| 1. |
- Aalberg Haugen, Inger Marie, 1980-
(author)
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The diapause switch : Evolution of alternative developmental pathways in a butterfly
- 2014
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Doctoral thesis (other academic/artistic)abstract
- Diapause decision is a classic example of a threshold switch mechanism with cascading effects on morphology, behaviour and life-history traits. This thesis addresses the downstream effects of the insect diapause switch, with the main focus on pathway-specific regulation of life-history traits, using the speckled wood butterfly (Pararge aegeria) as a study species. The ultimate pathway decision is made towards the end of larval development and allows the larvae to take into account up-to-date information from the environment about future conditions (Paper I, IV). However, already from an early point in development the larvae are sensitive to environmental cues and continuously adjust their growth trajectory in accordance to current information about the environmental conditions to be expected in future (Paper IV). An asymmetry in the ability to change from one developmental pathway to another at a late point in larval development suggests that the diapause and the direct pathway require different physiological preparations (Paper IV). Pathway-specific regulation of traits downstream of the diapause switch is maintained by ongoing selection. When the direct pathway is not regularly expressed, as with a shift from bivoltinism to univoltinism, relaxed selection on the unexpressed pathway leads to genetic drift and loss of protandry (Paper II, III). Natural populations display local adaptations in the diapause switch with an increase in critical daylengths as there is a gradual shift from bivoltinism to univoltinism (Paper III). This thesis highlights two aspects of the diapause decision, the determination of how and when this decision is made as well as the way the resulting pathways are moulded by selection in order to produce adaptive seasonal polyphenism in life-history traits.
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| 2. |
- Karlsson, Magnus
(author)
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Evolution in changing environments revealed by fire melanism in pygmy grasshopper
- 2010
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Doctoral thesis (other academic/artistic)abstract
- According to theory, genetic diversity can be maintained by environmental variation and the degree of genetic and phenotypic polymorphism may enhance the ability of populations to endure stress imposed by changing environments. In my thesis I used colour polymorphic pygmy grasshoppers (Tetrix subulata) as a model system to explore how environmental variation influenced genetic diversity. I compared population colour morph frequencies between populations in burnt and non-burnt areas and performed experiments to investigate to what extent colour patterns in these insects are determined by genes and influenced by phenotypic plasticity in response to environmental effects experienced during development. My results showed that the frequency of black individuals on average was much higher in recently fire ravaged areas, a condition known as fire melanism. The highest proportion of black individuals was reached within the first year after a fire. After the initial increase, the proportion of black individuals declined again and the distribution among alternative colour morphs became more even. Data for individuals raised in captivity revealed a high correspondence between maternal and offspring colour patterns, indicating a strong genetic influence on colour. Additional experiments demonstrated that the development of colour patterns in pygmy grasshoppers was not influenced by burnt material or high population densities, two environmental cues associated with post fire environments.To test if reduced competition among alternative colour morphs may contribute to the maintenance of colour pattern polymorphism in these insects I examined if average survival was higher in diverse compared to homogeneous groups of individuals. I found that survival increased with colour pattern diversity, presumably due to reduced competition among alternative colour morphs. Relaxation of competition may explain why the distribution among alternative colour patterns changed and became more even after the initial evolution of fire melanism. My results demonstrate that environmental change may cause extremely rapid and reversible evolution, indicate that fluctuating selection may preserve genetic variation and support the notion that polymorphism may increase average individual success and enable populations to withstand environmental change.
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| 3. |
- Lindestad, Olle, 1988-
(author)
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Geographic variation in life cycles : Local adaptation and ecological genetics in a temperate butterfly
- 2020
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Doctoral thesis (other academic/artistic)abstract
- Conditions in nature change with the seasons, necessitating seasonal adaptations that synchronize the life cycles of organisms with their surroundings. Such regulatory adaptations must vary between populations to track local variation in climate and seasonality; this local adaptation is facilitated by locally specific seasonal cues, but may be hampered by gene flow and genetic history. For populations of temperate insects, two central features of adaptation to local climate are voltinism, the yearly number of generations; and diapause, the state of arrested development and suppressed metabolism in which most temperate insects spend winter. Delaying diapause allows for an additional generation to be produced within the same year, but this is only adaptive if the season is sufficiently long to safely accommodate such a life cycle. Hence, selection to express a locally adaptive voltinism should drive divergence between populations in diapause regulation and associated life history traits. In this thesis, I investigate variation in voltinism and life cycle regulation in a set of populations of the butterfly Pararge aegeria. Population-level variation in seasonal plasticity was tested in two sets of experiments. The first (Paper I) focused on photoperiodic plasticity during the growing season, and revealed considerable differences between populations in diapause induction and developmental reaction norms. Mechanistic modeling based on the laboratory results indicated that differences in voltinism are actively maintained by these genetic differences. Next, I tested the idea that shorter diapause may help populations achieve higher voltinism through earlier emergence in the spring (Paper II). This idea was not supported; instead, populations differed in a manner that suggests that diapause duration is selected upon by the need to avoid premature development under warm autumn conditions. The genetic background of seasonal adaptation in these populations was also explored. Phylogeographic structures inferred from genome-wide data put the results of the laboratory experiments into a historic context, and were used as the basis for a scan for genetic loci showing signs of differential selection (Paper III). The scan revealed novel variation in two circadian genes that have been shown to be linked to diapause control in P. aegeria, including a large deletion in the gene timeless. Finally, a test of two previously described circadian mutations (Paper IV) showed that, while these mutations may affect photoperiodic plasticity on a between-population level, they seemingly have no effect within a single population located at intermediate latitudes. Closer inspection revealed novel, locally unique mutations in the same genes, possibly compensating for the effect of diapause-delaying variants in a setting where an attempted second generation is not adaptive. I have shown that voltinism variation in P. aegeria is enabled by population differences in seasonal plasticity, with population differences playing a greater role during some parts of the year than others. These results present voltinism as a complex trait resulting from plasticity acting at different levels of geographic specificity. Although much of the genetic variation enabling the observed local adaptation remains uncharacterized, the considerably variable circadian genes seen in these populations provide an intriguing target for future investigation.
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| 4. |
- Nilsson-Örtman, Viktor, 1984-
(author)
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Thermal adaptation along a latitudinal gradient in damselflies
- 2012
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Doctoral thesis (other academic/artistic)abstract
- Understanding how temperature affects biological systems is a central question in ecology and evolutionary biology. Anthropogenic climate change adds urgency to this topic, as the demise or success of species under climate change is expected to depend on how temperature affects important aspects of organismal performance, such as growth, development, survival and reproduction. Rates of biological processes generally increase with increasing temperature up to some maximal temperature. Variation in the slope of the initial, rising phase has attracted considerable interest and forms the focus of this thesis. I explore variation in growth rate-temperature relationships over several levels of biological organization, both between and within species, over individuals’ lifetime, depending on the ecological context and in relation to important life history characteristics such as generation length and winter dormancy. Specifically, I examine how a clade of temperate damselflies have adapted to their thermal environment along a 3,600 km long latitudinal transect spanning from Southern Spain to Northern Sweden. For each of six species, I sampled populations from close to the northern and southern range margin, as well from the center of the latitudinal range. I reared larvae in the laboratory at several temperatures in order to measure indiviudal growth rates. Very few studies of thermal adaptation have employed such an extensive sampling approach, and my finding reveal variation in temperature responses at several levels of organization. My main finding was that temperature responses became steeper with increasing latitude, both between species but also between latitudinal populations of the same species. Additional genetic studies revealed that this trend was maintained despite strong gene flow. I highlight the need to use more refined characterizations of latitudinal temperature clines in order to explain these findings. I also show that species differ in their ability to acclimate to novel conditions during ontogeny, and propose that this may reflect a cost-benefit trade-off driven by whether seasonal transitions occur rapidly or gradually during ontogeny. I also carried out a microcosm experiment, where two of the six species were reared either separately or together, to determine the interacting effects of temperature and competition on larval growth rates and population size structure. The results revealed that the effects of competition can be strong enough to completely overcome the rate-depressing effects of low temperatures. I also found that competition had stronger effects on the amount of variation in growth rates than on the average value. In summary, my thesis offers several novel insights into how temperature affects biological systems, from individuals to populations and across species’ ranges. I also show how it is possible to refine our hypotheses about thermal adaptation by considering the interacting effects of ecology, life history and environmental variation.
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| 5. |
- Pruisscher, Peter, 1988-
(author)
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Functional genomics of diapause in two temperate butterflies
- 2019
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Doctoral thesis (other academic/artistic)abstract
- Natural selection will act on a given phenotype to maximize fitness in a particular environment, even if this would result in reduced fitness in other environments. In insects some of the strongest selection pressures act on timing life cycles to seasonal variation in environmental conditions, in order to maximize growth, reproduction, and to anticipate the onset of winter. Many temperate insects survive winter by entering a pre-programmed state of developmental arrest, called diapause. The decision to induce diapause is predominantly based on measuring day length. Populations have adapted to latitudinal variation in photoperiod, thereby synchronizing with local seasonal variation. However, there is no general understanding of the genetic basis for controlling diapause induction, maintenance and termination. In this thesis I aimed to gain a better understanding of the genetic basis underlying variation in the induction decision, as well as to gain insights into gene expression changes during diapause in temperate butterflies. I started by revealing local adaptation in the photoperiodic response of two divergent populations of Pieris napi (Paper I). I found that variation in diapause induction among populations of both P. napi and Pararge aegeria showed strong sex-linked inheritance in inter-population crosses (Paper I and II). The genome-wide variation across populations was relatively low in both species. However, there was strong divergence in genomic regions containing the circadian clock genes timeless and period in P. aegeria, and period, cycle, and clock in P. napi. The genetic variation in these specific regions segregated between diapausing and direct developing individuals of inter-population crosses, showing that allelic variation at few genes with known functions in the circadian clock correlated to variation in diapause induction (Paper II and III). Furthermore, I investigated the transcriptional dynamics in two tissues (head and abdomen) during diapause (Paper IV). Already at the first day of pupal development there are on average 409 differentially expressed genes (DEG) each up and down regulated between the direct development and diapause pathways, and this increases dramatically across these formative stages to an average of 2695. Moreover, gene expression is highly dynamic during diapause, showing more than 2600 DEG’s in the first month of diapause development, but only 20 DEG’s in the third month. Moreover, gene expression is independent of environmental conditions, revealing a pre-programmed transcriptional landscape that is active during the winter. Still, adults emerging from either the direct or diapause pathways do not show large transcriptomic differences, suggesting the adult phenotype is strongly canalized. Thus, by integrating whole-genome scans with targeted genotyping and bulk-segregant analyses in population crosses, I demonstrate that adaptive variation in seasonal life cycle regulation in the two butterflies P. napi and P. aegeria both converge on genes of the circadian clock, suggesting convergent evolution in these distantly related butterflies.Moreover, the diapause program is a dynamic process with a distinct transcriptional profile in comparison to direct development, showing that on a transcriptome level diapause development and direct development are two distinct developmental strategies.
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| 6. |
- Stålhandske, Sandra, 1986-
(author)
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Spring Phenology of Butterflies : The role of seasonal variation in life-cycle regulation
- 2016
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Doctoral thesis (other academic/artistic)abstract
- Animals and plants in temperate regions must adapt their life cycle to pronounced seasonal variation. The research effort that has gone into studying these cyclical life history events, or phenological traits, has increased greatly in recent decades. As phenological traits are often correlated to temperature, they are relevant to study in terms of understanding the effect of short term environmental variation as well as long term climate change. Because of this, changes in phenology are the most obvious and among the most commonly reported responses to climate change. Moreover, phenological traits are important for fitness as they determine the biotic and abiotic environment an individual encounters. Fine-tuning of phenology allows for synchronisation at a local scale to mates, food resources and appropriate weather conditions. On a between-population scale, variation in phenology may reflect regional variation in climate. Such differences can not only give insights to life cycle adaptation, but also to how populations may respond to environmental change through time. This applies both on an ecological scale through phenotypic plasticity as well as an evolutionary scale through genetic adaptation. In this thesis I have used statistical and experimental methods to investigate both the larger geographical patterns as well as mechanisms of fine-tuning of phenology of several butterfly species. The main focus, however, is on the orange tip butterfly, Anthocharis cardamines, in Sweden and the United Kingdom. I show a contrasting effect of spring temperature and winter condition on spring phenology for three out of the five studied butterfly species. For A. cardamines there are population differences in traits responding to these environmental factors between and within Sweden and the UK that suggest adaptation to local environmental conditions. All populations show a strong negative plastic relationship between spring temperature and spring phenology, while the opposite is true for winter cold duration. Spring phenology is shifted earlier with increasing cold duration. The environmental variables show correlations, for example, during a warm year a short winter delays phenology while a warm spring speeds phenology up. Correlations between the environmental variables also occur through space, as the locations that have long winters also have cold springs. The combined effects of these two environmental variables cause a complex geographical pattern of phenology across the UK and Sweden. When predicting phenology with future climate change or interpreting larger geographical patterns one must therefore have a good enough understanding of how the phenology is controlled and take the relevant environmental factors in to account. In terms of the effect of phenological change, it should be discussed with regards to change in life cycle timing among interacting species. For example, the phenology of the host plants is important for A. cardamines fitness, and it is also the main determining factor for oviposition. In summary, this thesis shows that the broad geographical pattern of phenology of the butterflies is formed by counteracting environmental variables, but that there also are significant population differences that enable fine-tuning of phenology according to the seasonal progression and variation at the local scale.
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| 7. |
- Toftegaard, Tenna, 1982-
(author)
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Temperature and the synchrony of plant-insect interactions
- 2016
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Doctoral thesis (other academic/artistic)abstract
- Increasing temperatures resulting from climate change have within recent years been shown to advance phenological events in a large number of species worldwide. Species can differ in their response to increasing temperatures, and understanding the mechanisms that determine the response is therefore of great importance in order to understand and predict how a warming climate can influence both individual species, but also their interactions with each other and the environment. Understanding the mechanisms behind responses to increasing temperatures are however largely unexplored.The selected study system consisting of host plant species of the Brassicaceae family and their herbivore Anthocharis cardamines, is assumed to be especially vulnerable to climatic variations. Through the use of this study system, the aim of this thesis is to study differences in the effect of temperature on development to start of flowering within host plant species from different latitudinal regions (study I), and among host plant species (study II). We also investigate whether different developmental phases leading up to flowering differ in sensitivity to temperature (study II), and if small-scale climatic variation in spring temperature influence flowering phenology and interactions with A. cardamines (study III). Finally, we investigate if differences in the timing of A. cardamines relative to its host plants influence host species use and the selection of host individuals differing in phenology within populations (study IV).Our results showed that thermal reaction norms differ among regions along a latitudinal gradient, with the host plant species showing a mixture of co-, counter- and mixed gradient patterns (study I). We also showed that observed differences in the host plant species order of flowering among regions and years might be caused by both differences in the distribution of warm days during development and differences in the sensitivity to temperature in different phases of development (study II). In addition, we showed that small-scale variations in temperature led to variation in flowering phenology among and within populations of C. pratensis, impacting the interactions with the butterfly herbivore A. cardamines. Another result was that the less the mean plant development stage of a given plant species in the field deviated from the stage preferred by the butterfly for oviposition, the more used was the species as a host by the butterfly (study IV). Finally, we showed that the later seasonal appearance of the butterflies relative to their host plants, the higher butterfly preference for host plant individuals with a later phenology, corresponding to a preference for host plants in earlier development stages (study IV).For our study system, this thesis suggest that climate change will lead to changes in the interactions between host plants and herbivore, but that differences in phenology among host plants combined with changes in host species use of the herbivore might buffer the herbivore against negative effects of climate change. Our work highlights the need to understand the mechanisms behind differences in the responses of developmental rates to temperature between interacting species, as well as the need to account for differences in temperature response for interacting organisms from different latitudinal origins and during different developmental phases in order to understand and predict the consequences of climate change.
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