There is growing concern that the worldwide loss in biodiversity will harm the stability of the ecosystems, and thereby, the carrying capacity and critical ecosystem services the biosphere provides. Phytoplankton (microalgae) in lakes and oceans are arguably the most important primary producers. They are responsible for approximately half of the earth's primary production. However, there is little research into what influences the biodiversity of phytoplankton communities and of those studies the mechanisms for coexistence of phytoplankton have so far almost exclusively been studied on ecological time scales. We, therefore, explored how biodiversity and biomass (a proxy to primary production) of phytoplankton communities respond to co-varied environmental drivers over evolutionary time scales. We model adaptive evolution of phytoplankton' resource use, with a non-dimensionalized model of negatively buoyant phytoplankton inhabiting a partially mixed one-dimensional water column using reaction-advection-diffusion equations. We show that a number of environmental drivers have novel effects on biodiversity and biomass on evolutionary timescales. In contrast with previous non-evolutionary work, we found that decreasing light attenuation or increasing resource use efficiency can result in decreased biomass of plankton communities and nutrient-poor environments. One novel driver of species diversity was the combination of low rates of diffusion with relatively intermediate rates of sinking promote species diversity. Furthermore, we show that the phytoplankton turnover rate affects environmental heterogeneity and is, therefore, a contributing driver to species diversity.The evolution of half saturation constants can produce a variety of biodiversity-ecosystem function patterns as two positive, one unimodal, and one negative association were found when comparing biodiversity-ecosystem function. Collectively, our analyses suggest that environmental drivers can have substantially different effects over evolutionary timescales than those effects ecological modeling has previously shown.

Panarthropoda is a monophyletic group comprised of arthropods and lobopods, molting animals with a segmented body, paired appendages, dorsal brain, and ventral nerve cords. Evolutionary Developmental Biology (EvoDevo) is an interdisciplinary field that seeks to understand how changes in development form the basis for variations in morphology and phenotypic evolution, including the genetic network underlying these processes. To study the evolution of panarthropods from such an EvoDevo perspective, one typically uses standard molecular techniques. A first step here is to investigate the expression of a gene of interest in order to find out where and when it is transcribed during development. A hallmark of EvoDevo studies is its comparative character, often with respect to model organisms such as the fruit fly Drosophila melanogaster. Recently developed single-cell RNA sequencing technologies allow the profiling of a plethora of gene expression on the level of individual cells, and thus provide a much more detailed insight into gene expression. In Paper I, I applied standard molecular techniques used in EvoDevo research such as PCR, gene cloning, probe synthesis and whole mount in situ hybridization, to investigate the embryonic expression patterns of the tiptop/teashirt (tio/tsh) and spalt (sal) genes in a range of arthropods representing all main groups of this phylum, and an onychophoran. In the arthropod model Drosophila, these genes act as trunk-specifiers, and the objective of my work was to find out if this is conserved in Arthropoda or even Panarthropoda as a whole. I provide comprehensive data on arthropod tio/tsh and sal expression, including the first data from an onychophoran. The results support the idea that tio/tsh genes are involved in the development of ‘trunk’ segments by regulating limb development. In addition, my data suggest that the function of Sal is unlikely to be conserved in trunk vs head development. Early expression of sal, however, is in line with a potential homeotic function of this gene, at least in Arthropoda. In Paper II, I provide an embryonic tissue dissociation protocol for embryos of the common house spider Parasteatoda tepidariorum that I developed and that I successfully applied for single-cell RNA sequencing. In addition, I report on the progress of this experiment, and provide and discuss preliminary results.

Gymnasieelevers förståelse av naturligt urval och fylogenetiska träd har undersökts genom en enkätundersökning med 223 elever på fyra gymnasieskolor i Sverige. Resultatet från studien visar att elever oftare svarar rätt på flervalsfrågor med evolutionära träd än på frågor som behandlar naturligt urval. Det skulle kunna vara för att de frågor (och svar) som tar upp naturligt urval har mer text, där den relevanta informationen måste kunna tas ut först från frågan och sedan från svarsalternativen för att kunna välja det rätta alternativet. Resultatet visar även att det inte finns något samband mellan elevernas förståelse av evolutionära träd och naturligt urval. Eleverna kan alltså förstå det ena (evolutionära träd) men behöver inte nödvändigtvis förstå det andra (naturligt urval) och vice versa. / High school students understanding of natural selection and phylogenetic trees has been tested through a survey of 223 pupils in four upper secondary schools in Sweden. The results of the study show that students more often choose the correct answer on multiple choice questions with evolutionary trees than on questions that address natural selection. One reason for this could be that the questions (and answers) that address natural selection have more text, where the students must be able to take out the relevant information first from the question and then from the answers, in order to be able to choose the correct answer. The result also shows that there is no connection between the students understanding of evolutionary trees and natural selection. The pupils can thus understand one (evolutionary trees) but do not necessarily need to understand the other (natural selection) and vice versa.

Denna studie handlar om elevers kunskap om evolutionära mekanismer och hur biologilärare utformar sin pedagogik för att stärka elevers begreppsmässiga förståelse. Studien består av fem semistrukturerade intervjuer med gymnasielärare i biologi vid två svenska gymnasieskolor. Studien lägger särskilt fokus på active learning-pedagogik och undersöker huruvida active learning kan främja elevers förståelse för evolutionära kärnbegrepp (variation, selektion och ärftlighet). Studiens resultat indikerar att de vanligaste missförstånden är att individer kan anpassa sig för att överleva, att begreppet fitness biologiska innebörd misstolkas samt en bristande förståelse för begreppet selektionstryck. Studien sätter förståelsen för evolutionära kärnbegrepp i relation till så kallade tröskelbegrepp som är utgör viktig förförståelse samt kognitiva biaser vilka utgör vanliga hinder för att kunna ta till sig kunskap om evolutionära mekanismer. Studien kontextualiserar active learning med närliggande didaktiska teoribildningar såsom Piagets begrepp ackommodera och det övre stegen i Blooms taxonomi. Ett resultat är att de lärare som själva har ett intresse av evolutionsundervisning i större utsträckning utvärderar sin undervisning och aktivt söker nya, mer illustrativa laborationer och övningar. Eftersom undervisningens utformning skiljer sig betydligt lärare emellan föreslås skapande av ett forum där evidensbaserade active learning- övningar kan tillgängliggöras för alla biologilärare. En active learning-laboration som baserat på forskning visat goda resultat beskrivs i detalj och jämförs ingående med en laboration genomförd av tre av de 5 intervjuade lärarna. Båda dessa laborationer utgör exempel på active learning-pedagogik, vilket skulle kunna utgöra en källa till inspiration som biologilärare kan använda för vidareutveckling av sin egen undervisning.

The human gut microbiome is a complex ecosystem with hundreds of species interacting with each other and the host. One function of the microbiome is to break down undigested nutrients into smaller nutrients, sometimes available for uptake by the host. The digestion of such macromolecules can involve several species where one feeds on another’s byproducts, forming a large cross-feeding network. The method of digital evolution can be of great aid in studying such complex ecosystems by creating models of the studied system. In this study, the digital evolution software Avida was used to study the effects of perturbations in the system on byproduct cross-feeding chain length. Intense perturbations were found to shorten the chain lengths in general whereas weaker perturbations had either a small or no effect. When perturbations ceased, most byproduct chains displayed recovery to lengths similar to the preperturbation lengths. This indicates that byproduct chain lengths may be kept short by common ecological mechanisms alone, explaining why very long chains are rarely observed while still theoretically possible.

Selection pressure is a constant force in evolution, pushing birds and their wings towards an optimal shape and structure, were increasing performance, and minimizing the costs is central. But even though the science of aerodynamics can provide calculations of the optimal construction for the wing in different situations this rarely directly correspond to what is observed in nature. Since the optima are not the same for all birds due to different specifications and ecology this optimum becomes harder to determine and different functions can even have different optima, resulting in selection conflict. In the genius of birds there is an immense variation between species and their wings in everything from size, shape, and function.  The aim of this study is to investigate how wing morphology over a large phylogeny of bird species correlates to migration and habitat/ecology. Many studies have been done focusing on the effect of migration on the wing morphology, and some have been done focusing on other parameters such as display or daily usage. But by including the bird’s ecology and habitat related information with migration and morphology and looking at the selection from a broad perspective, can we uncover something more? The morphology of the wing cannot provide a perfect optimum for all circumstances since they require different specifications. What then, has the largest impact on the wing’s morphology? And does the relative length of the tail provide any correlations with its habitat and performance?  1185 birds of 137 species were caught at Ottenby, Öland and information on age, weight, sex, and tail length was collected for each individual bird. Photographs were taken of the back of the bird with the left wing outstretched 90 degrees from the body and analyzed in ImageJ to calculate aspect ratio and wing loading. Data on migration distance, foraging behavior, diet, and habitat density was then added for each of the species. Mean values of all parameters was calculated on species level creating a strong dataset with 137 data points. The species mean values dataset was used to test the interspecific effects and the dataset with all individuals was used to test intraspecific effects. ANOVAs, ANCOVAs, correlations tests and random slopes mixed models were performed revealing significant connections between wing morphology, migration, and habitat density. Correlations could also be observed between wing morphology, diet, and foraging behavior. Habitat density revealed the highest correlation with wing morphology, demonstrating a greater significance than migration and the other parameters. Effects that at first sight looked significant could later be excluded as they turned out to be dependent on other variables. The study therefore also highlights the importance of including alternative parameters for reliable conclusions.

No description available.

Selection and genetic drift are often competing forces in shaping genetic structure in populations. Genetic drift will often effectively cancel out the effect of selection when population sizes are small, such as in colonizing island populations. On a small island in the Skeppsvik Archipelago in northern Sweden, a newly founded population of Silene dioica has been monitored since it first established around 1993. Though inhabiting an area of merely 173 m2, the population has been shown to exhibit a genetically differentiated patch structure where closely related individuals are tightly grouped, distanced from other family groups. In this study, the effect of selection was evaluated as compared to that of genetic drift. Variation in phenotypic traits in flowers, leaves and stalks were compared to that of neutral markers, in the form of PST and FST measures, to assess a measure of what proportion of differentiation among patches in phenotypic traits could not be attributed to genetic drift. Males and females were analysed separately to obtain measures of sex specific selection. Signs of divergent and stabilizing selection were found in several traits in both males and females despite the small spatial scale and short time since colonization. Further analysis is needed to assess explanations for trait divergence among patches and direction of selection.

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>

Competition over access for mates has led to the evolution of many striking examples of morphological traits and behaviour in animals. The rapid development of the sexual selection field in recent decades have dramatically advanced our understanding of what traits make individuals more successful in attracting mates and how preferences for mates evolve over time. However, till now, research in this field has put less emphasis on the mechanisms that underlie variation in mate choice and sexual behaviour. Cognitive processes could potentially be key drivers of individual variation in mating preferences and sexual behaviours and therefore deserve further investigation. In this thesis, I used guppies artificially selected for relative brain size as the model system to study the association between brain size, cognitive ability and various aspects of mate choice. Previous studies in this model system showed that large-brained individuals of both sexes outperformed small-brained individuals in cognitive tests. Here I quantified their sexual behaviours and mating preferences to provide novel empirical data concerning the association between brain size, cognitive ability and sexual selection. In dichotomous choice preference tests based on visual cues, comparisons between large-brained and small-brained guppies showed important differences in their assessment of mate quality. These results are not driven by pre-existing visual biases caused by the artificial selection since further investigation of the visual capacity of these fish detected no differences between large-brained and small-brained individuals in their sensitivity to colour or in their capacity to resolve spatial detail. I also quantified sexual behaviour in male guppies artificially selected for relative brain size and found no difference in the behaviours of large-brained and small-brained males in a single male-single female non-competitive scenario. On the contrary, in a more complex social setting I found a reduction in large-brained males in the rate of courtship towards females and dominance displays towards other males when exposed to different degrees of predation threat and different numbers of male competitors. However, this reduction in behavioural intensity did not result in a lower access to copulation with females for large-brained males. I likewise evaluated female sexual behaviour and found that large-brained females had higher behavioural flexibility such that they decreased their receptiveness towards males more strongly under higher levels of predation threat. Together, these results provide novel empirical evidence that brain size and cognitive ability are tightly linked to mating preferences and sexual behaviours. These findings suggest that brain size and cognitive ability might be important mechanisms behind variation in mating preferences and in sexually selected traits across and within species. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 3: Manuscript. Paper 5: Manuscript.</p>