Avigdor Beiles

Microsatellites: genomic distribution, putative functions and mutational mechanisms: a review

Microsatellites, or tandem simple sequence repeats (SSR), are abundant across genomes and show high levels of polymorphism. SSR genetic and evolutionary mechanisms remain controversial. Here we attempt to summarize the available data related to SSR distribution in coding and noncoding regions of genomes and SSR functional importance. Numerous lines of evidence demonstrate that SSR genomic distribution is nonrandom. Random expansions or contractions appear to be selected against for at least part of SSR loci, presumably because of their effect on chromatin organization, regulation of gene activity, recombination, DNA replication, cell cycle, mismatch repair system, etc. This review also discusses the role of two putative mutational mechanisms, replication slippage and recombination, and their interaction in SSR variation.

The Evolutionary Significance of Genetic Diversity: Ecological, Demographic and Life History Correlates

The evolutionary significance of genetic diversity of proteins in nature remains controversial despite the numerous protein studies conducted electrophoretically during the last two decades. Ironically, the discovery of extensive protein polymorphisms in nature (reviewed by Lewontin, 1974; Powell, 1975; Selander, 1976; Nevo 1978, 1983b; Hamrick et al., 1979; Nelson and Hedgecock, 1980), did not resolve the disagreement between the die ho torn ou s explanatory models of selection (e.g., Ayala, 1977; Milkman, 1978; Clarke, 1979; Wills, 1981) versus neutrality (Kimura, 1968; Kimura and Chta, 1971; Nei, 1975; and modifications in Kimura, 1979atb). The more general problem of the relative importance of the evolutionary forces interacting in genetic population differentiation at the molecular levels of proteins and DNA, i.e., mutation, migration, natural selection and genetic drift, remains now as enigmatic as ever.

Genetic diversity of wild emmer wheat in Israel and Turkey : Structure, evolution, and application in breeding.

SummaryAllozyme variation in the tetraploid wild emmer wheat, Triticum dicoccoides, the progenitor of all cultivated wheats, was studied for the proteins encoded by 42 gene loci in 1815 plants representing 37 populations - 33 from Israel and 4 from Turkey - sampled in 33 localities from 1979 to 1987. The results showed that: (a) 6 loci (14%) were monomorphic in all populations, 15 loci (36%) were locally polymorphic, and 21 loci (50%) were regionally polymorphic. These results are similar to those obtained earlier on 12 Israeli populations. All polymorphic loci (except 4) displayed high local levels of polymorphism (>/ 10%). (b) The mean number of alleles per locus, A, was 1.252 (range: 1.050–1.634); the proportion of polymorphic loci per population averaged 0.220 (range: 0.050–0.415); genic diversity, He, averaged 0.059 (range: 0.002–0.119). (c) Altogether there were 119 alleles at the 42 putative loci tested, 114 of these in Israel, (d) Genetic differentiation was primarily regional and local, not clinal; 70% of the variant alleles were common (>/ 10%) and not widespread, but rather localized or sporadic, displaying an “archipelago” population genetics and ecology structure. The coefficients of genetic distance between populations were high and averaged D = 0.134; range: 0.018–0.297, an indication of sharp genetic differentiation over short distances, (e) Discriminant analyses differentiated Israeli from Turkish populations, and within Israel, between central and 3 marginal regions, as well as between different soil-type populations, (f) Allozymic variation comprised 40% within and 60% between populations, (g) Gametic phase disequilibria were abundant, their number being positively correlated (rs = 0.60, P<0.01) with the humidity, (h) Multilocus organization was substantive, also positively correlated with humidity, (i) Allozyme diversity, overall and at single loci, was significantly correlated with, and partly predictable by, climatic and edaphic factors, (j) The distrubition of the significant positive and negative values and the absence of autocorrelations in the correlogram revealed no similar geographic patterns across loci, eliminating migration as a prime factor of population genetic differentiation. These results suggest: (I) during the evolutionary history of wild emmer, diversifying natural selection, through climatic and edaphic factors, was a major agent of genetic structure and differentiation at both the single and multilocus levels; (II) wild emmer harbors large amounts of genetic diversity exploitable as genetic markers in sampling and abundant genetic resources utilizable for wheat improvement.

Genetic diversity and environmental associations of wild wheat, Triticum dicoccoides, in Israel

SummaryAllozyme variation in the tetraploid wild progenitor of wheat, Triticum dicoccoides, was studied for the proteins encoded by about 50 gene loci in 457 individuals representing 12 populations from Israel. Six spikelet morphological traits were measured in the same populations. The results indicate that: (a) 16 loci (= 32%) were monomorphic in all 12 populations, 15 loci (= 30%) were locally polymorphic, and 19 loci (= 38%) were regionally polymorphic. All polymorphic loci (but one) displayed high levels of polymorphism (≧ 10%). In Israel, the proportion of polymorphic loci per population, P, in wild wheat averaged 0.25 (range, 0.16–0.38), and the genetic diversity index, He averaged 0.07, (range, 0.03 – 0.12). (b) Altogether there were 110 alleles at the 50 putative loci tested (c) Genetic differentiation of populations included regional and local patterns: (i) The coefficients of genetic distance between populations were high (mean D = 0.10 range, 0.02 – 0.25), and indicated sharp genetic differentiation over short distances, (ii) Common (≧ 10%) but sporadic and localized alleles were frequent (76%), and (iii) Rare alleles were few (only 5 alleles). (d) The patterns of allozyme and spikelet variation in the wild gene pool were significantly correlated with, and partly predictable by, water factors, including those of precipitation, evaporation, and relative humidity as well as of soil type, (e) All six spikelet characters showed statistically significant variation among localities and (f) Allozymic variation was correlated with spikelet variation.These results suggest in T. dicoccoides: (i) the operation of natural selection in population genetic structure, (ii) local adaptive genetic differentiation caused by diversifying selection through climate and soil, and (iii) the guidelines for sampling these resources for use in wheat breeding programs.

RAPD polymorphism of wild emmer wheat populations, Triticum dicoccoides, in Israel

Abstract Genetic diversity in random amplified polymorphic DNAs (RAPDs) was studied in 110 genotypes of the tetraploid wild progenitor of wheat, Triticum dicoccoides, from 11 populations sampled in Israel and Turkey. Our results show high level of diversity of RAPD markers in wild wheat populations in Israel. The ten primers used in this study amplified 59 scorable RAPD loci of which 48 (81.4%) were polymorphic and 11 monomorphic. RAPD analysis was found to be highly effective in distinguishing genotypes of T. dicoccoides originating from diverse ecogeographical sites in Israel and Turkey, with 95.5% of the 100 genotypes correctly classified into sites of origin by discriminant analysis based on RAPD genotyping. However, interpopulation genetic distances showed no association with geographic distance between the population sites of origin, negating a simple isolation by distance model. Spatial autocorrelation of RAPD frequencies suggests that migration is not influential. Our present RAPD results are non-random and in agreement with the previously obtained allozyme patterns, although the genetic diversity values obtained with RAPDs are much higher than the allozyme values. Significant correlates of RAPD markers with various climatic and soil factors suggest that, as in the case of allozymes, natural selection causes adaptive RAPD ecogeographical differentiation. The results obtained suggest that RAPD markers are useful for the estimation of genetic diversity in wild material of T. dicoccoides and the identification of suitable parents for the development of mapping populations for the tagging of agronomically important traits derived from T. dicoccoides.

Tibet is one of the centers of domestication of cultivated barley

The Near East Fertile Crescent is well recognized as a primary center of barley origin, diversity, and domestication. A large number of wild barleys have been collected from the Tibetan Plateau, which is characterized by an extreme environment. We used genome-wide diversity array technology markers to analyze the genotypic division between wild barley from the Near East and Tibet. Our results confirmed the existence of Tibetan wild barley and suggested that the split between the wild barleys in the Near East and those in Tibet occurred around 2.76 million years ago (Mya). To test the concept of polyphyletic domestication of barley, we characterized a set of worldwide cultivated barley. Some Chinese hulless and six-rowed barleys showed a close relationship with Tibetan wild barley but showed no common ancestor with other cultivated barley. Our data support the concept of polyphyletic domestication of cultivated barley and indicate that the Tibetan Plateau and its vicinity is one of the centers of domestication of cultivated barley. The current results may be highly significant in exploring the elite germplasm for barley breeding, especially against cold and drought stresses.

Microgeographic edaphic differentiation in hordein polymorphisms of wild barley.

SummaryGenetic diversity in the storage protein hordein encoded by two loci, Hor1 and Hor2, was analyzed electrophoretically in seeds from 123 individual plants of wild barley, Hordeum spontaneum, the progenitor of cultivated barley. The test was conducted in two topographically different 100 meter transects in Israel, each equally divided into basalt and terra rossa soil types. Altogether 15 Hor1 and 16 Hor2 phenotypes were detected; 7 phenotypes in Hor1 and 5 in Hor2 were common. Significant differentiation of both Hor1 and Hor2 phenotypes and their associations was found with soil type and topography. Likewise, significant correlations were found between hordein phenotypes and allozyme types detected in a previous study. Our results suggest that at least part of the hordein polymorphisms in wild barley is adaptive and selected by soil and topographic differences over very short distances.

Evolution of wild cereals during 28 years of global warming in Israel

Climate change is a major environmental stress threatening biodiversity and human civilization. The best hope to secure staple food for humans and animal feed by future crop improvement depends on wild progenitors. We examined 10 wild emmer wheat (Triticum dicoccoides Koern.) populations and 10 wild barley (Hordeum spontaneum K. Koch) populations in Israel, sampling them in 1980 and again in 2008, and performed phenotypic and genotypic analyses on the collected samples. We witnessed the profound adaptive changes of these wild cereals in Israel over the last 28 y in flowering time and simple sequence repeat allelic turnover. The revealed evolutionary changes imply unrealized risks present in genetic resources for crop improvement and human food production.

Mercury selection of allozyme genotypes in shrimps

The effects of mercury pollution on the allozymic variation of 15 phosphoglucomutase (PGM) genotypes was tested in the Mediterranean shrimpPalaemon elegans in 79 laboratory tests involving 2765 shrimps, with 1560 survivors (767 test and 793 control). Our results indicate differential tolerance of genotypes in variable mercury concentrations, suggesting that they are adaptive. The genetic structure can possibly be explored and potentially be used as a monitoring system for the quality and quantity of marine pollutants.

Population genetic response to microsite ecological stress in wild barley, Hordeum spontaneum

Genetic diversity was studied in six subpopulations (a total of 60 individuals) of wild barley, Hordeum spontaneum, the progenitor of cultivated barley, sampled from six stations located along a transect of 300 m across the two opposing slopes of ‘Evolution Canyon’, a Mediterranean microsite at Lower Nahal Oren, Mt Carmel. The two opposing slopes are separated by between 100 and 400 m and designated SFS (South‐Facing Slope) and NFS (North‐Facing Slope) with each having three equidistant test stations. The SFS, which receives up to 300% more solar radiation, is drier, ecologically more heterogeneous, fluctuating, and more stressful than the NFS. Analysis of 12 RAPD primers, representing a total of 51 putative loci, revealed a significant inter‐ and intraslope variation in RAPD band polymorphism. A significantly higher proportion of polymorphic RAPD loci was found amongst the subpopulations on the SFS (mean P= 0.909) than on the NFS (mean P= 0.682), on the basis of the presence and absence of 22 strong bands. Polymorphism generally increased upwards from the bottom to the top of the SFS (0.636, 0.773, 0.955) and NFS (0.409, 0.500, 0.545), respectively. Gametic phase disequilibria estimates, D, revealed SFS and NFS unique predominant combinations which sharply differentiated the two slopes and indicated that there is differential interslope selection favouring slope‐specific multilocus combinations of alleles, or blocks of genes over tens to hundreds of meters. This suggests that selection overrides migration. RAPD polymorphism appears to parallel allozyme diversity which is climatically adaptive and driven by natural selection in the same subpopulations at the microsite.