Complex-trait analysis in plants

Identifying quantitative trait loci in Arabidopsis by population and pedigree studies

Inference of selection gradients using performance measures as fitness proxies

O.D.F. is supported by the Natural Sciences and Engineering Research Council of Canada and the Madame Vigdís Finnbogadóttir Scholarship. M.B.M. is supported by a University Research Fellowship from the Royal Society (London).1.  Selection coefficients, i.e., selection differentials and gradients, are useful for quantifying selection and for making comparisons across traits and organisms, because they appear in known equations for relating selection and genetic variation to one another and to evolutionary change. However, selection coefficients can only be estimated in organisms where traits and fitness (components) can be measured. This is probably a major contributor to taxonomic biases of selection studies. Aspects of organismal performance, i.e., quantities that are likely to be positively related to fitness components, such as body size, are sometimes used as proxies for fitness, i.e., used in place of fitness components in regression-based selection analysis. To date, little theory exists to inform empirical studies about whether such procedures may yield selection coefficients with known relationships to genetic variation and evolution. 2.  We show that the conditions under which performance measures can be used as proxies for fitness are very limited. Such analyses require that the regression of fitness on the proxy is linear and goes through the origin. We illustrate how fitness proxies may be used in conjunction with information about the performance-fitness relationship, and clarify how this is different from substituting fitness proxies for fitness components in selection analyses. 3.  We apply proxy-based and fitness component-based selection analysis to a system where traits, a performance measure (size; similar to proxies that are commonly used in place of fitness), and a more proximate fitness measure, are all available on the same set of individuals. We find that proxy-based selection gradients are poorly reflective of selection gradients estimated using fitness components, even when proxy-fitness relationships are quite strong and reasonably linear. 4.  We discuss the implications for proxy-based selection analysis. We emphasise that measures of organismal performance, such as size, may in many cases provide useful information that can contribute to quantitative inferences about natural selection, and their use could allow quantitative inference about selection to be conducted in a wider range of taxa. However, such inferences require quantitative analysis of both trait-performance and performance-fitness relationships, rather than substitution of performance for measures of fitness or fitness components.PostprintPeer reviewe

Mlcoalsim: Multilocus Coalescent Simulations

Coalescent theory is a powerful tool for population geneticists as well as molecular biologists interested in understanding the patterns and levels of DNA variation. Using coalescent Monte Carlo simulations it is possible to obtain the empirical distributions for a number of statistics across a wide range of evolutionary models; these distributions can be used to test evolutionary hypotheses using experimental data. The mlcoalsim application presented here (based on a version of the ms program, Hudson, 2002) adds important new features to improve methodology (uncertainty and conditional methods for mutation and recombination), models (including strong positive selection, finite sites and heterogeneity in mutation and recombination rates) and analyses (calculating a number of statistics used in population genetics and P-values for observed data). One of the most important features of mlcoalsim is the analysis of multilocus data in linked and independent regions. In summary, mlcoalsim is an integrated software application aimed at researchers interested in molecular evolution. mlcoalsim is written in ANSI C and is available at: http://www.ub.es/softevol/mlcoalsim

Cost of Defense in the Context of Plant Competition: Brassica Rapa May Grow and Defend

Theory on costs of plant defense against herbivory in stressful environments predicts that costs should increase when competition is intense. This amplifies a fundamental dilemma that plants are thought to face: allocate limited resources to grow fast enough to compete, or invest these resources in secondary metabolites to maintain defense. We studied costs associated with genetic and environmental variation in secondary metabolite production of Brassica rapa in the presence and absence of the generalist competitor Lolium perenne. We used experimental quantitative genetics (artificial selection) to manipulate genetic variation, and herbivore-induction treatments to produce environmental variation in myrosinase and glucosinolate concentrations and resistance. Glucosinolates, and their byproducts after breakdown by myrosinase, are known to affect herbivory on plants in the Brassicaceae family. Defense costs were significant in the absence of competitors, but in contrast to theoretical predictions, costs of constitutive defense (measured as growth rates) were not detectable and the cost of induced defense remained the same in the competitive environment. To understand what factors made constitutive defense costs not detectable under competition we conducted several experiments to assess the effects of limited resources and allelopathy on costs and benefits of the defense chemicals. None of the experiments involving nutrient supply and weak competition supported the hypothesis that the lack of defense costs in competitive environments was due to limited resources. Instead, the breakdown products of the glucosinolate–myrosinase reaction appeared to function as allelopathic agents, which may benefit B. rapa plants in competition, thereby reducing net costs of chemical defense. We found that: (1) the effects of exogenous glucosinolates on Lolium root length depended on the presence of myrosinase. (2) In the absence of nutrients, Lolium root lengths were shorter when seeds germinated with B. rapa. (3) Genetic increases in glucosinolate concentration negatively affected Lolium seedling growth only when there were simultaneous genetic increases in myrosinase concentration. Activated carbon treatments designed to neutralize allelopathic effects and restore costs in the competitive environments were, however, not statistically significant. When plant defenses also function to benefit plants in competitive interactions, plants may evolve to compete and defend

Understanding rice adaptation to varying agro-ecosystems: trait interactions and quantitative trait loci

Background: Interaction and genetic control for traits influencing the adaptation of the rice crop to varying environments was studied in a mapping population derived from parents (Moroberekan and Swarna) contrasting for drought tolerance, yield potential, lodging resistance, and adaptation to dry direct seeding. A BC2F3-derived mapping population for traits related to these four trait groups was phenotyped to understand the interactions among traits and to map and align QTLs using composite interval mapping (CIM). The study also aimed to identify QTLs for the four trait groups as composite traits using multivariate least square interval mapping (MLSIM) to further understand the genetic control of these traits. Results: Significant correlations between drought- and yield-related traits at seedling and reproductive stages respectively with traits for adaptation to dry direct-seeded conditions were observed. CIM and MLSIM methods were applied to identify QTLs for univariate and composite traits. QTL clusters showing alignment of QTLs for several traits within and across trait groups were detected at chromosomes 3, 4, and 7 through CIM. The largest number of QTLs related to traits belonging to all four trait groups were identified on chromosome 3 close to the qDTY3.2 locus. These included QTLs for traits such as bleeding rate, shoot biomass, stem strength, and spikelet fertility. Multivariate QTLs were identified at loci supported by univariate QTLs such as on chromosomes 3 and 4 as well as at distinctly different loci on chromosome 8 which were undetected through CIM. Conclusion: Rice requires better adaptation across a wide range of environments and cultivation practices to adjust to climate change. Understanding the genetics and trade-offs related to each of these environments and cultivation practices thus becomes highly important to develop varieties with stability of yield across them. This study provides a wider picture of the genetics and physiology of adaptation of rice to wide range of environments. With a complete understanding of the processes and relationships between traits and trait groups, marker-assisted breeding can be used more efficiently to develop plant types that can combine all or most of the beneficial traits and show high stability across environments, ecosystems, and cultivation practices