Hólar University College

The ability to predict evolutionary responses to environmental change is of central importance for future conservation efforts. Increases in temperature from anthropogenic climate change are arguably the most widespread and dramatic alteration of natural ecosystems, with its impact predicted to become more acute with time. These increases in temperature are likely to shift species distributions, or drive developmental and evolutionary changes in local populations. Therefore, knowing how currently extant populations can or have changed in response to warmer conditions would be extremely important. Here we propose to investigate how populations change with respect to their growth, physiology, and anatomy in response to long term exposure to increased temperature. We will also determine what genetic changes may have happened in addition to epigenetic changes (i.e. environmentally-induced structural alterations of the genome that affect its function but don't change the DNA sequence), and relate these to phenotypic variation. To address this we will use a series of unique study sites in Iceland where populations of threespine stickleback live both in geothermally heated ponds and in unheated (i.e. cool) ponds in close proximity. These pairs of adjacent populations (which occur in several places) have lived at these contrasting temperatures for thousands of generations and preliminary investigation shows evidence of evolutionary divergence between them. This broad-ranging project will use both comparative and experimental approaches to quantify how the phenotype, genome, and epigenome interact when the animal experiences rising temperatures. Lab experiments will use fish from these source populations to investigate 1) population level differences between ambient (cool) and 'warmed' populations at the genomic and epigenomic level, 2) the inheritance of phenotypic traits (including metabolic rate, morphology and locomotor performance) and epigenomic states in sticklebacks reared under reciprocal temperature regimes, 3) alterations of development under warm conditions that could limit evolution, and 4) the genetic and epigenetic basis for variation in phenotypic traits under different temperatures. This project will be holistic through its ability to allow plasticity and heritable epigenetic effects to be assessed together with genetic variation. It will reveal whether there are consistent (and hence predictable) evolutionary and epigenetic changes when populations are exposed to warmer temperatures, and will explore the role of plasticity in shaping adaptive responses. As a result, it will provide novel information on the extent to which populations can adapt to a warming world.

Most human disorders are complex and involve a multitude of genes, environmental inputs, and change in prevalence with age. To understand such disorders clinical researchers often use a specific set of lab animals (model organisms) that display similarity to human disease. However, while this approach has provided major advances for understanding some of the causal mutations underlying human disease there can be limitations in terms of how translatable results are. This is often attributed to differences between humans and other animals but it may also be the result of the approach used. Normally, lab-based research on such model organisms standardizes environmental conditions, as well as genetic variation within a lab line. This can simplify comparisons with an 'all else being equal' approach that allows for single mutations to be compared (i.e. this is often referred to as the mutant model approach). However, as mentioned above, many diseases manifest themselves from a multitude of genes, environments, and age. An emerging alternative to the traditional mutant model approach is now arriving from nature where in some cases evolutionary adaptations can actually resemble human disease. What is particularly exciting in that such populations can be more in line with how humans live, in that they experience environmental variation, and often have a diversity of genetic variation. Thus, they could be particularly effective for understanding complex human disease. This project will take advantage of a natural set of populations of fish, specifically threespine sticklebacks experiencing warmed habitats as a result of geothermal activity in Iceland. Such warmed habitats present a unique challenge to these fish as the higher temperature raises their metabolism, including in winter when prey are limited. This appears to have caused 'energy-sparing' adaptations to evolve in these fish, such as increased fat deposition, indications of glucose tolerance, and higher appetites. This can be considered to resemble metabolic syndrome in humans, but it is unclear what underlying mechanisms determine these changes, and whether a wider suite of traits are involved. To determine the utility of this system as a model for human disease will require deeper investigation, but could be especially valuable if these fish have also evolved the mitigate the negative effects of these traits. Therefore, we will aim to ascertain how the body composition of these fish is determined using a comprehensive approach that accounts for genetic, epigenetic, and environmental cues. This approach should more closely match how traits are determined in nature and lead to accurate insights about their mechanisms. We will also assess further aspects of the metabolic divergence between geothermal and ambient populations of stickleback, including glucose and insulin tolerance. Lastly, we will examine whether the expected negative effects of metabolic syndrome occur with associated traits in sticklebacks. Specifically, we will look for signs of non-alcoholic fatty liver disease, and test whether blood concentrations of triglycerides and cholesterol occur in geothermal fish (which would be expected to occur with higher levels of body fat). We predict that geothermal fish will show signs of mitigating these negative effects, and if supported it could provide the basis for further insight and even therapies for humans.

The studies of ecology and evolution are closely related. Ecologists seek to understand the environmental factors that explain the distribution and abundance of species, while evolutionary biologists investigate the process of natural selection and the evolution that results, by examination of adaptation in phenotypes and genotypes. It is curious in these times of environmental change that one of the biggest gaps in our understanding of the natural world falls exactly at the intersection between ecology and evolution: we know less than we should about how the environment shapes the evolution of biodiversity. Although it is generally understood that the environment is the cause of adaptation, the links between them have seldom been explicitly explored. Many ecological studies do not consider how the environmental variation that they measure affects evolution, while many studies of evolution measure selection or adaptation without considering their environmental causes, concentrating instead on the consequences for evolution of what is genetically possible. Explicit study of the involvement of the environment in evolution has the potential to fuel a paradigm shift in our comprehension of fundamental evolutionary patterns. For example: (i) Divergence. Evolution has resulted in abundant diversity in the natural world, but the extent of this divergence within related groups of organisms is often circumscribed. Are these limits, on the kind of organisms that evolve, a consequence of what is genetically possible, or do they result from similarities in the environments to which the organisms are exposed? (ii) Convergence. Within the greater divergence, organisms have often apparently converged on similar evolutionary solutions, suggesting that evolutionary outcomes are to some extent repeatable. Is the repeated evolution of similar organisms in different places the result of genetic biases or environmental determinants? If the latter, do similar organisms always evolve in similar environments, or can different environments favour the same outcome of organismal form? Vice versa, do similar environmental combinations always result in essentially the same organism, or are there different evolutionary solutions to similar environmental problems? (iii) Novelty. Although similar organisms in different places often converge on repeated evolutionary solutions, evolution also occasionally comes up with solutions that are different from the general pattern, by dint of developing, or having lost, some distinguishing feature or combination of features. Is such evolutionary novelty the result of particularly unusual environments? Most previous studies of how the environment affects evolution have measured only a single, or small number of aspects of both the organism and the environment, but thorough answers to the questions we pose require a more comprehensive understanding of multiple different aspects of organism and environment, and of how they interact and affect other. Our approach requires the use of recently developed multivariate statistical methods that allow the simultaneous analysis of many organismal traits and many environmental variables. Adaptive radiation is the differentiation of an ancestral species into divergent new populations or species. The abundance of variation in both environment and biodiversity make adaptive radiations the perfect natural laboratories to address our questions. We will use data from replicated adaptive radiations of three-spined stickleback fish in Scotland, Iceland, western Canada and Alaska in order to answer our questions and achieve a comprehensive understanding of how the environment affects evolution. Three-spined stickleback are originally marine fish that have invaded freshwater throughout the northern hemisphere since the last ice age. Freshwater stickleback can occupy contrasting environments and exhibit great phenotypic variation, providing a perfect system for our study.

Increasing global competition for natural resources poses major challenges to the Arctic. ArcticHubs will develop sustainable solutions for reconciliation of competing livelihoods and land-use modes in key Arctic ‘hubs’—important socioeconomic nodes in a geographical network—and their surroundings, considering in particular the needs and cultures of local communities (incl. indigenous people). This will be achieved by applying multi- and interdisciplinary, multi-actor participatory approaches to systematically map, identify and analyse global drivers and pressures with high environmental, societal and economic impacts affecting 33 key hubs examining sustainability of fish farming, multiple use of forests, tourism, mining and indigenous cultures. The outcome of ArcticHubs will be the provision of solution-oriented tools, such as improved public participatory geographical information systems, guidelines for ‘social license to operate’, and future scenarios to Arctic communities, industrial stakeholders, decision- and policymakers, and other relevant actors. This will enable creation and implementation of regional development strategies that reconcile new economic opportunities with traditional livelihoods, and increase the resilience of both new and existing industries and livelihoods against environmental, economic and political changes in the Arctic. The impact of the project will be long-term sustainability and resilience of future environmental, socio-cultural, economic and political factors in the increasingly competitive and globalised Arctic, enhancing social acceptance of increased economic activity. These impacts will contribute to the implementation of the new integrated EU policy for the Arctic, IPPC assessments and other major regional and global initiatives, provide support to the EU Arctic Research Cluster, and enhance engagement of and interaction between local (incl. indigenous), national and global actors.