Mapping quantitative trait loci for salt tolerance at germination and the seedling stage in barley (Hordeum vulgare L.)

Quantitative trait loci (QTLs) controlling salt tolerance at germination and the seedling stage in barley (Hordeum vulgare L.) were identified by interval mapping analysis using marker information from two doubled haploid (DH) populations derived from the crosses, Steptoe/Morex and Harrington/TR306. Interval mapping analysis revealed that the QTLs for salt tolerance at germination in the DH lines of Steptoe/Morex were located on chromosomes 4 (4H), 6(6H), and 7(5H), and in the DH lines of Harrington/TR306 on chromosomes 5(1H) and 7(5H). In both DH populations, the most effective QTLs were found at different loci on chromosome 7(5H). Genetic linkage between salt tolerance at germination and abscisic acid (ABA) response was found from QTL mapping. The QTLs for the most effective ABA response at germination were located very close to those for salt tolerance on chromosome 7 (5H) in both crosses. The QTLs for salt tolerance at the seedling stage were located on chromosomes 2(2H), 5(1H), 6(6H), and 7(5H) in the DH lines of Steptoe/Morex, and on chromosome 7(5H) in the DH lines of Harrington/TR 306. Their positions were different from those of QTLs controlling salt tolerance at germination, indicating that salt tolerance at germination and at the seedling stage were controlled by different loci. This revised version was published online in July 2006 with corrections to the Cover Date.     

不同磷水平下大麦分蘖期磷效率相关性状QTL定位分析

磷素营养与大麦品质及产量密切相关,磷高效遗传机制和品种改良是近年的研究热点之一。本研究利用由大麦栽培品种Baudin和种质材料CN4079杂交构建的重组自交系(RIL)群体,低磷胁迫(0.02 mmol L~(-1) KH_2PO_4)与正常供磷(0.2 mmol L~(-1) KH_2PO_4)条件下,对地上部和地下部磷素利用效率、磷素吸收效率和干重,以及分蘖数相关的QTL定位,并预测相关位点基因。表型鉴定结果表明,各性状在RIL群体中表现连续变异,并存在超亲分离。两种磷水平下,共检测到16个QTL,分布在2H、3H和5H染色体上,表型贡献率14.1%~28.5%。3H染色体上含有3个磷素利用效率位点,其增效等位基因均来源于Baudin,其中Qspue.sau-3H.1和Qrpue.sau-3H与控制磷素吸收效率的Qspae.sau-3H和Qrpae.sau-3H处于同一区段,而Qspue.sau-3H.2与控制分蘖数的位点Qtn.sau-3H处于同一区段。5H染色体上含有3个磷素吸收效率位点,其中Qspae.sau-5H.2和Qrpae.sau-5H的增效等位基因来自CN4079,且与控制磷素利用效率的Qspue.sau-5H和Qrpue.sau-5H,以及控制干重的Qsdw.sau-5H和Qrdw.sau-5H处于同一区段。在磷效率相关的4个区段中,除Qspue.sau-3H.1所处区间仅含有磷酸代谢与磷脂代谢相关基因外,其他区间均包含磷酸盐转运蛋白基因、磷酸代谢与磷脂代谢相关基因。     

Adaptation of barley (Hordeum vulgare L.) to harsh Mediterranean environments

Summary Differences in development of the apex may be the reason for cultivar differences in adaptation of barley to terminal drought in Mediterranean environments. The present study was conducted to identify apical development patterns of barley adapted to terminal drought stressed Mediterranean environments and to determine plant characteristics which can be used as criteria to select for an adapted development. Thirty-five two-rowed barley (Hordeum vulgare L.) entries were grown at two sites in northern Syria (Tel Hadya and Breda) in 1988/89. Four apical development patterns were observed: a slow or fast vegetative development, depending on the vernalization requirement, combined with a slow or fast generative development, depending on the daylength response of the crop. Early heading was related to fast generative development. Leaf appearance rates on the main shoot were constant during a major part of the pre-anthesis period, but significant differences were observed among development patterns. Genotypic differences in main shoot tiller number were associated with differences in the onset of tiller appearance and not with differences in tiller appearance rate or final leaf number on the main shoot. Since vernalization requirements and daylength responses are largely independent of terminal drought stress, selection for an adapted phenology can be done in favourable environments. Morphological traits related to these responses (winter growth habit, cold tolerance, plant colour, growth vigour, heading date) can be used as criteria for selection for adaptation to low-rainfall Mediterranean environments.     

Genomewide association analysis of salt tolerance in soybean [Glycine max (L.) Merr.]

Salinity is a common abiotic stress causing soybean [Glycine max (L.) Merr.] yield loss worldwide. The use of tolerant cultivars is an effective and economic approach to coping with this stress. Towards this, research is needed to identify salt‐tolerant germplasm and better understand the genetic and molecular basis of salt tolerance in soybean. The objectives of this study were to identify salt‐tolerant genotypes, to search for single‐nucleotide polymorphisms (SNPs) and QTLs associated with salt tolerance. A total of 192 diverse soybean lines and cultivars were screened for salt tolerance in the glasshouse based on visual leaf scorch scores after 15–18 days of 120 mM NaCl stress. These genotypes were further genotyped using the SoySNP50K iSelect BeadChip. Genomewide association mapping showed that 62 SNP markers representing six genomic regions on chromosomes (Chr.) 2, 3, 5, 6, 8 and 18, respectively, were significantly associated with salt tolerance (p < 0.001). A total of 52 SNP markers on Chr. 3 are mapped at or near the major salt tolerance QTL previously identified in S‐100 (Lee et al., 2014). Three SNPs on Chr. 18 map near the salt tolerance QTL previously identified in Nannong1138‐2 (Chen, Cui, Fu, Gai, & Yu, 2008). The other significant SNPs represent four putative minor QTLs for salt tolerance, newly identified in this study. The results above lay the foundation for fine mapping, cloning and molecular breeding for soybean salt tolerance.     

A skeletal linkage map of Hordeum bulbosum L. and comparative mapping with barley (H. vulgare L.)

A restriction fragment length polymorphism (RFLP) based linkage map of a cross between two diploid Hordeum bulbosum (2n = 2x = 14) clones, PB1 and PB11, was constructed from 46 recombinant progeny clones. Since both parents are heterozygous, separate and combined parental maps were constructed. All of the RFLP markers screened had previously been mapped in barley (H. vulgare L.) so that comparative maps could be produced. The PB1 linkage map consists of 20 RFLP marker loci assigned to four linkage groups covering 94.3 cM. The PB11 linkage map consists of 27 RFLP marker loci assigned to six linkage groups covering 149.1 cM. Thirteen markers polymorphic in both parents were used as ‘anchors’ to create a combined linkage map consisting of 38 loci assigned to six linkage groups and covering a genetic distance of 198 cM. Marker order was highly conserved in a comparison with the linkage map of H. vulgare (Laurie etal., 1995). However, in contrast, the genetic distances for the same markers were very different being 649 cM and 198 cM respectively, a genetic distance ratio of 1: 3.3. Thus although the map was short, it can be presumed to cover half the genome of H. bulbosum. This study provides further confirmation of the close relationship between the two species and gives a basis for the development of marker mediated introgression through interspecific hybridisation between the two species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Variation in developmental patterns of wild barley (Hordeum spontaneum L.) and cultivated barley (H. vulgare L.)

Summary Six wild barley (Hordeum spontaneum) accessions, from a diverse range of habitats, and two spring-cultivated barleys, were examined for variation in durations of development phases. The durations of the leaf initiation and spikelet initiation phases were longer and spikelet growth phases shorter, in wild than in cultivated barley. Across all wild and cultivated barleys the rate and duration of spikelet initiation were negatively correlated, but neither was related to the number of spikelets per spike. The number of spikelets was positively correlated with the number of leaves and the ratio of the number of spikelets to the number of leaves declined with increasing time to anthesis, indicating that each successive leaf was associated with a diminishing increase in the number of spikelets. The duration of culm elongation and final culm length were shorter in accessions of cultivated barley compared with wild barley. This paper also discusses the feasibility for increasing the number of spikelets per spike through breeding for genetic changes in lengths of pre-anthesis phases of development.Abbreviations ANOVA Analysis of variance - HV Hordeum vulgare - CE Culm elongation - DR Double ridge - HS Hordeum spontaneu - ° Cd Degree days     

The grain yield of uniculm barley (Hordeum vulgare L.) in two contrasting environments

Summary Four six-rowed uniculm lines of spring barley were grown in two yield trials at different sowing densities. One trial was grown on light sandy soil in 1990. The second trial was grown on clay soil in 1991 and included the two-rowed, tillering cultivar Golf. In the trial on sandy soil, the grain yield of the uniculm lines approached that of Golf grown in an adjacent trial, but in 1991 when the growing conditions were more favourable, Golf yielded significantly more than the uniculm lines. Uniculm lines apparently perform relatively better under marginal growing conditions than in high yielding environments.The uniculm lines do not conform to the ideotype proposed for wheat by Donald (1968) but the results show that a drastic change in plant type need not to imply a large drop in grain yield.     

The effect of anther orientation on microspore-derived plant production in barley (Hordeum vulgare L.)

Summary The effect of anther orientation on culture response has been examined in a range of barley cultivars of agronomic importance. Anther orientation did not significantly affect the percentage of anthers responding. However, orientation of the anthers in the up position with one lobe in contact with the medium significantly increased the number of green plants produced when compared to anthers cultured in the flat position. Green plantlet production was observed directly from embryoids i.e. without an intermediate callus phase. Cytological examination of the regenerants indicated that the majority of the plantlets possessed the normal barley karyotype. The implications of these results for Barley breeding are discussed.     

Effects of plot type on detection of quantitative-trait-locus effects in barley (Hordeum vulgare L.)

Hill plots offer advantages in testing large germplasm arrays with limited area and seed. However, the relationship of trait expression between hill vs. row plots remains in question. The development of quantitative-trait-locus (QTL) analysis offers alternative methods of testing hypotheses regarding levels of coincident gene effects in divergent plots. The objective of this study was to compare hill and row plots in terms of QTL detection, number of common QTLs and plot-type-specific QTLs for various characters in barley. Assessments were based on sets of 100 F₁-derived doubled haploid progeny from two environments of hill-plot data and two sets of multiple-environment row-plot data. Common large-effect QTLs for height, heading date, and 1000-kernel weight were detected in both plot types. Fewer QTLs were detected in the hill-plot data, with only one QTL detected for grain yield. There were no hill-plot-specific QTL effects. Hill plots appear to be best suited to highly heritable characters such as height, heading date, and 1000-kernel weight. Evaluation of grain yield is best deferred until larger experimental units can be employed.     

Cytogenetic variation in somatic tissue cultures and regenerated plants of barley (Hordeum vulgare L.)

Summary Chromosome number of morphogenic and non-morphogenic calli and regenerated plants of barley were determined. Cultures were obtained from two kinds of explants, immature embryos and seedling leaves from three cultivars, Ingrid, Dissa and Golden Promise. Callus chromosome analyses were carried out during a 12 month period in a medium containing 2 mg/l of 2,4-D. Diploid cells were predominant in all cases; although in leaf-derived cultures, retraploid cells (2n=4x=28) showed a tendency to increase as time in culture increased and after more than six months in culture, diploid cells decreased to percentages of almost 70%. Aneuploid cells were generally infrequent in all cases. The source of explant has been more important than the genotype (cultivar) and the type of callus (morphogenic vs. non-morphogenic) in the chromosomal stability of cultures as time increases. From short term cultures, only 1.85% of the regenerated plants were tetraploid, the remaining were diploids. The ability of morphogenic calli to regenerate plants decreased before any significant reduction of diploid cells were observed.     

QTL analysis of salt tolerance in Sorghum bicolor during whole-plant growth stages

To study the salt tolerance genetics of sorghum, 181 recombinant inbred lines (RILs) were used to locate quantitative trait loci (QTLs) underlying salt stress adaptability. Six traits, namely, plant height (PH), stem diameter (SD), total biomass (TB), stem fresh weight (SFW), juice weight (JW) and Brix, were investigated under normal and salt stress conditions in two years. A total of 53 QTLs for the six traits under both conditions and their corresponding salt tolerance index (STI) were detected and phenotypic variation explained (PVE) ranged from 4.16% to 20.42%. Six of the QTLs, qTB6, qSFW9, qJW9, qBrix2, qBrix10 and qSTI-Brix9, were the main effect QTLs controlling salt tolerance and had a PVE more than 10%. qSFW9 and qJW9 colocalized in the same marker interval as SB5069-UGSM18 and had PVEs of 17.70% and 14.20%, respectively, with positive effects from L-Tian. QTL clusters controlling PH, TB, STI-TB, SFW and JW were consistently mapped in the marker interval of Xcup19-SB4177 on chromosome 7. These locations might serve as target sites for marker-assisted selection (MAS) in improving salt tolerance of sorghum.     

The influence of genotype and environment on seed and embryo development in barley (Hordeum vulgare L.) after crossing with Hordeum bulbosum L.

Summary Seed and embryo development was studied in crosses between H. bulbosum and the barley genotypes VK 16032, Vada and Vogelsanger Gold and subsequently the F₁ and F₂ progenies from VK 16032 x Vada and VK 16032 x Vogelsanger Gold. Both seed and embryo development are strongly influenced by the barley genotype. Favourable environmental conditions can promote seed and embryo development in genotypes with existing good characters. Dominant inheritance for good seed development and incomplete dominance for large embryos is evident. Linkage between small embryo size and winter habit exists in Vogelsanger Gold.     

Segregation distortion in doubled haploid lines of barley (Hordeum vulgare L.) detected by simple sequence repeat (SSR) markers

A total of 147 simple sequence repeat (SSR) markers (including86 barley and 61 wheat microsatellite markers) were tested for their segregation in a doubled haploid (DH) and an F₂ population of barley. The DH population consisted of 71 doubled haploid lines, developed from F₁ plants of a cross between Tadmor and WI2291using isolated microspore culture technique. A genetic linkage map consisting of 43 microsatellite markers was constructed using the DH population. Particularly on chromosome 4H microsatellite markers showed distorted segregation ratios. Segregation of DH lines based on molecular markers were compared with segregation of 92 F₂ lines from the same cross. The proportion of loci deviating from the expected monogenic segregation ratios in the DH population was significantly higher (19/43loci, 44%) than in the F₂ population (7/43 loci, 16%). The deviation was biased towards the WI2291 parent alleles. In line with this observation, WI2291 was found to perform better than Tadmor in regenerating green plantlets with the isolated microspore-culture technique. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evaluation of fineness of wrinkles on husks of malting barley (Hordeum vulgare L.) by texture analysis of digital image data

Summary It has been suggested that the selection based on the wrinkles on the husks of malting barley is useful especially during the early generation in breeding program, because the wrinkles can be an indicator of malting quality of grains. However, it can be done only visually and requires much labor. In this study, the evaluation of the wrinkles by texture analysis of digital image data, was examined for the quantitative evaluation of the fineness of the wrinkles and its automatic selection. The texture analysis based on co-occurrence matrices was applied and it was revealed that finely wrinkled grains can be clearly discriminated by discriminant analysis on several parameters estimated from the co-occurrence matrices.     

Association of molecular markers with cold tolerance and green period in zoysiagrass (Zoysia Willd.)

Cold tolerance and the green period are key traits in the breeding of zoysiagrass (Zoysia Willd.). Identification of molecular markers associated with cold tolerance and the green period of zoysiagrass will contribute to efficient selection of elite cultivars. These two traits were measured in 96 zoysiagrass accessions in 2004 and 2005–2006, respectively. The mapping population was screened with 29 pairs of simple sequence repeat (SSR) primers and 54 pairs of sequence-related amplified polymorphism (SRAP) primers. A multi-loci in silico mapping approach implemented with an empirical Bayes method was applied for association mapping of cold tolerance and green period. We detected 254 SSR polymorphic loci and 338 SRAP polymorphic loci, among which three SSR loci (Xgwm131-3B-187, Xgwm469-6D-194 and Xgwm234-5B-244) and one SRAP locus (Me11Em7-406) were significantly associated with cold tolerance with effect values of 57.83%, 38.05%, 36.92% and 37%, respectively. Three SSR loci (Xgwm132-6B-225, Xgwm111-7D-34 and Xgwm102-2D-97) and two SRAP loci (Me19Em5-359 and Me16Em8-483) were significantly associated with the green period with effect values of 79.54%, 62.59%, 99.04%, 49.01% and 82.57%. These markers will be useful for genetic improvement of the cold tolerance and green period of zoysiagrass by marker-assisted breeding.     

Genes for scald resistance from wild barley (Hordeum vulgare ssp spontaneum) and their linkage to isozyme markers

Summary Accessions of Hordeum vulgare ssp. spontaneum, the wild progenitor of barley, collected in Israel (70), Iran (15) and Turkey (6) were screened for seedling response to four isolates of Rhynchosporium secalis, the pathogen causing leaf scald in barley. Resistance was very common in the collection (77%) particularly among accessions from the more mesic sites (90%). The genetics of this resistance were investigated in fifteen backcross (BC₃) lines that contained an isozyme variant from H.v. ssp. spontaneum in a H.v. ssp. vulgare (cv. Clipper) background and were resistant to scald. Segregation in the BC₃F₂ families conformed with a single dominant resistance gene in 9 of the 15 lines. Scald resistance and the isozyme marker were closely linked in three of the BC₃-lines, loosely linked in four and unlinked in the remaining eight. Scald resistance genes were identified on barley chromosomes 1, 3, 4 and 6. Crosses between several of the scald resistant BC-lines together with the linkage data indicated that at least five genetically independent resistances are available for combining together for deployment in barley. The linkage of scald resistance in several BC₃-lines to the isozyme locus Acp2 is of special interest as this locus is highly polymorphic in wild barley.     

Conservation of diversity in bulk populations of barley (hordeum vulgare L.)

Summary A series of experiments was conducted to investigate the effectiveness of mass propagated heterogeneous populations (bulk populations) in preserving genetic and phenotypic diversity. Five genetically broad-based bulk populations of cultivated barley (Hordeum vulgare L.) were employed. All of them were produced originally through mass hybridization that was facilitated by male sterility. Four of these populations had a common origin, but were subsequently propagated in different North American locations. Comparisons between early and advanced generations of these bulk populations revealed loss of variability in all populations for morphological and agronomic characters and very little to none for eight isozyme characters. Populations propagated in different locations differed in levels of residual diversity. The bulk populations were less variable than a random sample of parental accessions. The rapid loss of diversity in bulk populations was considered detrimental to genetic conservation.     

Characterization of resistance genes against scald (Rhynchosporium secalis (Oudem.) J.J. Davis) in barley (Hordeum vulgare L.) lines from central Norway, by means of genetic markers and pathotype tests

Rhynchosporium secalis is a serious pathogen of barley (Hordeum vulgare L.) in central Norway. A breeding effort was initiated in 1977 to introduce resistance from different sources into adapted genotypes, and the first cultivar from the program was recently released. However, little is known about the resistance genes introgressed in this cultivar or in advanced breeding lines. An effort was made to address this issue through a set of isolates and available molecular markers. Fourteen breeding lines and their resistance donors were investigated by evaluating their reactions to 11 R. secalis isolates. Bulked segregant analysis was used to identify molecular markers linked to resistance genes in 12 of the breeding lines. The isolates were found to be of less discriminating value than the markers. Useful information has been obtained as to the nature of several of the resistance genes introgressed. Eight of the 12 breeding lines contained introgressed genes that were located at the `complex Rh' locus on chromosome 3H and hence may not easily be pyramided into the same genotype. Previous information about the nature of the resistance in `Jet' is questioned. Neither of the resistance genes Rh or Rh2 seems to have been incorporated into Norwegian breeding material.     

Diallel analysis of partial resistance to powdery mildew caused by Erysiphe graminis f. sp. hordei in spring barley (Hordeum vulgare L.)

Summary Six spring barley genotypes with growth stage dependent expression of partial resistance to powdery mildew were crossed following a half diallel scheme. Data for percentage infected leaf area at different growth stages from the F1 and F2 generation and the F2 offspring were analyzed. The absence of a contrast between parents and offspring indicated that dominance effects were not important at the young growth stages. Diallel analysis showed that general combining ability effects (GCA) were important at all growth stages. Although significant, specific combining ability effects were of minor importance. The cross between the most resistant parents with the largest, negative GCA resulted in the most resistant progeny.