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Genetic architecture of adaptation is traditionally studied in the context of local adaptation, viz. spatially varying conditions experienced by the species. However, anthropogenic changes in the natural environment pose a new context to this issue, that is, adaptation to an environment that is new for the species. In this study, we used crossbreeding to analyze genetic architecture of adaptation to conditions not currently experienced by the species but with high probability of encounter in the near future due to global climate change. We performed targeted interpopulation crossing using genotypes from two core and two peripheral Triticum dicoccoides populations and raised the parents and three generations of hybrids in a greenhouse under simulated desert conditions to analyze the genetic architecture of adaptation to these conditions and an effect of gene flow from plants having different origin. The hybrid (F1) fitness did not differ from that of the parents in crosses where both plants originated from the species core, but in crosses involving one parent from the species core and another one from the species periphery the fitness of F1 was consistently higher than that of the periphery-originated parent. Plant fitness in the next two generations (F2 and F3) did not differ from the F1, suggesting that effects of epistatic interactions between recombining and segregating alleles of genes contributing to fitness were minor or absent. The observed low importance of epistatic gene interactions in allopolyploid T. dicoccoides and low probability of hybrid breakdown appear to be the result of permanent fixation of heterozygosity and lack of intergenomic recombination in this species. At the same time, predominant but not complete selfing combined with an advantage of bivalent pairing of homologous chromosomes appears to maintain high genetic variability in T. dicoccoides, greatly enhancing its adaptive ability.

Genetic differentiation in 20 hierarchically sampled populations of wild barley was analyzed with quantitative traits, allozymes and Random Amplified Polymorphic DNAs (RAPDs), and compared for three marker types at two hierarchical levels. Regional subdivision for both molecular markers was much lower than for quantitative traits. For both allozymes and RAPDs, most loci exhibited minor or no regional differentiation, and the relatively high overall estimates of the latter were due to several loci with exceptionally high regional differentiation. The allozyme- and RAPD-specific patterns of differentiation were concordant in general with one another, but not with quantitative trait differentiation. Divergent selection on quantitative traits inferred from very high regional Q(ST) was in full agreement with our previous results obtained from a test of local adaptation and multilevel selection analysis. In contrast, most variation in allozyme and RAPD variation was neutral, although several allozyme loci and RAPD markers were exceptional in their levels of regional differentiation. However, it is not possible to answer the question whether these exceptional loci are directly involved in the response to selection pressure or merely linked to the selected loci. The fact that Q(ST) and F(ST) did not differ at the population scale, that is, within regions, but differed at the regional scale, for which local adaptation has been previously shown, implies that comparison of the level of subdivision in quantitative traits, as compared with molecular markers, is indicative of adaptive population differentiation only when sampling is carried out at the appropriate scale.

The adaptive potential of a population defines its importance for species survival in changing environmental conditions such as global climate change. Very few empirical studies have examined adaptive potential across species' ranges, namely, of edge vs core populations, and we are unaware of a study that has tested adaptive potential (namely, variation in adaptive traits) and measured performance of such populations in conditions not currently experienced by the species but expected in the future. Here we report the results of a Triticum dicoccoides population study that employed transplant experiments and analysis of quantitative trait variation. Two populations at the opposite edges of the species range (1) were locally adapted; (2) had lower adaptive potential (inferred from the extent of genetic quantitative trait variation) than the two core populations; and (3) were outperformed by the plants from the core population in the novel environment. The fact that plants from the species arid edge performed worse than plants from the more mesic core in extreme drought conditions beyond the present climatic envelope of the species implies that usage of peripheral populations for conservation purposes must be based on intensive sampling of among-population variation.

Populations of predominantly selfing plant species often show spatial genetic structure but little is known whether epistatic gene interactions are spatially structured. To detect a possible epistatic effect and a spatial scale at which it operates, we created artificial crosses between plants spanning a range of fixed distances from 1 to 400 m in three populations of wild barley. The self-pollinated and crossed progeny (F(1)) and two generations of segregated progeny (F(2) and F(3)) were tested in experimentally simulated population environments for relative performance (RP). The measured fitness traits included number of seeds, total seed weight and seed germination. For any of these traits, there was no association between RP of F(1), F(2) and F(3) plants and either pairwise kinship coefficients or crossing distance. In contrast, in all three populations, we found lower seed viability of outcrossed as compared with self-pollinated genotypes in the first generation of segregation. However, in the F(3) generation this outbreeding effect disappeared in the two populations and greatly decreased in the third population. For seed production, heterosis in F(1) and outbreeding depression in F(2) were observed only in the population with unusually high number of heterozygotes. Our findings support the view that in selfing species a spatial mosaic of various locally abundant genotypes represents not randomly fixed combinations of alleles but the co-adapted gene complexes that were sieved by selection, while heterozygotes are characteristic for the transient phase of this process, when segregation and purging of maladaptive genotypes have not yet occurred.