Inbreeding depression and partitioning of genetic load in the invasive biennial Alliaria petiolata (Brassicaceae).

PREMISE OF THE STUDY: Invasive species are nonnative species that enter novel environments, establish sustained populations, and can negatively impact native species. Here we assess a potential weakness of invasive species (genetic load) and show how species might overcome genetic barriers. Colonization of novel habitats by invasive species typically involves few individuals, exposing populations to founder effects. We empirically tested a central Illinois population of an invasive biennial plant, Alliaria petiolata, for evidence of a founder effect by assessing the pattern of genetic load. • METHODS: To estimate genetic load, we assayed offspring from three cross types (self-pollinated, outcrossed within- and between-populations) in a greenhouse. Vegetative and reproductive traits were measured on first-year plants grown with or without intraspecific competition. • KEY RESULTS: We found substantial genetic load in this population of A. petiolata, which can mostly be attributed to genetic drift (founder effect) and not inbreeding depression. Between-population heterosis was expressed more than inbreeding depression under intraspecific competition. • CONCLUSIONS: Inbreeding may be adaptive for A. petiolata in its introduced range by providing reproductive assurance, with limited inbreeding load. Nevertheless, most of the genetic load in this population of A. petiolata is due to fixation of deleterious alleles. Drift load is expected, given that this population is near the edge of its continuous geographic range in highly fragmented habitats, and gene flow between isolated populations is likely highly limited. Preventing additional introduction and movement of propagules between isolated local populations should reduce heterosis and A. petiolata competitiveness.

The scale of population structure in Arabidopsis thaliana.

The population structure of an organism reflects its evolutionary history and influences its evolutionary trajectory. It constrains the combination of genetic diversity and reveals patterns of past gene flow. Understanding it is a prerequisite for detecting genomic regions under selection, predicting the effect of population disturbances, or modeling gene flow. This paper examines the detailed global population structure of Arabidopsis thaliana. Using a set of 5,707 plants collected from around the globe and genotyped at 149 SNPs, we show that while A. thaliana as a species self-fertilizes 97% of the time, there is considerable variation among local groups. This level of outcrossing greatly limits observed heterozygosity but is sufficient to generate considerable local haplotypic diversity. We also find that in its native Eurasian range A. thaliana exhibits continuous isolation by distance at every geographic scale without natural breaks corresponding to classical notions of populations. By contrast, in North America, where it exists as an exotic species, A. thaliana exhibits little or no population structure at a continental scale but local isolation by distance that extends hundreds of km. This suggests a pattern for the development of isolation by distance that can establish itself shortly after an organism fills a new habitat range. It also raises questions about the general applicability of many standard population genetics models. Any model based on discrete clusters of interchangeable individuals will be an uneasy fit to organisms like A. thaliana which exhibit continuous isolation by distance on many scales.

Studying plant-pollinator interactions in a changing climate: A review of approaches.

Plant-pollinator interactions are potentially at risk due to climate change. Because of the spatial and temporal variation associated with the effects of climate change and the responses of both actors, research to assess this interaction requires creative approaches. This review focuses on assessments of plants' and pollinators' altered phenology in response to environmental changes, as phenology is one of the key responses. I reviewed research methods with the goal of presenting the wide diversity of available techniques for addressing changes in these interactions. Approaches ranged from use of historical specimens to multisite experimental community studies; while differing in depth of historical information and community interactions, all contribute to assessment of phenology changes. Particularly insightful were those studies that directly assessed the environmental changes across spatial and temporal scales and the responses of plants and pollinators at these scales. Longer-term studies across environmental gradients, potentially with reciprocal transplants, enable an assessment of climate impacts at both scales. While changes in phenology are well studied, the impacts of phenology changes are not. Future research should include approaches to address this gap.

Studying plant-pollinator interactions facing climate change and changing environments.

Plant-pollinator interactions are essential for successful plant reproduction in both natural and agricultural systems. These interactions are negatively impacted by recent large-scale alterations of the environments, particularly climate change. The responses of plants and pollinators to changing abiotic conditions that vary seasonally and geographically are often uncoordinated, potentially leading to the breakdown of this interaction. The complexity of the responses of plants and pollinators to our changing climate necessitates creative approaches. The six articles in this special issue directly address this need by providing a variety of key methods and reviews of current methodology. The articles include: DNA barcoding methods for use on pollen collected from visiting bees; methods for assessment of plant attraction traits (nectar and review of floral volatiles methods); a field sampling method for ground nesting bees; a review of using spatial and temporal transplants for addressing changing dynamics of plant-pollinator interactions; and a review of approaches used to assess potential shifts in phenology of plants and pollinators. Collectively, these articles illustrate some of the breadth of approaches needed to address the changing dynamics of plant-pollinator interactions.

Methamphetamine-induced neurotoxicity disrupts pharmacologically evoked dopamine transients in the dorsomedial and dorsolateral striatum.

Phasic dopamine (DA) signaling, during which burst firing by DA neurons generates short-lived elevations in extracellular DA in terminal fields called DA transients, is implicated in reinforcement learning. Disrupted phasic DA signaling is proposed to link DA depletions and cognitive-behavioral impairment in methamphetamine (METH)-induced neurotoxicity. Here, we further investigated this disruption by assessing effects of METH pretreatment on DA transients elicited by a drug cocktail of raclopride, a D2 DA receptor antagonist, and nomifensine, an inhibitor of the dopamine transporter (DAT). One advantage of this approach is that pharmacological activation provides a large, high-quality data set of transients elicited by endogenous burst firing of DA neurons for analysis of regional differences and neurotoxicity. These pharmacologically evoked DA transients were measured in the dorsomedial (DM) and dorsolateral (DL) striatum of urethane-anesthetized rats by fast-scan cyclic voltammetry. Electrically evoked DA levels were also recorded to quantify DA release and uptake, and DAT binding was determined by means of autoradiography to index DA denervation. Pharmacologically evoked DA transients in intact animals exhibited a greater amplitude and frequency and shorter duration in the DM compared to the DL striatum, despite similar pre- and post-drug assessments of DA release and uptake in both sub-regions as determined from the electrically evoked DA signals. METH pretreatment reduced transient activity. The most prominent effect of METH pretreatment on transients across striatal sub-region was decreased amplitude, which mirrored decreased DAT binding and was accompanied by decreased DA release. Overall, these results identify marked intrastriatal differences in the activity of DA transients that appear independent of presynaptic mechanisms for DA release and uptake and further support disrupted phasic DA signaling mediated by decreased DA release in rats with METH-induced neurotoxicity.

Evolution in heterogeneous environments and the potential of maintenance of genetic variation in traits of adaptive significance.

The maintenance of genetic variation in traits of adaptive significance has been a major dilemma of evolutionary biology. Considering the pattern of increased genetic variation associated with environmental clines and heterogeneous environments, selection in heterogeneous environments has been proposed to facilitate the maintenance of genetic variation. Some models examining whether genetic variation can be maintained, in heterogeneous environments are reviewed. Genetic mechanisms that constrain evolution in quantitative genetic traits indicate that genetic variation can be maintained but when is not clear. Furthermore, no comprehensive models have been developed, likely due to the genetic and environmental complexity of this issue. Therefore, I have suggested two empirical approaches to provide insight for future theoretical and empirical research. Traditional path analysis has been a very powerful approach for understanding phenotypic selection. However, it requires substantial information on the biology of the study system to construct a causal model and alternatives. Exploratory path analysis is a data driven approach that uses the statistical relationships in the data to construct a set of models. For example, it can be used for understanding phenotypic selection in different environments, where there is no prior information to develop path models in the different environments. Data from Brassica rapa grown in different nutrients indicated that selection changed in the different environments. Experimental evolutionary studies will provide direct tests as to when genetic variation is maintained.

VARIATION IN SEED CHARACTERS IN NEMOPHILA MENZIESII: EVIDENCE OF A GENETIC BASIS FOR MATERNAL EFFECT.

A growing body of evidence indicates that phenotypic selection on juvenile traits of both plants and animals may be considerable. Because juvenile traits are typically subject to maternal effects and often have low heritabilities, adaptive responses to natural selection on these traits may seem unlikely. To determine the potential for evolutionary response to selection on juvenile traits of Nemophila menziesii (Hydrophyllaceae), we conducted two quantitative genetic studies. A reciprocal factorial cross, involving 16 parents and 1960 progeny, demonstrated a significant maternal component of variance in seed mass and additive genetic component of variance in germination time. This experiment also suggested that interaction between parents, though small, provides highly significant contributions to the variance of both traits. Such a parental interaction could arise by diverse mechanisms, including dependence of nuclear gene expression on cytoplasmic genotype, but the design of this experiment could not distinguish this from other possible causes, such as effects on progeny phenotype of interaction between the environmental conditions of both parents. The second experiment, spanning three generations with over 11,000 observations, was designed for investigation of the additive genetic variance in maternal effect, assessment of paternal effects, as well as further partitioning of the parental interaction identified in the reciprocal factorial experiment. It yielded no consistent evidence of paternal effects on seed mass, nor of parental interactions. Our inference of such interaction effects from the first experiment was evidently an artifact of failing to account for the substantial variance among fruits within crosses. The maternal effect was found to have a large additive genetic component, accounting for at least 20% of the variation in individual seed mass. This result suggests that there is appreciable potential for response to selection on seed mass through evolution of the maternal effect. We discuss aspects that may nevertheless limit response to individual selection on seed mass, including trade-offs between the size of individual seeds and germination time and between the number of seeds a maternal plant can mature and their mean size.