No abstract

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 allozymeand 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. Heredity (2005) 95, 466-475.

Reciprocal introduction of seeds and seedlings was used to test for local adaptation and to identify a set of co-adapted traits of Mediterranean and desert ecotypes of wild barley Hordeum spontaneum. Evidence for local adaptation was found in seedling introductions into intact environments and from ecotype colonization success in the first generation after seed dispersal. Estimates of fitness were obtained at particular stages of the life cycle (seed, seedling and adult). Experiments that manipulated the environment (vegetation removal, different plant density) demonstrated the intensity and direction of natural selection in different life history episodes, but there was no strong evidence for local adaptation under these circumstances. The observed genetically determined differences between Mediterranean and desert ecotypes can be summarized as the following: reproductive output was higher in desert plants, with smaller seeds than in Mediterranean plants. There was a higher competitive ability of Mediterranean than desert plants. Plants of desert origin had significant reductions in yield when grown in mixed stands with Mediterranean plants; no such effect was observed for plants of Mediterranean origin. Seed germination and seedling survival was lower in seeds of desert origin. This was due to both: genetically determined higher dormancy of desert seeds and a trade-off between no. of seeds and their size (directly related to seed/seedling vigour).

Reciprocal introduction of seeds and seedlings of wild barley, Hordeum spontaneum , originating in four different environments of Israel was used to: (1) test for local adaptation, (2) make inferences about environmental effects on life‐history and reproductive traits, and (3) identify trait combinations with recognizable ‘strategies’. The four populations examined represented the following environments: (1) desert − low productivity and predictability, drought stress; (2) semi‐steppe batha − moderate productivity and predictability; (3) grassland − high productivity and predictability; and (4) mountain − high productivity and predictability but with severe frost stress. Significant genotype‐by‐environment interactions were observed for yield and reproductive biomass, seedling biomass and percentage germinated and survived seeds, suggesting local ecotype adaptation. Increasing productivity and predictability of environment in respect to rainfall, without concomitant frost stress, was found to select for high reproductive biomass and large seeds, a high fraction of germinating seeds and high vigour of seedlings. The optimal strategy changes with increasing productivity and predictability and involves a trade‐off between seed size and number, with reduced yield but increased seed mass, consistent with competition selection (or K‐selection sensu MacArthur & Wilson (1967)) type. No specific life‐history adaptations to predictable frost stress were detected for the mountain ecotype, but there was higher survival of seedlings in their indigenous (mountain) environment compared with other ecotypes. The latter appears to be a physiological adaptation to frost, which is consistent with selection for stress tolerance (or S‐selection sensu Grime (1977)) type. The other stress factor, drought, which is very unpredictable in deserts, was associated with high seed dormancy, small seed size and low vigour of seedlings, but relatively high yield, which is consistent with a stress‐escape bet‐hedging strategy. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77, 479–490.

Plants from four populations of Hordeum spontaneum originating in distinct environments of Israel were compared for stress induced phenotypic plasticity. The environments ranged along a gradient of increasing rainfall amount and predictability from low (desert) to moderate (semisteppe batha) to high (Mediterranean grassland and mountain, the latter also experiencing frost stress). The plants were exposed to a set of four treatments: no stress (optimum water and nutrients), water, nutrient and both water and nutrient stress. Plants from the four populations (or ecotypes) exhibited different patterns of plasticity in response to the different stresses (water and nutrients) and in different trait categories (reproductive, fitness and resource allocation). The importance of plasticity in response to water stress appears to decrease, and to nutrient stress appears to increase along the increasing rainfall gradient. The mountain ecotype, growing in an area with high potential productivity (amount of rainfall) but experiencing periodic frosts, was the most plastic among ecotypes in resource allocation under both water and nutrient stress, but exhibited low plasticity in other trait categories. In contrast, the desert ecotype had low plasticity in resource allocation under water stress and the lowest plasticity among the four ecotypes in all trait categories in response to nutrient stress. The ecotype originating in Mediterranean grassland, a predictable and most favourable environment, was highly plastic in fitness and allocation traits in response to low nutrient levels which is likely to occur due to competition in productive environment. We discuss the observed differences in ecotype plasticity as part of their environmentally induced adaptive ‘strategies’. We found no support for the hypothesis that plants originating in environments with greater variation and unpredictability are more plastic. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society 2002, 75, 301–312.