The extent to which a species' environmental range reflects adaptive differentiation remains an open question. Environmental gradients can lead to adaptive divergence when differences in stressors ...among sites along the gradient place conflicting demands on the balance of stress responses. The extent to which this is accomplished through stress tolerance vs stress avoidance is also an open question.
We present results from a controlled environment study of 48 lineages of Arabidopsis thaliana collected along a gradient in northeastern Spain across which temperatures increase and precipitation decreases with decreasing elevation. We tested the extent to which clinal adaptive divergence in heat and drought is explained through tolerance and avoidance traits by subjecting plants to a dynamic growth chamber cycle of increasing heat and drought stress analogous to low elevation spring in northeastern Spain.
Lineages collected at low elevation were the most fit and fitness scaled with elevation of origin. Higher fitness was associated with earlier bolting, greater early allocation to increased numbers of inflorescences, reduction in rosette leaf photosynthesis and earlier fruit ripening.
We propose that this is a syndrome of avoidance through early flowering accompanied by restructuring of the organism that adapts A. thaliana to low-elevation Mediterranean climates.
Widespread specimen digitization has greatly enhanced the use of herbarium data in scientific research. Publications using herbarium data have increased exponentially over the last century. Here, we ...review changing uses of herbaria through time with a computational text analysis of 13,702 articles from 1923 to 2017 that quantitatively complements traditional review approaches. Although maintaining its core contribution to taxonomic knowledge, herbarium use has diversified from a few dominant research topics a century ago (e.g., taxonomic notes, botanical history, local observations), with many topics only recently emerging (e.g., biodiversity informatics, global change biology, DNA analyses). Specimens are now appreciated as temporally and spatially extensive sources of genotypic, phenotypic, and biogeographic data. Specimens are increasingly used in ways that influence our ability to steward future biodiversity. As we enter the Anthropocene, herbaria have likewise entered a new era with enhanced scientific, educational, and societal relevance.
Although adaptive plasticity would seem always to be favored by selection, it occurs less often than expected. This lack of ubiquity suggests that there must be trade-offs, costs, or limitations ...associated with plasticity. Yet, few costs have been found. We explore one type of limitation, a correlation between plasticity and developmental instability, and use quantitative genetic theory to show why one should expect a genetic correlation. We test that hypothesis using the Landsberg erecta × Cape Verde Islands recombinant inbred lines (RILs) of Arabidopsis thaliana. RILs were grown at four different nitrogen (N) supply levels that span the range of N availabilities previously documented in North American field populations. We found a significant multivariate relationship between the cross-environment trait plasticity and the within-environment, within-RIL developmental instability across 13 traits. This genetic covariation between plasticity and developmental instability has two costs. First, theory predicts diminished fitness for highly plastic lines under stabilizing selection, because their developmental instability and variance around the optimum phenotype will be greater compared to nonplastic genotypes. Second, empirically the most plastic traits exhibited heritabilities reduced by 57% on average compared to nonplastic traits. This demonstration of potential costs in inclusive fitness and heritability provoke a rethinking of the evolutionary role of plasticity.
Background: Understanding the relationship between environment and genetics requires the integration of knowledge on the demographic behavior of natural populations. However, the demographic ...performance and genetic composition of Arabidopsis thaliana populations in the species' native environments remain largely uncharacterized. This information, in combination with the advances on the study of gene function, will improve our understanding on the genetic mechanisms underlying adaptive evolution in A. thaliana. Methodology/Principal Findings: We report the extent of environmental, demographic, and genetic variation among 10 A. thaliana populations from Mediterranean (coastal) and Pyrenean (montane) native environments in northeast Spain. Geographic, climatic, landscape, and soil data were compared. Demographic traits, including the dynamics of the soil seed bank and the attributes of aboveground individuals followed over a complete season, were also analyzed. Genetic data based on genome-wide SNP markers were used to describe genetic diversity, differentiation, and structure. Coastal and montane populations significantly differed in terms of environmental, demographic, and genetic characteristics. Montane populations, at higher altitude and farther from the sea, are exposed to colder winters and prolonged spring moisture compared to coastal populations. Montane populations showed stronger secondary seed dormancy, higher seedling/juvenile mortality in winter, and initiated flowering later than coastal populations. Montane and coastal regions were genetically differentiated, montane populations bearing lower genetic diversity than coastal ones. No significant isolation-by-distance pattern and no shared multilocus genotypes among populations were detected. Conclusions/Significance: Between-region variation in climatic patterns can account for differences in demographic traits, such as secondary seed dormancy, plant mortality, and recruitment, between coastal and montane A. thaliana populations. In addition, differences in plant mortality can partly account for differences in the genetic composition of coastal and montane populations. This study shows how the interplay between variation in environmental, demographic, and genetic parameters may operate in natural A. thaliana populations.
Early life-history transitions are crucial determinants of lifetime survival and fecundity. Adaptive evolution in early life-history traits involves a complex interplay between the developing plant ...and its current and future environments. We examined the plant's earliest life-history traits, dissecting an integrated suite of pregermination processes: primary dormancy, thermal induction of secondary dormancy, and seasonal germination response. We examined genetic variation in the three processes, genetic correlations among the processes, and the scaling of germination phenology with the source populations' climates. A spring annual life history was associated with genetic propensities toward both strong primary dormancy and heat-induced secondary dormancy, alone or in combination. Lineages with similar proportions of winter and spring annual life history have both weak primary dormancy and weak thermal dormancy induction. A genetic bias to adopt a spring annual strategy, mediated by rapid loss of primary dormancy and high thermal dormancy induction, is associated with a climatic gradient characterized by increasing temperature in summer and rainfall in winter. This study highlights the importance of considering combinations of multiple genetically based traits along a climatic gradient as adaptive strategies differentiating annual plant life-history strategies. Despite the genetic-climatic cline, there is polymorphism for life-history strategies within populations, classically interpreted as bet hedging in an unpredictable world.
• Understanding the adaptive basis of life history variation is a central goal in evolutionary ecology. The use of model species enables the combination of molecular mechanistic knowledge with ...ecological and evolutionary questions, but the study of life history variation in natural environments is required to merge these disciplines. • Here, we tested for clinal variation in life history and associated traits along an environmental and altitudinal gradient in the model species Arabidopsis thaliana. Seventeen natural populations of A. thaliana were geo-referenced in north-eastern Spain on a gradient in which precipitation increases but maximum spring temperature and minimum winter temperature decrease with altitude. • One hundred and eighty-nine genotypes from the 17 populations were grown under uniform controlled conditions. Variations in traits related to biomass allocation, fecundity, phenology and vegetative growth were tested for relationships with the altitude and climatic variables associated with the home sites. Above-ground mass, number of rosette leaves at bolting, developmental time and seed weight increased with the home site's altitude. Root allocation, vegetative growth during winter and number of seeds decreased with altitude. • We suggest that the differences among home sites provide clues to the variation in adaptive strategies associated with the climatic gradient. We compared these results with adaptations and clinal relationships reported for other species and with molecular mechanisms described in Arabidopsis.
Explaining the diversity in geographic range sizes among species is a central goal of ecological and evolutionary studies. We tested species age as an explanation of range size variation within a ...group of understory shrubs in the Neotropics (Psychotria subgenus Psychotria, Rubiaceae). We distinguish between range occupancy (filling an occupied area) and range extent (maximum distances dispersed). We used Bayesian relaxed-clock dating of molecular sequence data to estimate the relative age of species, and we used species distribution modeling to predict species' potential ranges. If the range sizes of species are limited by time for dispersal, we hypothesize that older species should have (1) larger realized range occupancies and realized range extents than younger species, (2) filled a greater proportion of their potential range occupancies, and (3) colonized a greater proportion of their potential range extents. We found (1) a significant but weak positive relationship between species age versus both realized range occupancy and realized range extent, (2) no relationship between species age and filling of potential range occupancies, but (3) that older species had colonized a significantly greater proportion of their potential range extents than younger species. Our results indicate that a time-for-dispersal effect can limit the extent of ranges of species but not necessarily their occupancies.
Exactly 50 years ago, a revolution in empirical population genetics began with the introduction of methods for detecting allelic variation using protein electrophoresis (Throckmorton 1962; Hubby ...1963; Lewontin & Hubby 1966). These pioneering scientists showed that populations are chock‐full of genetic variation. This variation was a surprise that required a re‐thinking of evolutionary genetic heuristics. Understanding the causes for the maintenance of this variation became and remains a major area of research. In the process of addressing the causes, this same group of scientists documented geographical genetic structure (Prakash et al. 1969), spawning the continued accumulation of what is now a huge case study catalogue of geographical differentiation (e.g. Loveless & Hamrick 1984; Linhart & Grant 1996). Geographical differentiation is clearly quite common. Yet, a truly general understanding of the patterns in and causes of spatial genetic structure across the genome remains elusive. To what extent is spatial structure driven by drift and phylogeography vs. geographical differences in environmental sources of selection? What proportion of the genome participates? A general understanding requires range‐wide data on spatial patterning of variation across the entire genome. In this issue of Molecular Ecology, Lasky et al. (2012) make important strides towards addressing these issues, taking advantage of three contemporary revolutions in evolutionary biology. Two are technological: high‐throughput sequencing and burgeoning computational power. One is cultural: open access to data from the community of scientists and especially data sets that result from large collaborative efforts. Together, these developments may at last put answers within reach.
•We find that wild Arabidopsis genotypes hyperaccumulate camalexin, a major defense chemical against bacterial infection.•Lines with hyperaccumulation of camalexin were more resistant to bacterial ...infection.•Camalexin production was dramatically upregulated in all plants following infection.•We report the first evidence of a significant tradeoff between high inducibility and high constitutive production of camalexin, as predicted by optimal defense theory.
Optimal defense theory predicts that induction of defensive secondary metabolites in plants will be inversely correlated with constitutive expression of those compounds. Here, we asked whether camalexin, an important defense against fungal and bacterial pathogens, support this prediction in structured natural populations of Arabidopsis thaliana from the Iberian Peninsula. In common garden experiments, we found that genotypes from the VIE population constitutively hyper-accumulated camalexin. Camalexin concentrations were not induced significantly when plants were exposed to a temperature of 10°C for 48h. However, they were induced when plants were exposed to 48h of infection by the virulent bacterial pathogen, Pseudomonas syringae pv. tomato DC3000. Genotypes from the VIE population with the hyper-accumulation of camalexin were significantly more resistant to bacterial growth. Induction of camalexin was negatively correlated with constitutive camalexin concentrations following log transformation and two different corrections for autocorrelation, thus supporting the tradeoff predicted by optimal defense theory. Constitutive overexpression of camalexin was not explained by the only known natural genetic polymorphism at the Accelerated Cell Death 6, ACD6, locus. Collectively, the results support an important role of camalexin in defense against P. syringae as well as significant structured variation in defense levels within wild populations.
A metamorphosis from rosette to inflorescence in many annuals shifts photosynthetic tissue from a two-dimensional array in the soil boundary layer during cool months to a three-dimensional structure ...in the troposphere as spring progresses. We propose that this shift allows escape from both self-shading and an increasingly stressful boundary layer microclimate, permitting continued increases in growth. As a first step in exploring this hypothesis, we compared the lifetime C gain, water loss, and instantaneous water use efficiency (WUE) of five Arabidopsis thaliana genotypes by measuring gas exchange across the life cycle. On average, the inflorescence contributed 55% (± 5% SE) of lifetime C gain, but only 25% of lifetime water loss. Mean inflorescence WUE was nearly fourfold that of the rosette. The inflorescence continued to fix C after rosette senescence. The percentage inflorescence: total C gain varied among genotypes, from 36% to 93%. Genotypes differed in WUE for both structures. We suggest that local climates may have selected for divergence in these traits. For many annuals and winter annuals, understanding C and water budgets and their evolution must include measures of both rosette and inflorescence gas exchange.
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