Allelic Diversity of Hrd A and Hrd B Hordein-Coding Loci in Wild (Hordeum spontaneum C. Koch) and Cultivated (Hordeum vulgare L.) Barley from Israel and Palestine

Russian Journal of Genetics - Polymorphism of hordeins encoded by the Hrd A and Hrd B loci was studied using starch gel electrophoresis in 13 barley landrace accessions and in 31 wild barley...     

Tests for adaptive RAPD variation in population genetic structure of wild barley, Hordeum spontaneum Koch.

We examined the adaptive importance of RAPD variation in the population genetic structure of wild barley, Hordeu m spontaneum. The test involved (1) a nested sampling design with four population groups representing four distinct environments; and (2) a comparison of observed variation with that expected as a result of natural selection. Analyses of selection on fitness-related traits by reciprocal introductions served as guidelines for the expected pattern of RAPD variation. We found no concordance between the observed pattern of population genetic structure and that expected under the null hypothesis of environment-specific natural selection. There was no relationship between genetic distance and environmental similarity; none of 54 putative loci exhibited an allele distribution in accordance with that expected and no favoured epistatic allele combinations were detected across the four environments. The fact that environmentally induced adaptation, detected by fitness-related traits, was not reflected in inter-population RAPD structure (1) strongly enhances the neutralist viewpoint and (2) casts doubt on the notion that significant correlations between some environmental parameters and allele frequencies in one or more loci are evidence of selection on the latter.     

Phenotypic and allozyme variation in Mediterranean and desert populations of wild barley,Hordeum spontaneum Koch

Populations of wild barley, Hordeum spontaneum Koch, were collected in two distinct climatic regions, desert and Mediterranean. Plants from five desert and five Mediterranean populations were compared and contrasted for extent and structure of phenotypic variation. These same 10 and one other population from each region were analyzed for allozyme variation. In a field trial of phenotypic diversity, two phenological and 14 morphological traits were examined. Study of allozyme variation was performed using eight enzyme systems encoding for 13 loci. Plants from the desert and Mediterranean regions were significantly different in seven of 16 phenotypic traits, exhibited a high (30%) interregional component of phenotypic variation, and showed a high degree of segregation on a principal component scattergram indicating ecotypic differentiation. Mediterranean populations were twice as variable as desert populations in reproductive growth parameters (stem and spike length) and grain filling (spikelet weight), but half as variable for onset of reproduction. The extent and structure of phenotypic and allozyme variation did not match. The Mediterranean and desert populations did not differ in amount of allozyme variation as estimated by mean number of alleles per locus, effective number of alleles, polymorphism, and gene diversity (n(a), n(e), P, and H(e)), did not segregate on the basis of population genetic distances, and exhibited a low proportion of interregion allozyme diversity (2%). No effect of selection on allozyme distribution was detected. Our results suggest that the adaptation of plants originating from desert and Mediterranean environments is reflected in phenotypic but not in allozyme variation.     

野生二棱大麦种子休眠型态与农艺性状及生态地理因素相关性研究

对来自以色列不同地区16个生态型野生二棱大麦种子的休眠型态与其农艺性状及起源地生态地里因素的相关性进行了研究。结果表明：高温（40℃）储藏可以打破种子的休眠;16个生态型种子在高温处理下的萌发率表现出显著差异,其休眠打破过程显示出不同型态的对数生长曲线：8个旱生生态型为S型,而8个湿生生态型为倒L型。休眠深度用实际达到最大萌发率的时间度量,最低休眠深度（15·6d）是来自湿润地区＂进化峡谷＂的生态型37-N,而最深休眠深度（103·1d）是来自干旱地区Ein-Zukim（死海附近）的生态型32-6。此外,对11个物候及农艺性状指标与休眠深度做斯皮尔曼秩相关分析,结果有9个显示出显著相关,尤其是粒重与休眠深度有极显著相关性。同时,休眠深度与起源地15个生态地理因素中的9个有显著相关,种子休眠主要受其起源地的地理位置、温度和水分条件等影响。可见,野生二棱大麦自然选择进化了休眠特性去应对干热环境而繁衍生息。本研究结果可用于进一步遗传研究和现代栽培大麦品种的改良。     

Induced Androgenesis in vitro in Mutated Populations of Barley,Hordeum vulgare

Graduated concentrations of chemomutagens ethylmethanesulphonate (EMS), N-nitroso-N-methylurea (MNU) and N-nitroso-N-ethylurea (ENU) and one concentration of sodium azide (NaN₃) were used to treat seeds of spring barley cultivars Heris, Tolar, Granát and Novum. Androgenesis in vitro was induced in mutagenised plant populations. The significant stimulation effect of mutagenic treatment on the mean number of androgenic anthers from all 36 treated variants was registered in 17 variants, on the mean number of regenerated plants in 15 variants and on the mean number of regenerated plants per androgenic anthers in seven variants only. Genotypes with a lower androgenic response were relatively more stimulated. Evaluation was made of the frequency of chlorophyll mutations within androgenic regenerants and in seed progeny of androgenic donors. Androgenesis was also induced in vital chlorophyll mutants and in the variants where the mutagen treatment resulted in less than 50% survival of the donor plants. Ratios between frequency of haploids and spontaneous dihaploids were similar in green and in albino androgenic plants. The results confirm that in barley it is possible to enhance the frequency of in vitro pollen embryogenesis by mutagenic treatment of seeds.     

Genetic diversity of four esterase loci in natural populations of Hordeum spontaneum C. Koch from Jordan

Summary The diversity of four esterase loci was studied electrophoretically in 690 individual spikes representing 12 populations of wild barley (Hordeum spontaneum C. Koch.) collected from central, peripheral and marginal regions of its distribution in Jordan. A minimum of 6, 10, 5 and 5 alleles were observed at the Est-1, Est-2, Est-4 and Est-5 loci, respectively. Est-2 and Est-4 were the most diverse loci (H_c=0.53±0.05 and 0.46±0.07, respectively). Est-5 was intermediate (H_c=0.33+0.07) and Est-1 was the lowest (H_c=0.22±0.04). Polymorphism was highest in the central populations (H_e=0.52±0.04), followed by the peripheral (H_e=0.40±0.05) and the marginal (H_e=0.22±0.05) populations. Average allelic diversity between (G_st=0.49) and within (H_s=0.51) populations reflects a high allelic differentiation among these populations. Log-linear analyses revealed that four two-locus terms and two three-locus terms were significantly associated (P<0.05). Geographical distances between populations were not significantly correlated with Nei's genetic similarity index (r=0.16; P<0.19). It is postulated that diversifying selection is a major factor in the population genetic differentiation of these esterase loci.     

Natural Variation in Grain Selenium Concentration of Wild Barley, <Emphasis Type="Italic">Hordeum spontaneum</Emphasis>, Populations from Israel

Wild barley (Hordeum spontaneum), the progenitor of cultivated barley, is an important genetic resource for cereal improvement. Selenium (Se) is an essential trace mineral for humans and animals with antioxidant, anticancer, antiarthropathy, and antiviral effects. In the current study, the grain Se concentration (GSeC) of 92 H. spontaneum genotypes collected from nine populations representing different habitats in Israel was investigated in the central area of Guizhou Province, China. Remarkable variations in GSeC were found between and within populations, ranging from 0 to 0.387 mg kg⁻¹ among the 92 genotypes with an average of 0.047 mg kg⁻¹. Genotype 20_C from the Sede Boqer population had the highest GSeC, while genotype 25_1 from the Atlit population had the lowest. The mean value of GSeC in each population varied from 0.010 to 0.105 mg kg⁻¹. The coefficient of variation for each population ranged from 12% to 163%. Significant correlations were found between GSeC and 12 ecogeographical factors out of 14 studied. Habitat soil type also significantly affected GSeC. The wild barley exhibited wider GSeC ranges and greater diversity than its cultivated counterparts. The higher Se grain concentrations found in H. spontaneum populations suggest that wild barley germplasm confer higher abilities for Se uptake and accumulation, which can be used for genetic studies of barley nutritional value and for further improvement of domesticated cereals.     

Allozyme Genotype-Environment Relationships. I. Variation in Natural Populations of DROSOPHILA PACHEA

Remarkable geographic and seasonal uniformities in gene and genotype frequencies were found for four enzyme loci. Principal component analysis revealed two patterns of allele frequency changes in three of the allozyme loci (Est-b, Est-c and Acph) and one pattern in the Mdh-d locus. These patterns accounted for 90% to 100% of the variability at these loci. Significant positive correlations were obtained between several of the components and the variability in the non-saponifiable and alkaloid fractions of the senita cactus host plant. The genetic variance within a single locality (Guaymas, Mexico) was as great as the variance between all localities sampled. This is interpreted as a local founder effect. The temporary nature of the rotting cactus breeding site, the delayed maturation and the differential maturation rate of females and males are three factors proposed to cause forced outbreeding and counter the local founder effect. A combination of selection resulting from a direct interaction of the genotype and the environment, in addition to migration, is proposed to maintain the uniform gene frequencies.     

Grain protein variability among populations of wild barley (Hordeum spontaneum C. Koch.) from Jordan

Summary Protein content, kernel weight, and genetic diversity in the storage protein hordein, encoded by the Hor 1 and Hor 2 loci, were assessed in 12 populations of wild barley (Hordeum spontaneum C. Koch.) collected from central, peripheral, and marginal areas of its distribution in Jordan. Protein content ranged from 106.3 to 239.1 g kg^-1, and kernel weight ranged from 21.17 to 31.8 mg. Populations with high protein content and heavy kernels have been identified. Electrophoretic analysis of the storage protein hordein showed that the two hordein loci, Hor 1 and Hor 2, are highly polymorphic, having 34 and 38 alleles, respectively. Polymorphism (H_e) was highest in central populations (H_e Hor 1=0.859, H_e Hor 2=0.782), intermediate in peripheral populations (H_e Hor 1=0.566, H_e Hor 2=0.509), and lowest in marginal populations (H_e Hor 1=0.392, H_e Hor 2=0.349). Geographical distances between populations were not indicative of Nei's genetic similarity (NI). NI values averaged 0.209 and ranged from 0.0 to 0.83, supporting the hypothesis of an island population model for the species. The high proportion of allelic diversity, apportioned among populations for Hor 1 (0.584) and Hor 2 (0.495) loci, indicates that these natural populations are a rich reserve of genetic variability for protein. This variability is readily exploitable in breeding.     

A Novel and Major Quantitative Trait Locus for Fusarium Crown Rot Resistance in a Genotype of Wild Barley (Hordeum spontaneum L.)

Fusarium crown rot (FCR), caused by various Fusarium species, is a destructive disease of cereal crops in semiarid regions worldwide. As part of our contribution to the development of Fusarium resistant cultivars, we identified several novel sources of resistance by systematically assessing barley genotypes representing different geographical origins and plant types. One of these sources of resistance was investigated in this study by generating and analysing two populations of recombinant inbred lines. A major locus conferring FCR resistance, designated as Qcrs.cpi-4H, was detected in one of the populations (mapping population) and the effects of the QTL was confirmed in the other population. The QTL was mapped to the distal end of chromosome arm 4HL and it is effective against both of the Fusarium isolates tested, one F. pseudograminearum and the other F. graminearum. The QTL explains up to 45.3% of the phenotypic variance. As distinct from an earlier report which demonstrated co-locations of loci conferring FCR resistance and plant height in barley, a correlation between these two traits was not detected in the mapping population. However, as observed in a screen of random genotypes, an association between FCR resistance and plant growth rate was detected and a QTL controlling the latter was detected near the Qcrs.cpi-4H locus in the mapping population. Existing data indicate that, although growth rate may affect FCR resistance, different genes at this locus are likely involved in controlling these two traits.     

Chloroplast DNA Diversity in Populations of Wild and Cultivated Barley

Chloroplast DNA (cpDNA) diversity was found within and among populations (245 accessions total) of wild barley, Hordeum vulgare L. ssp. spontaneum Koch from Israel and Iran. Three polymorphic restriction sites (HindIII, EcoRI, BclI) which define three distinct cpDNA lineages were detected. One lineage is common to populations in the Hule Valley and Kinneret of northern Israel, and in Iran. The second lineage is found predominantly in the Lower Jordan Valley and Negev. The distribution of the third lineage is scattered but widespread throughout Israel. Sixty two accessions of cultivated barleys, H. vulgare L., were found, with two exceptions, to belong to just one cpDNA lineage of wild barley, indicating that the cpDNA of cultivated barley is less variable than its wild ancestor. These results demonstrate the need for assessing intraspecific cpDNA variability prior to choosing single accessions for phylogenetic constructions at the species level and higher.     

Differentiation along a gradient of environmental productivity and predictability in populations of Hordeum spontaneum Koch: multilevel selection analysis

A contextual analysis combined with path analysis was applied to detect ecotype-specific past selection in hierarchically structured populations of wild barley, Hordeum spontaneum . In our analysis a multiple regression model incorporated several individual and ecotype-level unmeasured (derived) traits obtained by factor analysis from 20 measured morphological and phenological traits. Under favourable conditions (high water and nutrients) both individual and ecotype plant size (RF1) were significant predictors of individual plant fitness, estimated by either reproductive biomass or yield. Both individual and ecotype size of reproductive structures (RF2) were significantly related to individual reproductive biomass. Individual yield, however, significantly correlated with ecotype RF2 only. Transition to reproduction (RF3) correlated with neither reproductive biomass nor yield at individual level, but correlated with two estimates of fitness at ecotype level. In all cases, selection at the individual and ecotype levels was in opposition. We interpret the observed effect of ecotype identity on individual fitness not as a current group selection, but as a constraining effect of ecotype-specific past selection. The four ecotypes went through an environmentally specific selection process in their own environments with the optimal strategy evolved. Consequently, this strategy may have a constraining effect on plant performance in other environments. Under conditions of either low water or low nutrients the ecotype level did not contribute to individual fitness. The latter may suggest that a mechanism for plant responses to stress is largely independent of plant origin, with a difference between ecotypes under stressful conditions due entirely to the difference in amount, not architecture, of plasticity. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 313–318.     

Allozyme Variation among Populations of Rana pirica (Amphibia: Anura)

Genetic variation of Rana pirica , an east Asian brown frog of the R. temporaria group, was elucidated by analysing 140 specimens from 11 populations from Hokkaido and Sakhalin, both locating near the eastern coasts of the Asian continent, and 12 specimens of R. ornativentris from Honshu, Japan mainland, as an outgroup, through horizontal starch gel electrophoresis. Rana pirica shows relatively low genetic differentiation, and high genetic affinities are found between Hokkaido and Sakhalin populations. Populations from these islands are morphologically somewhat differentiated but should be regarded as conspecific. Degree of genetic divergence among populations of amphibian species from Hokkaido, including R. pirica , is not so extensive as that reported for species from Japan mainland and relatively recent formation of amphibian fauna in Hokkaido is suggested.     

Allozyme Variation within and between Populations of Coreopsis latifolia (Asteraceae)

Abstract Enzyme electrophoresis was used to measure genetic variation in five populations of the rare diploid species Coreopsis latifolia which appears to be a relict taxon endemic to a small area of the southeastern United States. Gene diversity within the species as a whole is low compared to other species with similar ecological and life history traits. Also, gene diversity in C. latifolia is lower than nearly all other species of Coreopsis that have been examined. Larger populations contain significantly more variation at isozyme loci than do smaller populations. Populations of C. latifolia are deficient in heterozygotes relative to expected equilibrium values.     

Allozyme Variation within and between Populations of Coreopsis intermedia (Asteraceae)

Abstract Enzyme electrophoresis was used to measure genetic diversity in two populations of Coreopsis intermedia of section Coreopsis , a rare species endemic to a small area of Texas and Louisiana. Genetic diversity is higher in C. intermedia than has been reported for many other geographically restricted species. C. intermedia contains levels of genetic variation comparable to those of other species in section Coreopsis , and a greater amount of diversity than either the rare congener, C. latifolia (section Silphidium ), or the Florida endemic, C. leavenworthii (section Calliopsis ). The two populations, although greatly different in size, exhibited similar levels of genetic diversity.     

Latitudinal Variation of Allozyme Frequencies in Populations of Two Freshwater Isopods in Western Europe

Populations of the freshwater isopods, Proasellus meridianus Racovitza and P. coxalis Dollfus were found to be polymorphic at loci coding for the enzymes glucose phosphate isomerase (GPI) and phosphoglucomutase (PGM). PGM allele frequencies in populations of both species were significantly associated with latitude as has been previously reported for the closely related Asellus aquaticus (L.). Co-variation of PGM allele frequencies was also found in sympatric populations of P. meridianus and A. aquaticus. Significant interpopulation variation in GPI allele frequencies occurred in P. coxalis but not P. meridianus, however, this variation was not associated with latitude.     

Correlation between hordatine accumulation, environmental factors and genetic diversity in wild barley (Hordeum spontaneum C. Koch) accessions from the Near East Fertile Crescent

Wild barley shows a large morphological and phenotypic variation, which is associated with ecogeographical factors and correlates with genotypic differences. Diversity of defense related genes and their expression in wild barley has been recognized and has led to attempts to exploit genes from H. spontaneum in breeding programs. The aim of this study was to determine the variation in the accumulation of hordatines, which are Hordeum-specific preformed secondary metabolites with strong and broad antimicrobial activity in vitro, in 50 accessions of H. spontaneum from different habitats in Israel. Differences in the accumulation of hordatines in the seedling stage were significant between different H. spontaneum genotypes from different regional locations and micro-sites. Variation in the hordatine accumulation within genotypes was between 9% and 45%, between genotypes from the same location between 13% and 38%, and between genotypes from different locations up to 121%. Principal component analysis showed that water related factors explain 39%, temperature related factors explain 33% and edaphic factors account for 11% of the observed variation between the populations of H. spontaneum. Genetic analysis of the tested accessions with LP-PCR primers that are specific for genes involved in the biosynthetic pathway of hordatines showed tight correlations between hordatine abundance and genetic diversity of these markers. Multiple regression analyses indicated associations between genetic diversity of genes directly involved in hordatine biosynthesis, ecogeographical factors and the accumulation of hordatines.     

Ecological-genomic diversity of microsatellites in wild barley,Hordeum spontaneum,populations in Jordan

We analyzed the ecological-genomic diversity of microsatellites of wild barley, Hordeum spontaneum (C. Koch) Thell., at 18 loci in 306 individuals of 16 populations from Jordan across a southward transect of increasing aridity. The 18 microsatellites revealed a total of 249 alleles, with an average of 13.8 alleles per locus (range 3-29), with nonrandom distribution. The proportion of polymorphic loci per population averaged 0.91 (range 0.83-1.00); gene diversity, He, averaged 0.512 (range 0.38-0.651). We compared the number of alleles of the 18 loci to those found in Israel populations by Turpeinen et al. Out of the 280 alleles, 138 (49.3%) were unique (i.e. occurred in only one of the countries). The percentage of unique alleles in Jordan and Israel populations was 43.0% and 17.9%, respectively, suggesting that Jordan is an important center of origin and diversity of wild barley. Estimates of mean gene diversity were highest in the populations collected near the Golan Heights, such as Shuni North, Shuni South and Jarash. Sixty nine percent of the microsatellite variation was partitioned within populations and 31% between populations. Associations between ecogeographical values and gene diversity were established for eight microsatellite loci. The cluster produced by simple sequence repeat (SSR) data is mostly coincidence with the result of the dendrogram of the Spalax ehrenbergi superspecies of subterranean mole rats in Jordan based on allozyme gene loci. The major soil type in the wild barley habitat of each ecological group was different. Stepwise multiple regression analysis indicated that the variance of gene diversity was explained by altitude (R(2) = 0.362**). These observations suggest that microsatellites are at least partly adaptive and subject to natural selection.     

Growth characteristics in Hordeum spontaneum populations from different habitats

Hordeum spontaneum shows a large genetic variation and occupies a wide range of different habitats. The aim of this study was to quantify variation in growth characteristics of H. spontaneum from different sites in Israel and to relate this variation to different environmental conditions. To this end, 84 accessions of 21 populations were grown in a growth chamber in near-optimal conditions and a range of physiological, morphological, allocation- related and chemical characteristics were measured. These parameters included rates of photosynthesis, shoot and root respiration, specific leaf area, biomass allocation and seed mass. Averaged over all traits variation explained by differences between populations was 26%, between accessions 21%, whereas that within accessions was 53%. By contrast with most genetic studies, we found variation between populations larger than between accessions. The largest between-population variation (46%) was for morphological traits. In particular, seed mass, leaf thickness and leaf width differed strongly between populations. Variation in growth characteristics between populations was poorly related to mean annual rainfall, mean humidity or January temperature at the sites of origin. We expect that differences between populations to be larger and correlation with environmental parameters stronger in plants grown in stressful conditions. According to our study, seed mass is more important than relative growth rate in determining variation in early plant biomass in H. spontaneum .     

High crossability of wild barley (Hordeum spontaneum C. Koch) with bread wheat and the differential elimination of barley chromosomes in the hybrids

Four bread wheat (Triticum aestivum L.) cultivars, Aobakomugi, Chinese Spring, Norin 61 and Shinchunaga, were pollinated with five barley lines/cultivars consisting of three cultivated barley (Hordeum vulgare L.) lines, Betzes, Kinai 5 and OHL089, and two wild barley (Hordeum spontaneum C. Koch) lines, OUH602 and OUH324. Crossability, expressed as the percentage of embryo formation, varied from 0 to 55.4% among the cross combinations. The two wild barley lines generally had a higher crossability than the previously reported best pollinator, Betzes, and some Japanese wheat cultivars were better as the female parent than Chinese Spring. Ninety four hybrid plants were obtained from 250 embryos cultured, and their somatic chromosome numbers ranged from 21 to 36. Eighteen plants were mosaic in chromosome number. Twenty one-chromosome plants appeared most frequently (45.7%) followed by 28-chromosome plants (14.9%). C-banding analysis revealed that elimination of barley chromosomes was mainly responsible for the occurrence of aneuploid plants. In hypoploids derived from Betzes-crosses, chromosome 5 was preferentially eliminated as previously reported, while in hypoploids derived from OUH602-crosses, chromosome 4 was preferentially eliminated. The wild barley line OUH602 may be a useful parent for producing a new wheat-barley addition set because of its high crossability with wheat and a different pattern of chromosome elimination.