[show abstract][hide abstract] ABSTRACT: Understanding how genetic variation is organized over geography has long been of interest to evolutionary biologists given
that traits can vary within and among populations, across regions, and at continental or global scales. The pattern of regional
variation can have an important impact on trait evolution at the local or population level. Using a common garden, we asked
whether a geographically variable mosaic of tolerance to the widely applied herbicide RoundUp® existed in two closely related co-occurring species of morning glory, Ipomoea purpurea and I. hederacea. We assayed RoundUp tolerance in over 1,700 plants representing 290 families from 29 populations in the southeastern United
States. Our findings suggest that the two species of morning glory partition their respective levels of genetic variation
for tolerance to glyphosate differently. Variation for tolerance in I. purpurea appears to exist among maternal lines and regions, whereas in I. hederacea, variation in tolerance existed only among populations. In addition, we find a significant hotspot of tolerance or positive
spatial aggregation of this trait on a local scale in I. purpurea populations from the Coastal Plain. This suggests that either similar regimes of selection or gene flow between populations
can produce a geographic mosaic of tolerance. These results highlight the fact that the genetic variation underlying an adaptive
trait can exist at many different scales, whether it be within- or among-populations, among geographical ‘hotspots,’ or among
distinct ecological regions. Given these results, the partitioning of genetic variation should be considered before making
predictions about an adaptive trait’s evolutionary trajectory.
[show abstract][hide abstract] ABSTRACT: Reproductive barriers play a major role in the origin and maintenance of biodiversity by restricting gene flow between species. Although both pre- and postzygotic barriers often isolate species, prezygotic barriers are thought to contribute more to reproductive isolation. We investigated possible reproductive barriers between Leavenworthia alabamica and L. crassa, parapatric species with high morphological and ecological similarity and the ability to hybridize. Using greenhouse and field experiments, we tested for habitat isolation and genetic incompatibilities. From controlled crosses, we identified unilateral incompatibility (a partial prezygotic barrier associated with the self-incompatibility system), but no evidence of other genetic incompatibilities. We found a small reduction in pollen viability of F(1) hybrids and early germination of F(1), F(2), and BC hybrids relative to L. alabamica and L. crassa in a common garden experiment, but the effect on fitness was not tested. Field studies of hybrid pollen viability and germination are needed to determine if they contribute to reproductive isolation. In a reciprocal transplant, we found no evidence of habitat isolation or reduced hybrid survival (from seedling to adult stage) or reproduction. These data suggest unilateral incompatibility partially reproductively isolates L. alabamica and L. crassa, but no other reproductive barriers could be detected.
American Journal of Botany 03/2010; 97(3):412-22. · 2.59 Impact Factor
[show abstract][hide abstract] ABSTRACT: Evolutionary biologists explain the maintenance of intermediate levels of defense in plant populations as being due to trade-offs, or negative genetic covariances among ecologically important traits. Attempts at detecting trade-offs as constraints on the evolution of defense have not always been successful, leading some to conclude that such trade-offs rarely explain current levels of defense in the population. Using the agricultural pest Ipomoea purpurea, we measured correlations between traits involved in defense to glyphosate, the active ingredient in Roundup, a widely used herbicide. We found significant allocation costs of tolerance, as well as trade-offs between resistance and two measures of tolerance to glyphosate. Selection on resistance and tolerance exhibited differing patterns: tolerance to leaf damage was under negative directional selection, whereas resistance was under positive directional selection. The joint pattern of selection on resistance and tolerance to leaf damage indicated the presence of alternate peaks in the fitness landscape such that a combination of either high tolerance and low resistance, or high resistance and low tolerance was favored. The widespread use of this herbicide suggests that it is likely an important selective agent on weed populations. Understanding the evolutionary dynamics of herbicide defense traits is thus of increasing importance in the context of human-mediated evolution.
[show abstract][hide abstract] ABSTRACT: The extent to which epistasis contributes to adaptation, population differentiation, and speciation is a long-standing and important problem in evolutionary genetics. Using recombinant inbred (RI) lines of Arabidopsis thaliana grown under natural field conditions, we have examined the genetic architecture of fitness-correlated traits with respect to epistasis; we identified both single-locus additive and two-locus epistatic QTL for natural variation in fruit number, germination, and seed length and width. For fruit number, we found seven significant epistatic interactions, but only two additive QTL. For seed germination, length, and width, there were from two to four additive QTL and from five to eight epistatic interactions. The epistatic interactions were both positive and negative. In each case, the magnitude of the epistatic effects was roughly double that of the effects of the additive QTL, varying from -41% to +29% for fruit number and from -5% to +4% for seed germination, length, and width. A number of the QTL that we describe participate in more than one epistatic interaction, and some loci identified as additive also may participate in an epistatic interaction; the genetic architecture for fitness traits may be a network of additive and epistatic effects. We compared the map positions of the additive and epistatic QTL for germination, seed width, and seed length from plants grown in both the field and the greenhouse. While the total number of significant additive and epistatic QTL was similar under the two growth conditions, the map locations were largely different. We found a small number of significant epistatic QTL x environment effects when we tested directly for them. Our results support the idea that epistatic interactions are an important part of natural genetic variation and reinforce the need for caution in comparing results from greenhouse-grown and field-grown plants.
[show abstract][hide abstract] ABSTRACT: The extent to which epistasis contributes to adaptation and speciation has been a controversial topic in evolutionary genetics. One experimental approach to study epistasis is based on quantitative trait locus (QTL) mapping using molecular markers. Comparisons can be made among all possible pair-wise combinations of the markers, irrespective of whether an additive QTL is associated with a marker; several software packages have been developed that facilitate this. We review several examples of using this approach to identify epistatic QTLs for traits of evolutionary or ecological interest. While there is variability in the results, the number of epistatic QTL interactions is often greater than or equal to the number of additive QTLs. The magnitude of epistatic effects can be larger than the additive effects. Thus, epistatic interactions seem to be an important part of natural genetic variation. Future studies of epistatic QTLs could lead to descriptions of the genetic networks underlying variation for fitness-related traits.
Genetics Research 11/2005; 86(2):89-95. · 2.00 Impact Factor
[show abstract][hide abstract] ABSTRACT: Hybridization is increasingly recognized as a significant creative force in evolution. Interbreeding among species can lead to the creation of novel genotypes and morphologies that lead to adaptation. On the Hawaiian island of O'ahu, populations of two species of plants in the endemic genus Lipochaeta grow at similar elevations in the northern Wai'anae Mountains. These two species represent extremes of the phenotypic distribution of leaf shape: the leaves of Lipochaeta tenuifolia individuals are compound and highly dissected while leaves of L. tenuis are simple. Based primarily on leaf shape morphology, a putative hybrid population of Lipochaeta located at Pu'u Kawiwi was identified. Individuals in this population exhibit a range of leaf shapes intermediate in varying degrees between the leaf shapes of the putative parental species. We analyzed individuals from pure populations of L. tenuifolia, L. tenuis and the putative hybrids using 133 AFLP markers. Genetic analysis of these neutral markers provided support for the hybrid origin of this population. The correlation between genetic background and leaf morphology in the hybrids suggested that the genome of the parental species with simple leaves might have significantly contributed to the evolution of a novel, compound leaf morphology.
[show abstract][hide abstract] ABSTRACT: Although much is known about the molecular genetic basis of trichome development in Arabidopsis thaliana, less is known about the underlying genetic basis of continuous variation in a trait known to be of adaptive importance: trichome density. The density of leaf trichomes is known to be a major determinant of herbivore damage in natural populations of A. thaliana and herbivores are a significant selective force on genetic variation for trichome density. A number of developmental changes occur during ontogeny in A. thaliana, including changes in trichome density. I used multiple interval mapping (MIM) analysis to identify QTL responsible for trichome density on both juvenile leaves and adult leaves in replicate, independent trials and asked whether those QTL changed with ontogeny. In both juvenile and adult leaves, I detected a single major QTL on chromosome 2 that explained much of the genetic variance. Although additional QTL were detected, there were no consistent differences in the genetic architecture of trichome density measured on juvenile and adult leaves. The finding of a single QTL of major effect for a trait of known adaptive importance suggests that genes of major effect may play an important role in adaptation.
[show abstract][hide abstract] ABSTRACT: Ecologists study the rules that govern processes influencing the distribution and abundance of organisms, particularly with respect to the interactions of organisms with their biotic and abiotic environments. Over the past decades, using a combination of sophisticated mathematical models and rigorous experiments, ecologists have made considerable progress in understanding the complex web of interactions that constitute an ecosystem. The field of genomics runs on a path parallel to ecology. Like ecology, genomicists seek to understand how each gene in the genome interacts with every other gene and how each gene interacts with multiple, environmental factors. Gene networks connect genes as complex as the 'webs' that connect the species in an ecosystem. In fact, genes exist in an ecosystem we call the genome. The genome as ecosystem is more than a metaphor--it serves as the conceptual foundation for an interdisciplinary approach to the study of complex systems characteristic of both genomics and ecology. Through the infusion of genomics into ecology and ecology into genomics both fields will gain fresh insight into the outstanding major questions of their disciplines.
[show abstract][hide abstract] ABSTRACT: Glyphosate, the active ingredient in the herbicide RoundUp, has increased dramatically in use over the past decade and constitutes a potent anthropogenic source of selection. In the southeastern United States, weedy morning glories have begun to develop tolerance to glyphosate, representing a unique opportunity to examine the evolutionary genetics of a novel trait. We found genetic variation for tolerance, indicating the potential for the population to respond to selection by glyphosate. However, the following significant evolutionary constraint exists: in the absence of glyphosate, tolerant genotypes produced fewer seeds than susceptible genotypes. The combination of strong positive directional selection in the presence of glyphosate and strong negative directional selection in its absence may indicate that the selective landscape of land use could drive the evolutionary trajectory of glyphosate tolerance. Understanding these evolutionary forces is imperative for devising comprehensive management strategies to help slow the rate of the evolution of tolerance.
Proceedings of the National Academy of Sciences 10/2004; 101(36):13386-90. · 9.74 Impact Factor
[show abstract][hide abstract] ABSTRACT: Pathogen resistance is an ecologically important phenotype increasingly well understood at the molecular genetic level. In this article, we examine levels of avrRpt2-dependent resistance and Rps2 locus DNA sequence variability in a worldwide sample of 27 accessions of Arabidopsis thaliana. The rooted parsimony tree of Rps2 sequences drawn from a diverse set of ecotypes includes a deep bifurcation separating major resistance and susceptibility clades of alleles. We find evidence for selection maintaining these alleles and identify the N-terminal part of the leucine-rich repeat region as a probable target of selection. Additional protein variants are found within the two major clades and correlate well with measurable differences among ecotypes in resistance to the avirulence gene avrRpt2 of the pathogen Pseudomonas syringae. Long-lived polymorphisms have been observed for other resistance genes of A. thaliana; the Rps2 data suggest that the long-term maintenance of phenotypic variation in resistance genes may be a general phenomenon and are consistent with diversifying selection acting in concert with selection to maintain variation.
[show abstract][hide abstract] ABSTRACT: Measuring natural selection has been a fundamental goal of evolutionary biology for more than a century, and techniques developed in the last 20 yr have provided relatively simple means for biologists to do so. Many of these techniques, however, share a common limitation: when applied to phenotypic data, environmentally induced covariances between traits and fitness can lead to biased estimates of selection and misleading predictions about evolutionary change. Utilizing estimates of breeding values instead of phenotypic data with these methods can eliminate environmentally induced bias, although this approach is more difficult to implement. Despite this potential limitation to phenotypic methods and the availability of a potential solution, little empirical evidence exists on the extent of environmentally induced bias in phenotypic estimates of selection. In this article, we present a method for detecting bias in phenotypic estimates of selection and demonstrate its use with three independent data sets. Nearly 25% of the phenotypic selection gradients estimated from our data are biased by environmental covariances. We find that bias caused by environmental covariances appears mainly to affect quantitative estimates of the strength of selection based on phenotypic data and that the magnitude of these biases is large. As our estimates of selection are based on data from spatially replicated field experiments, we suggest that our findings on the prevalence of bias caused by environmental covariances are likely to be conservative.
The American Naturalist 11/2002; 160(4):511-23. · 4.55 Impact Factor