The changes in physiological and biochemical traits of Tibetan wild and cultivated barley in response to low phosphorus stress

Low phosphorus (LP) limits crop growth and productivity in the majority of arable lands worldwide. Here, we investigated the changes in physiological and biochemical traits of Tibetan wild barleys (Hordeum vulgare L. ssp. spontaneum) XZ99 (LP tolerant), XZ100 (LP sensitive), and cultivated barley ZD9 (moderately LP tolerant) under two phosphorus (P) levels during vegetative stage. These genotypes showed considerable differences in the change of biomass accumulation, root/shoot dry weight ratio, root morphology, organic acid secretion, carbohydrate metabolism, ATPase (Adenosine triphosphatase) activity, P concentration and accumulation under LP in comparison with CK (control) condition. The higher LP tolerance of XZ99 is associated with more developed roots, enhanced sucrose biosynthesis and hydrolysis of carbohydrate metabolism pathway, higher APase (Acid phosphatase) and ATPase activity, and more secretion of citrate and succinate in roots when plants are exposed to LP stress. The results prove the potential of Tibetan wild barley in developing barley cultivars with high tolerance to LP stress and understanding the mechanisms of LP tolerance in plants.     

Difference in response to aluminum stress among Tibetan wild barley genotypes

Wild barley (Hordeum sp.) germplasm is rich in genetic diversity and provides a treasure trove of useful genes for crop improvement. We carried out a comprehensive program combining short‐term hydroponic screening via hematoxylin‐staining of root‐regrowth procedure and filter paper–based evaluation of diverse germplasm in response to Al/acid stress using 105 annual Tibetan wild barley and 45 cultivated barley genotypes. Root elongation among the 105 Tibetan wild barley genotypes varied significantly after Al exposure, ranging from 62.9% to 80.0% in variation coefficients and 4.35 to 4.45 in diversity index. These genotypic differences in Al resistance were fairly consistent in both the hydroponic and filter paper–based evaluations: XZ16, XZ166, and XZ113 were selected as Al‐resistant genotypes, and XZ61, XZ45, and XZ98 as Al‐sensitive wild genotypes. Furthermore, significantly lower Al concentrations in roots and shoots were detected in the three selected Al‐resistant genotypes than in the three sensitive genotypes in the filter paper–based experiment. Meanwhile, XZ16 was the least affected by Al toxicity in regard to reduced SPAD value (chlorophyll meter readings), plant height, root length, dry biomass, tillers per plant, and chlorophyll fluorescence (Fv/Fm) in the long‐term hydroponic experiment compared with the Al‐resistant cultivated barley cv. Dayton, while XZ61 had the severest stress symptoms.     

Nitrogen nutrition index and leaf chlorophyll concentration and its relationship with nitrogen use efficiency in barley (Hordeum vulgare L.)

Nitrogen (N) is one of the most important nutrients for barley, and at the same time excessive N fertilization can have a negative impact on the environment. A 2-year field study was conducted with the objective to determine the relationship of N Use Efficiency (NUE) and its components N utilization efficiency (NUtE), and N uptake efficiency (NUpE) with N diagnostic tools N nutrition index (NNI) and chlorophyll meter (CM) readings of barley. CM readings were negatively correlated with NUE and NUpE. NNI varied from 0.79 to 1.00 across years and cultivars. This study provides new information about the effect of N application on NUE and its components and its relationship with CM readings and NNI of barley which can be used for proper N management and protection of the environment.     

Allelic diversity of a beer haze active protein gene in cultivated and Tibetan wild barley and development of allelic specific markers

The formation of haze is a serious quality problem in beer production. It has been shown that the use of silica elute (SE)-ve malt (absence of molecular weight (MW) ～14000 Da) for brewing can improve haze stability in the resultant beer, and the protein was identified as a barley trypsin inhibitor of the chloroform/methanol type (BTI-CMe). The objectives of this study were to determine (1) the allelic diversity of the gene controlling BTI-CMe in cultivated and Tibetan wild barley and (2) allele-specific (AS) markers for screening SE protein type. A survey of 172 Tibetan annual wild barley accessions and 71 cultivated barley genotypes was conducted, and 104 wild accessions and 35 cultivated genotypes were identified as SE+ve and 68 wild accessions and 36 cultivated genotypes as SE-ve. The allelic diversity of the gene controlling BTI-CMe was investigated by cloning, alignment, and association analysis. It was found that there were significant differences between the SE+ve and SE-ve types in single-nucleotide polymorphisms at 234 (SNP(234)), SNP(313), and SNP(385.) Furthermore, two sets of AS markers were developed to screen SE protein type based on SNP(313). AS-PCR had results very similar to those obtained by immunoblot method. Mapping analysis showed that the gene controlling the MW～14 kDa band was located on the short arm of chromosome 3H, at the position of marker BPB-0527 (33.302 cM) in the Franklin/Yerong DH population.     

野生大麦氮素吸收利用的基因型差异

采用土培盆栽,研究16份野生大麦在同一低氮处理下全生育期氮素吸收利用的基因型差异,旨在筛选出具有高效利用氮素能力的基因型,为氮素高效利用大麦育种提供种质材料。结果表明：（1）生物量、氮含量和氮素利用效率的基因型差异最大都在抽穗期（CV=33.55%,18.93%,21.88%）,氮含量和氮素利用效率基因型差异拔节期最小...     

Phenotypic diversity in wild barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis> L. ssp. <Emphasis Type="Italic">spontaneum</Emphasis> (C. Koch) Thell.) accessions collected in Jordan

Wild barley, Hordeum vulgare L. ssp. spontaneum (C. Koch) Thell., is the progenitor of cultivated barley. Almost unanimously the center of diversity is considered to be in the Fertile Crescent of the Near East, where wild barley grows under a wide range of environmental and climatic conditions. Jordanian wild barley is expected to harbor genes useful for the improvement of cultivated barley, particularly those associated with tolerance to drought. This study evaluated 103 wild barley accessions collected from different areas of Jordan along with 29 cultivated barley genotypes for several morphological and agronomical traits. The Hordeum vulgare ssp. spontaneum C. Koch accessions were grouped into six populations according to the longitude, latitude, altitude, and rainfall zone of the collection site, and the cultivated barley in one population. The evaluation was conducted during the 2004–2005 growing season under field conditions in three locations in Jordan; namely, Khanasri, Ramtha, and Maru with 123.0, 222.9, and 429.2 mm annual rainfall, respectively. We used an unreplicated design with two systematic checks (the cultivars Rum and Mu’ta) each repeated 15 times. The results showed the existence of high variability among the Hordeum vulgare ssp. spontaneum C. Koch accessions for most of the traits, especially for plant height, tiller number, days to heading, days to anthesis, peduncle length, and peduncle extrusion. Plant height, earliness, peduncle length, and peduncle extrusion were found to be adaptive traits under drought conditions and several superior genotypes for each trait were identified. Genetic variation within population was much higher than between populations. Clustering of populations was according to their ecological geographical pattern.     

Identification and selection of low-phosphate-tolerant germplasm in barley (Hordeum vulgare L.)

Phosphorus (P) deficiency is one of the major constraints to crop yield worldwide, and genotypes or cultivars with high phosphate use efficiency (PUE) sustain growth when exposed to phosphate stress. Therefore, it is imperative to develop the genotypes or cultivars with high PUE. A pot experiment was conducted to evaluate the PUE among 150 barley (Hordeum vulgare L.) genotypes. Two high-tolerant and -sensitive accessions were selected. These two candidate materials were used to investigate the differences among the root morphology characteristics, antioxidant enzyme activity, inorganic phosphate (Pi) content and gene expression of HvPT5 under P-deficiency and P-sufficiency conditions. The values of these parameters were higher in the low-P-tolerant genotype than in the sensitive one. In pot experiment 1, all genotypes showed a significant difference in low-P tolerance, with variety GN121 achieving the highest tolerance, and GN42 being most sensitive. The results of this study may provide elite genetic germplasms for future work on isolation of P-related genes, and the improvement of PUE in barley.

Abbreviations: PUE: phosphate use efficiency; CAT: catalase; POD: peroxidase; SOD: superoxide dismutase; DMSO: dimethyl sulphoxide; MDA: malondialdehyde; TOPSIS: technique for order preference by similarity to an ideal solution; MCDM/MADM: multi-criteria (or attribute) decision making     

Molecular analysis of the DNA polymorphism of wild barley (Hordeum spontaneum) germplasm using the polymerase chain reaction

Summary Hordeum spontaneum is the progenitor of cultivated barley (H. vulgare) and is an important source of genetic variation for barley breeding programs. The genetic diversity ofH. spontaneum in the Australian germplasm collection was investigated using the polymerase chain reaction with random and semi-random primers. This approach was found to be robust in respect of reaction conditions. Genetic dissimilarity values between genotypes were used to produce a phenogram of the relationships among the accessions using the unweighted pair group method with arithmetic mean. The largest divergence was observed among Israeli accessions, whereas the Turkish and Iranian samples clustered as distinct subsets, each apparently related to portion of the Israeli material. The results indicate that the genetic diversity of the wild barleys is broadly correlated with geographic distribution.     

Monitoring the photosystem II behavior of wild and cultivated barley in response to progressive water stress and rehydration using OJIP chlorophyll a fluorescence transient

To study the effects of progressive drought stress on photosystem II behavior of wild type (Spantaneum) and cultivated barley (Morocco), different levels of soil water availability, including control, moderate, mild, and severe water stress (70%, 50%, 30%, and 10% water holding capacity of soil, respectively) and rehydration were used. Polyphasic OJIP fluorescence transient of Morocco plants exhibited a considerable increase in fluorescence intensity at O, J, and I steps under mild and severe stress relative to slight increase in wild barley. Values of fluorescence parameters and quantum efficiencies, including minimal fluorescence, relative variable fluorescence at phase J and I, maximal quantum yield of photosystem II (PSII), performance index, electron transport yield, and excitation transfer efficiency were influenced by water stress in both genotypes. These parameters were significantly less affected in wild type barley by progressive drought stress compared to Morocco. After re-watering, both genotypes were able to restore from severe drought in most of the traits. Based on our findings, highly correlated values of relative water content (RWC) and independent JIP-test parameters (P < 0.01) indicate that the chlorophyll a fluorescence induction technique is sensitive to plant water status and performance index represent an accurate and reliable indicator for early stress detection and also explore plant vitality under water stress.     

不同基因型油菜氮素利用效率的差异及其与农艺性状和氮素营养性状的关系

通过田间试验对50个甘蓝型油菜基因型进行氮利用效率筛选,并将不同氮利用效率基因型在施氮（N 180 kg/hm2）和不施氮（N 0 kg/hm2）条件下的农艺性状和氮营养性状作了比较研究。结果表明,50个油菜基因型的氮利用效率具有显著差异,最大相差2.5倍（施氮）和1.7倍（不施氮）。油菜的农艺性状及氮素营养性状指标均表现出一定的基因型变异,其中第一个有效分枝高度变异系数最大,超过 50%,而籽粒含氮量的变异系数最小,只有 6% 左右。不论供氮水平高低,氮高效基因型的每角粒数,籽粒生物量,籽粒占地上部生物量的比例以及籽粒氮占地上部总氮的比例均高于氮低效基因型,第一个有效分枝高度则显著低于氮低效基因型。另外,油菜氮素利用效率与各器官含氮量、 茎叶氮素累积量、 果荚氮素累积量、 茎叶氮占地上部氮的比例和果荚氮占地上部氮的比例呈显著或极显著负相关,而不同氮效率基因型的地上部总生物量和地上部氮素累积总量则无显著差异。不同氮利用效率基因型的农艺性状和氮营养性状对氮肥的敏感性有显著差异。     

不同氮效率油菜SPS和PEPC活性差异及其对籽粒产量与油分含量的影响

【目的】蔗糖磷酸合成酶(sucrose phosphate synthase,SPS)与磷酸烯醇式丙酮酸羧化酶(phosphoenolpyruvate carboxylase,PEPC)分别控制着植物体内的碳骨架向碳代谢和氮代谢的流转,影响作物的产量与品质。为探明氮高效油菜品种在高效利用氮素的同时协调籽粒蛋白与油分累积矛盾的机理,研究了不同氮效率油菜品种的SPS与PEPC活性差异及其对籽粒油分含量的影响。【方法】采用土培试验,以氮高效品种27号(H)与氮低效品种6号(L)为试验材料,在常氮(N)与低氮(S)条件下,研究不同氮效率油菜品种苗期到花期叶片与角果发育初期的角果、角果发育中期的角果皮与籽粒中SPS与PEPC活性变化及差异、生长后期碳素转运量与转运率以及收获期籽粒油分含量的差异。【结果】两种供氮水平下,氮高效品种27号的产量与籽粒油分含量均显著高于氮低效品种6号,品种优势明显;且氮高效品种27号苗期到花期叶片与角果发育初期的角果、角果发育中期的角果皮与籽粒中的SPS与PEPC活性均高于氮低效品种6号,两种供氮水平的规律相同,但是SPS与PEPC活性的比值(SPS/PEPC)却因生育期不同而异,营养生长期叶片中氮高效品种27号的SPS/PEPC高于氮低效品种6号,开花期品种间叶片SPS/PEPC相近,角果发育期主要生殖器官中的SPS/PEPC值氮高效品种反而低于氮低效品种。说明氮高效品种向碳代谢和氮代谢输送的碳骨架在全生育期均多于氮低效品种,而碳代谢对氮代谢的响应只在生育前期强于氮低效品种,生育后期则相反。碳素转运量与转运率、籽粒油分含量与产量也是氮高效品种大于氮低效品种,这可能为氮高效品种协调籽粒蛋白与油分累积矛盾的重要生理机制。供氮水平对上述各指标有不同的影响,籽粒产量、PEPC活性、碳素转运量及转运率以常氮处理高于低氮处理,而油分含量、SPS活性及SPS/PEPC以常氮处理低于低氮处理,但不改变以上指标的品种间差异。【结论】与氮低效品种相比,氮高效品种全生育期向碳、氮代谢均输送更多的碳骨架,这是氮高效品种缓解碳、氮代谢矛盾的重要前提;碳代谢对氮代谢的响应生育前期较高、生育后期较低,同时生育后期有更多营养器官的碳素转运到籽粒,也为油菜生育后期满足籽粒碳、氮代谢所需要的碳骨架,并协调籽粒油分与蛋白质含量的矛盾提供了条件。     

Genetic diversity in wild barley (Hordeum spontaneum C. Koch) in the Near East: a molecular analysis using Random Amplified Polymorphic DNA (RAPD) markers

We analyzed the genetic diversity in 88 genotypes from 20 populations of wild barley (Hordeum spontaneum C. Koch) from Israel, Turkey and Iran, by randomly amplified polymorphic DNA (RAPD). Twenty two of the 33 primers used yielded scorable products with 1–11 polymorphic bands. No duplicate patterns were found except for four haplotypes.When the total genetic diversity was estimated, 75% of the variation detected was partitioned within the 88 genotypes and 25% among the populations. When variation between countries was assessed, no substantial differences were found, because most of the variation detected (97%) was partioned within the 20 populations and the remainder among countries. The results of this limited survey indicate that the extensive genetic diversity is present in natural stands of wild barley throughout the Fertile Crescent.     

Leaf photosynthesis,chlorophyll fluorescence and ion content of barley (Hordeum vulgare) in response to salinity

Eight barley genotypes contrasting in their salinity tolerance were assessed for their chlorophyll fluorescence, photosynthetic performance, lipid peroxidation level and ionic content. A pot experiment was conducted in Borj-Cedria (Tunisia), in a wire house with a glass roof to avoid rainfall. The assay consisted of three treatments (0, 100 and 200 mM NaCl) with eight barley genotypes following a completely randomized design. Each pot was considered as one replicate and nine replicates were used for each genotype and treatment. The salt-tolerant genotypes Kerkna and Tichedrett recorded the highest tolerance for photosynthesis and potassium accumulation, whereas the susceptible genotypes were mostly affected at severe salinity. Contrasting behavior was noted for fluorescence attributes, while PSII yield was unchanged reflecting good protection against photodamage. Photosynthetic performance, enhanced water use efficiency, maintained leaf K⁺ and oxidative defense remain the key components for tolerance mechanisms. Salt-tolerant barley could be suitable for management of salt-affected soils.     

Differences in phytase activity and phytic acid content between cultivated and Tibetan annual wild barleys

The Qinghai-Tibetan Plateau in China is considered to be one of the original centers of cultivated barley. At present, little is known about the phytase activity (Phy) or phytic acid content (PA) in grains of Tibetan annual wild barley. Phy and PA were determined in grains of 135 wild and 72 cultivated barleys. Phy ranged from 171.3 to 1299.2 U kg(-1) and from 219.9 to 998.2 U kg(-1) for wild and cultivated barleys, respectively. PA and protein contents were much higher in wild barley than in cultivated barley. Tibetan annual wild barley showed a larger genetic diversity in phytase activity and phytic acid and protein contents and is of value for barley breeding. There is no significant correlation between phytase activity and phytic acid or protein content in barley grains, indicating that endogenous phytase activity had little effect on the accumulation of phytic acid.     

Solute adjustment to salinity and calcium supply in cultivated and wild barley

Ion relations, water content, leaf water potential, and osmotic adjustment were determined for cultivated barley (cv Harrington) and wild barley grown under mixed sulphate (SO₄) salts with varied calcium (Ca) supply using a hydroponic system. Salinity induced significant increases of leaf, stem and root sodium (Na) concentrations in both species. Salt‐stressed wild barley roots accumulated more Na than shoots, and transport of Na from roots to shoots was low compared to Harrington. Cultivated barley had lower Ca concentrations than wild barley, especially in the low Ca salt treatment. Although potassium (K/Na) and Ca/Na ratios were higher in control wild barley plants than in Harrington, they declined under salt stress, irrespective of Ca supply. Major osmotica in wild barley leaves were K, sugars, organic acids, and quaternary ammonium compounds, while in Harrington they were cations, including Na, K and Mg, and anions such as phosphate (PO₄) and SO₄. Wild barley maintained better water status than Harrington under low Ca salt treatment. Supplemental Ca improved water status more in Harrington than in wild barley. Lack of osmotic adjustment to salinity in wild barley apparently resulted from its ion exclusion. Low Ca salt treatment caused Ca deficiency, Na toxicity, and loss of turgor in Harrington. In the high Ca salt treatment, Harrington had improved water and ion relations, as well as positive turgor.     

Genetic diversity of Acid-PAGE monomeric prolamins in cultivated hulless barley (Hordeum vulgare L.) from Qinghai–Tibet plateau in China

Naked barley, due to its favorable attributes such as high feed value, good human nutrition and easy processing, increasingly attracts people’s attention. Qinghai–Tibet Plateau is very rich in naked barley resources and has a long growing history. Genetic diversity of these cultivated naked barley is, however, poorly documented. This study analyzed the genetic diversity of monomeric prolamins (protein fraction corresponding to wheat gliadins) using the Acid -PAGE technique in eighty-six cultivated naked barley cultivars from Qinghai–Tibet Plateau. Extensive diversity was observed. A total of 43 different bands and 76 distinct patterns were found. Jaccard’s coefficient of similarity was calculated, and the accessions were divided into five main groups by cluster analysis using UPGMA. Differentiation among the populations from different collecting regions was investigated, based on the polymorphism of monomeric prolamins. The genetic diversity within the accessions from Tibet was slightly higher with an average diversity index of 0.29 than those from Sichuan. This study supports Tibetan Plateau is the diversity center of the cultivated naked barley and suggests that cultivated naked barley from Qinghai–Tibet Plateau is an important pool of variability that could be used in cereal breeding.     

Durum Wheat Genotypic Variation of Yield and Nitrogen use Efficiency and Its Components Under Different Water and Nitrogen Regimes in the Mediterranean Region

Farmers in dry areas of the Mediterranean region are reluctant to apply nitrogen (N) fertilizer to rainfed wheat because of frequent drought. So, it is desirable to select varieties with high nitrogen use efficiency (NUE). The objectives of this study, conducted in 2009/10 and 2010/11 in Syria, were to study the response of genotypes of durum wheat to low and high N applications and to evaluate the contribution of N uptake efficiency and utilization efficiency to NUE under rainfed and supplemental irrigation conditions. Under the rainfed regime, grain yield decreased significantly in year 1 and year 2 when applying N fertilizer at a high rate. The early maturing genotypes gave the highest average yields under rainfed and irrigated regimes. High N fertilizer rate decreased significantly NUE from 36.1 to 24.3 in year 1 and 37.0 to 5.8 in year 2. Under irrigation, NUE fell from 84.6 to 67.1 in year 1 and from 117.7 to 33.3 in year 2. The contribution of N uptake efficiency and utilization efficiency to NUE varied from one year to another. In year 1, the contribution of N utilization efficiency was more important at all nitrogen levels; while the opposite was observed in year 2 when more N was applied. The fractions of the genotype sum of squares, respectively, for N uptake efficiency and utilization efficiency were in average 0.15 and 0.78 in year 1 and 0.75 and 0.25 in year 2. From this study we conclude that high N levels in the soil exacerbate the effect of water stress on productivity and NUE of wheat. Early maturing new genotypes tend to be better adapted and to use nitrogen more efficiently under limited water conditions. N utilization efficiency contributes more to NUE under high N availability than N uptake efficiency and vice-versa.     

ADAPTABILITY MECHANISM OF NITROGEN-EFFICIENT GERMPLASM OF NATURAL VARIATION TO LOW NITROGEN STRESS IN BRASSICA NAPUS

Previous studies showed that wide genotype differences in nitrogen (N) efficiency exists among cultivars of rapeseed (Brassica napus L.), but the mechanisms behind those differences are still unknown. In the present study, our aim was to analyze the adaptability mechanism of N-efficient rapeseed to low-N stress by employing two genotypes of natural variation in N efficiency. Nitrogen-efficient genotype, ‘BG51’, and N-inefficient genotype, ‘BG88’, were grown in a solution culture experiment under conditions of high-N (6.0 mM N) and low-N (0.6 mM N) supply. After growing 30 d, roots and shoots were sampled for the analysis of dry weight, N concentration and accumulation, N use efficiency (NUE), N transport efficiency (NTE), root system vigor parameters, nitrate redutase (NR) activity, and glutamine synthetase (GS) activity. Nitrogen deficiency decreased shoot and root dry weight significantly, but ‘BG51’ exhibited a significantly lower decrease in shoot dry weight and had significantly higher biomass production than ‘BG88’. Under low N supply ‘BG51’ accumulated more N in shoot, root and whole plant than ‘BG88’, and presented higher NUE in both shoot and root. Low-N stress induced an increase in maximum root length by 28.3% for ‘BG88’ and 55.1% for ‘BG51’ compared with the high-N treatment. And ‘BG51’ presented larger root volume, higher root vigor, larger root total absorbing area and root active absorbing area than ‘BG88’ in low-N treatment. Furthermore, ‘BG51’ had significantly higher NR and GS activity in both leaf and root in low N treatment than ‘BG88’, while there was no evident difference between them in high N treatment. These results suggested that N-efficient rapeseed germplasm of natural variation involves an integrated adaptability mechanism responding to low-N stress. Namely, N-efficient genotype could form more developed root system to accumulate more N, and presented efficient N assimilation by higher NR activity and GS activity than N-inefficient genotype. These ultimately resulted in high tolerance of N-efficient genotype to low-N stress and high biomass production.     

不同品种油菜氮效率差异及其生理基础研究

采用盆栽土培试验,以氮胁迫与正常供氮条件下的子粒产量比值作为氮效率系数,探讨了不同品种油菜氮效率差异及其生理基础。结果表明,供试5个油菜品种的氮效率有显著差异,其氮效率系数的变化范围是0.37~0.69。在氮胁迫条件下,氮高效品种的植株含氮量与氮素累积吸收量、叶片叶绿素含量与硝酸还原酶活性、茎叶可溶性糖含量与硝态氮再利用量高于氮低效品种。在正常供氮条件下,高潜力品种的植株含氮量低于低潜力品种,但由于其生物量较高,氮素累积吸收量并不低于低潜力品种,甚至苗期的氮素累积吸收量高于低潜力品种;高潜力品种的叶片叶绿素含量与硝酸还原酶活性、茎叶可溶性糖含量与硝态氮再利用量高于低潜力品种。说明上述4种生理指标均可作为评价油菜氮效率及增长潜力差异的间接指标。     

Ecological correlates of RAPD DNA diversity of wild barley, Hordeum spontaneum, in the Fertile Crescent

Genetic variability in RAPDs (Randomly Amplified Polymorphic DNA) was studied in 104 genotypes of wild barley, Hordeum spontaneum from 21 populations sampled in Israel, Turkey and Iran, seven population from each country. The band (= loci) frequencies were calculated for each population and correlated with ecogeographical variables. In general, high RAPD genetic diversity indices were associated with stressful environments, either with hot or cold steppes and deserts. Interpopulational genetic distances showed no association with the geographic distance between the populations' provenance. Significant Spearman rank correlations between RAPD band frequencies and ecogeographical parameters of provenance occured. Frequencies of RAPD bands were significantly correlated with the principal component factors of allozymes. The correlation data indirectly suggest that natural selection appears to be the major determinant of both RAPD and allozyme diversities both being correlated with environmental stress.