna förändras. Trots vetskapen om detta finns det kunskapsluckor kring spiggens ekologi och dess möjliga påverkan på sin omgivning. Litteratur säger att spiggen till största del bara befinner sig inne längs kusterna under maj-juli. Observationer från sportfiskare tyder på att detta inte stämmer, och att vuxen storspigg befinner sig inne längs kusten och i skärgårdsvikar året om. Ett syfte med studien var att undersöka när spiggen kom in till de grunda vikarna i Kalmarsund. Studien jämförde även spiggtätheterna mellan två närliggande områden där den största skillnaden var födan. Ett månadslångt provfiske inleddes den första isfria dagen i två skyddade vikar söder om Kalmar, där en av lokalerna var en dokumenterad leklokal för gäddor. Resultaten visade att spiggen fanns i vikarna direkt efter islossningen, och vid fisket sammanföll de största spiggfångsterna med gäddleken. Maganalyser visade att en stor andel av födan bestod av olika kräftdjur, men även romkorn hittades. Resultaten tyder på andra förutsättningar för många av de hårt ansatta vårlekande fiskarterna än vad som tidigare är dokumenterat.

Throughout the animal kingdom, individuals often differ consistently from one another in how they cope with their environment. In particular, consistent behavioural variation, known as animal personality, is a substantial driver of a range of important ecological and evolutionary processes. As most animal species are social for at least part of their lives and group living is common, a crucial link between personality and the social context may be expected. In this thesis I systematically investigate this link, using three-spined stickleback fish (Gasterosteus aculeatus) as my model system. I begin by showing that fish vary consistently in their boldness and sociability, with only boldness being positively linked to food intake for fish at rest. This finding provides support for evolutionary theory that links personality variation to life-history strategies, and lays the basis for work related to the social context. I continue by investigating how the social context may modulate personality variation and show that short contact with a social group may have carry-over effects and obfuscate personality expression when individuals are alone. Next, I observed fish in different pairs over time and found that social experience from both the current as well as previous social contexts are integrated in the risk-taking and leadership decisions of individuals but also depends on their boldness type. This result provides support for the importance of social feedback in the expression of personality differences. I go on to demonstrate that, in a pair, bolder fish have lower social attraction, with positive effects on individual’s leadership but negative effects on social coordination. Finally, by detailed tracking of the collective movements and group foraging of free-swimming shoals, I reveal boldness and sociability have complementary driving effects of on social structure, collective behaviour, and group functioning. Furthermore, I show that in turn the group composition determines the performance of individual personality types, providing a potential adaptive explanation for the maintenance of personality variation. Taken together, these studies provide an integrated account of animal personality and the social context and highlight the presence of a feedback loop between them, with personality variation being a key driver of collective behaviour and group functioning but also strongly affected and potentially maintained by it.