Plasticity to wind is modular and genetically variable in Arabidopsis thaliana

Thigmomorphogenesis, the characteristic phenotypic changes by which plants react to mechanical stress, is a widespread and probably adaptive type of phenotypic plasticity. However, little is known about its genetic basis and population variation. Here, we examine genetic variation for thigmomorphogenesis within and among natural populations of the model system Arabidopsis thaliana. Offspring from 17 field-collected European populations was subjected to three levels of mechanical stress exerted by wind. Overall, plants were remarkably tolerant to mechanical stress. Even high wind speed did not significantly alter the correlation structure among phenotypic traits. However, wind significantly affected plant growth and phenology, and there was genetic variation for some aspects of plasticity to wind among A. thaliana populations. Our most interesting finding was that phenotypic traits were organized into three distinct and to a large degree statistically independent covariance modules associated with plant size, phenology, and growth form, respectively. These phenotypic modules differed in their responsiveness to wind, in the degree of genetic variability for plasticity, and in the extent to which plasticity affected fitness. It is likely, therefore, that thigmomorphogenesis in this species evolves quasi-independently in different phenotypic module

Experimental alteration of DNA methylation affects the phenotypic plasticity of ecologically relevant traits in Arabidopsis thaliana

Heritable phenotypic variation in plants can be caused not only by underlying genetic differences, but also by variation in epigenetic modifications such as DNA methylation. However, we still know very little about how relevant such epigenetic variation is to the ecology and evolution of natural populations. We conducted a greenhouse experiment in which we treated a set of natural genotypes of Arabidopsis thaliana with the demethylating agent 5-azacytidine and examined the consequences of this treatment for plant traits and their phenotypic plasticity. Experimental demethylation strongly reduced the growth and fitness of plants and delayed their flowering, but the degree of this response varied significantly among genotypes. Differences in genotypes' responses to demethylation were only weakly related to their genetic relatedness, which is consistent with the idea that natural epigenetic variation is independent of genetic variation. Demethylation also altered patterns of phenotypic plasticity, as well as the amount of phenotypic variation observed among plant individuals and genotype means. We have demonstrated that epigenetic variation can have a dramatic impact on ecologically important plant traits and their variability, as well as on the fitness of plants and their ecological interactions. Epigenetic variation may thus be an overlooked factor in the evolutionary ecology of plant population

Navigating economic governance: German export and investment promotion between varieties of capitalism and multi-level governance

This thesis examines the German system of Export and Investment Promotion Agencies (EIPAs) using a combination of the Varieties of Capitalism (VOC) and Multi-Level Governance (MLG) approaches. It challenges the traditional application of VOC and using the MLG approach argues for an expansion of its theoretical framework to account for a hybrid type of actor organisation and governance interactions across multiple levels. Accordingly, this thesis examines the theoretical framework of both approaches before their utilisation. It then proposes to address their shortcomings by examining whether the VOC approach can be refined and further improved by combining it with aspects of MLG to better reflect the complex interactions between different state and non-state actors and their type or organisation within a country’s economy.This is accomplished in several ways. First, an in-depth case study of the Export and Investment Promotion Agencies (EIPA) sector in Germany on multiple, vertical levels of governance is conducted. To do so, a combination of 28 elite interviews and key primary sources such as financial documents, official program reports and parliamentary hearings were used. This is followed by a case within a case example examining Germany’s single largest state-funded program for export promotion, called the Export Initiative Renewable Energy (Exportinitiative Erneuerbare Energien).Overall, the thesis argues for a combined VOC and MLG approach and rejects VOC's traditional dualism. It instead recognises the possibility of a hybrid type of actor organisation within market economies. The analysis identified a new type of actor organisation, referred to as “coordinated competition”. This was based on emerging evidence showing that regional, federal and international level actors display complex and opportunistic behaviours which are best understood in the context of a hybrid type of VOC market system. The results also show the role of the state as a direct market actor on both a regional and federal level. These findings both challenge the traditional understanding of VOC and confirm several of the criticisms of the approach found in the academic literature. The case supports the argumentation that both VOC and MLG are valuable heuristic analytical devices which can benefit from an expansion of their theoretical frameworks.Based on the results of the analysis of the German EIPA system, this thesis addresses three main gaps in the academic literature. First, it provides a detailed analysis of the setup of the German EIPA system. Despite its size, history and economic significance, Germany’s EIPA system, along with EIPAs in general, has received little academic attention. This has resulted in them being somewhat of a ‘black box’ to academic research. The results of this thesis provide a rich understanding of the governance systems and actor behaviours that could add to the overall academic understanding of EIPAs as actors in the political economy literature. Second, it illustrates that a combined VOC and MLG approach should be applied to accurately understand the complex governance relationships and forms of actor organisation that shape this sector of the German economy. This is critically important, as it allows one to take into consideration and incorporate different levels of governance structures and spheres of influence that were missing in the VOC literature. Finally, based on the case analysis of Germany, it argues that the VOC approach should be expanded to incorporate a new, ideal-type hybrid category, in which actors are organised via a combination of Coordinated Market Environment (CME) and Liberal Market Environment (LME) characteristics. This new hybrid type of VOC allows for a more precise reflection and analysis of the complex actor relationships found within Germany’s export and investment promotion system

Epigenetic variation in plant responses to defence hormones

Background and Aims There is currently much speculation about the role of epigenetic variation as a determinant of heritable variation in ecologically important plant traits. However, we still know very little about the phenotypic consequences of epigenetic variation, in particular with regard to more complex traits related to biotic interactions. Methods Here, a test was carried out to determine whether variation in DNA methylation alone can cause heritable variation in plant growth responses to jasmonic acid and salicylic acid, two key hormones involved in induction of plant defences against herbivores and pathogens. In order to be able to ascribe phenotypic differences to epigenetic variation, the hormone responses were studied of epigenetic recombinant inbred lines (epiRILs) of Arabidopsis thaliana - lines that are highly variable at the level of DNA methylation but nearly identical at the level of DNA sequence. Key Results Significant heritable variation was found among epiRILs both in the means of phenotypic traits, including growth rate, and in the degree to which these responded to treatment with jasmonic acid and salicylic acid. Moreover, there was a positive epigenetic correlation between the responses of different epiRILs to the two hormones, suggesting that plant responses to herbivore and pathogen attack may have a similar molecular epigenetic basis. Conclusions This study demonstrates that epigenetic variation alone can cause heritable variation in, and thus potentially microevolution of, plant responses to defence hormones. This suggests that part of the variation of plant defences observed in natural populations may be due to underlying epigenetic, rather than entirely genetic, variatio

Phenotypic and genetic differentiation between native and introduced plant populations

Plant invasions often involve rapid evolutionary change. Founder effects, hybridization, and adaptation to novel environments cause genetic differentiation between native and introduced populations and may contribute to the success of invaders. An influential idea in this context has been the Evolution of Increased Competitive Ability (EICA) hypothesis. It proposes that after enemy release plants rapidly evolve to be less defended but more competitive, thereby increasing plant vigour in introduced populations. To detect evolutionary change in invaders, comparative studies of native versus introduced populations are needed. Here, we review the current empirical evidence from: (1) comparisons of phenotypic variation in natural populations; (2) comparisons of molecular variation with neutral genetic markers; (3) comparisons of quantitative genetic variation in a common environment; and (4) comparisons of phenotypic plasticity across different environments. Field data suggest that increased vigour and reduced herbivory are common in introduced plant populations. In molecular studies, the genetic diversity of introduced populations was not consistently different from that of native populations. Multiple introductions of invasive plants appear to be the rule rather than the exception. In tests of the EICA hypothesis in a common environment, several found increased growth or decreased resistance in introduced populations. However, few provided a full test of the EICA hypothesis by addressing growth and defence in the same species. Overall, there is reasonable empirical evidence to suggest that genetic differentiation through rapid evolutionary change is important in plant invasions. We discuss conceptual and methodological issues associated with cross-continental comparisons and make recommendations for future research. When testing for EICA, greater emphasis should be put on competitive ability and plant tolerance. Moreover, it is important to address evolutionary change in characteristics other than defence and growth that could play a role in plant invasion

Climate warming can reduce biocontrol efficacy and promote plant invasion due to both genetic and transient metabolomic changes.

Climate change may affect plant-herbivore interactions and their associated ecosystem functions. In an experimental evolution approach, we subjected replicated populations of the invasive Ambrosia artemisiifolia to a combination of simulated warming and herbivory by a potential biocontrol beetle. We tracked genomic and metabolomic changes across generations in field populations and assessed plant offspring phenotypes in a common environment. Using an integrated Bayesian model, we show that increased offspring biomass in response to warming arose through changes in the genetic composition of populations. In contrast, increased resistance to herbivory arose through a shift in plant metabolomic profiles without genetic changes, most likely by transgenerational induction of defences. Importantly, while increased resistance was costly at ambient temperatures, warming removed this constraint and favoured both vigorous and better defended plants under biocontrol. Climate warming may thus decrease biocontrol efficiency and promote Ambrosia invasion, with potentially serious economic and health consequences

Climate warming can reduce biocontrol efficacy and promote plant invasion due to both genetic and transient metabolomic changes

Climate change may affect plant-herbivore interactions and their associated ecosystem functions. In an experimental evolution approach, we subjected replicated populations of the invasive Ambrosia artemisiifolia to a combination of simulated warming and herbivory by a potential biocontrol beetle. We tracked genomic and metabolomic changes across generations in field populations and assessed plant offspring phenotypes in a common environment. Using an integrated Bayesian model, we show that increased offspring biomass in response to warming arose through changes in the genetic composition of populations. In contrast, increased resistance to herbivory arose through a shift in plant metabolomic profiles without genetic changes, most likely by transgenerational induction of defences. Importantly, while increased resistance was costly at ambient temperatures, warming removed this constraint and favoured both vigorous and better defended plants under biocontrol. Climate warming may thus decrease biocontrol efficiency and promote Ambrosia invasion, with potentially serious economic and health consequences

Environment-Sensitive Epigenetics and the Heritability of Complex Diseases

Genome-wide association studies have thus far failed to explain the observed heritability of complex human diseases. This is referred to as the “missing heritability” problem. However, these analyses have usually neglected to consider a role for epigenetic variation, which has been associated with many human diseases. We extend models of epigenetic inheritance to investigate whether environment-sensitive epigenetic modifications of DNA might explain observed patterns of familial aggregation. We find that variation in epigenetic state and environmental state can result in highly heritable phenotypes through a combination of epigenetic and environmental inheritance. These two inheritance processes together can produce familial covariances significantly higher than those predicted by models of purely epigenetic inheritance and similar to those expected from genetic effects. The results suggest that epigenetic variation, inherited both directly and through shared environmental effects, may make a key contribution to the missing heritability

Fine-scale population epigenetic structure in relation to gastrointestinal parasite load in red grouse (Lagopus lagopus scotica)

Acknowledgements This study was funded by a BBSRC studentship (MA Wenzel) and NERC grants NE/H00775X/1 and NE/D000602/1 (SB Piertney). The authors are grateful to Mario Röder and Keliya Bai for fieldwork assistance; Alex Douglas for statistical advice; Tyler Stevenson, Heather Ritchie and three anonymous reviewers for helpful comments on manuscript drafts; and all estate owners, factors and keepers for access to field sites, most particularly MJ Taylor and Mike Nisbet (Airlie), Neil Brown (Allargue), RR Gledson and David Scrimgeour (Delnadamph), Andrew Salvesen and John Hay (Dinnet), Stuart Young and Derek Calder (Edinglassie), Kirsty Donald and David Busfield (Glen Dye), Neil Hogbin and Ab Taylor (Glen Muick), Alistair Mitchell (Glenlivet), Simon Blackett, Jim Davidson and Liam Donald (Invercauld) Richard Cooke and Fred Taylor (Invermark), Shaila Rao and Christopher Murphy (Mar Lodge), and Ralph Peters and Philip Astor (Tillypronie).Peer reviewedPublisher PD

Invasion of Europe by the western corn rootworm, Diabrotica virgifera virgifera: multiple transatlantic introductions with various reductions of genetic diversity

The early stages of invasion involve demographic bottlenecks that may result in lower genetic variation in introduced populations as compared to source population/s. Low genetic variability may decrease the adaptive potential of such populations in their new environments. Previous population genetic studies of invasive species have reported varying levels of losses of genetic variability in comparisons of source and invasive populations. However, intraspecific comparisons are required to assess more thoroughly the repeatability of genetic consequences of colonization events. Descriptions of invasive species for which multiple introductions from a single source population have been demonstrated may be particularly informative. The western corn rootworm (WCR), Diabrotica virgifera virgifera, native to North America and invasive in Europe, offers us an opportunity to analyse multiple introduction events within a single species. We investigated within- and between-population variation at eight microsatellite markers in WCR in North America and Europe to investigate the routes by which WCR was introduced into Europe, and to assess the effect of introduction events on genetic variation. We detected five independent introduction events from the northern USA into Europe. The diversity loss following these introductions differed considerably between events, suggesting substantial variation in introduction, foundation and/or establishment conditions. Genetic variability at evolutionarily neutral loci does not seem to underlie the invasive success of WCR in Europe. We also showed that the introduction of WCR into Europe resulted in the redistribution of genetic variance from the intra- to the interpopulational level contrary to most examples of multiple introductions