Associations between three ml-o powdery mildew resistance genes and agronomic traits in barley

Summary A population of 198 chromosome-doubled haploid lines of spring barley was scored for segregation in locus ml-o (powdery mildew reaction) on chromosome 4 and in the linked loci s (rachilla hair length) and ddt (reaction to the insecticide DDT) on chromosome 7. They were also tested in a disease-free field trial for the agronomic traits: grain yield, thousand grain weight, lodging, and necrotic leaf spotting. The three mutagen-induced resistance genes ml-o5, ml-o6 (from Carlsberg II) and ml-10 (from Foma) showed no detectable differences with respect to effects on agronomic traits. They all conferred a four per cent reduction in grain yield caused mainly by lower thousand grain weight, and an increase in necrotic leaf spotting. The two original mutants of Carlsberg II had additional mutant genes affecting agronomic traits. Lines with gene S (long hair) had on average a three per cent higher thousand grain weight than those with s. The alleles in locus ddt showed no association with the agronomic traits. It is concluded i) that the associations between the three ml-o alleles and agronomic traits are caused by pleiotropy, ii) that ml-o resistant, high-yielding lines may be selected, and iii) that the association between gene s and thousand grain weight may be due to genetic linkage.Abbreviations DH-lines chromosome-doubled haploid lines     

Further evidence of polygenic inheritance of partial resistance in barley to leaf rust,Puccinia hordei

Summary The latent period (LP) is a crucial component of partial resistance. Five cultivars, L94, Sultan (Su), Volla (Vl), Julia (Ju) and Vada (Va), representing the known range in partial resistance and LP were crossed in a diallel, and the F₁, F₂ and F₃ tested. The LP effectuated by the five cultivars is about 9, 10^1/2, 10^1/2, 13 and 15^1/2 days, respectively. The crosses Su×L94, Vl×L94 and Ju×L94 had an F₂ positively skewed. Their F₂ means were similar or only slightly larger than the F₁ means. The F₂ frequency distributions in the crosses Vl×Su, Ju×Su and Ju×Vl were normal or nearly so with F₁ and F₂ means similar to each other and to the mid-parent value. The crosses involving Va as a parent again showed a positive skewness but with F₂ means considerably larger than the F₁ moans.Most F₂'s ranged from the low parent to the high parent values without transgression. In the crosses Va×L94 (reported earlier) and Ju×L94 the parental values were not recovered among 216 and 154 F₂ plants, respectively. The cross Ju×Va showed transgression beyond the low parent, Ju.From these data it is concluded, assuming no linkage, that seven loci are involved. The + alleles (governing a longer LP) are thought to be distributed over the parents as follows: % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGceaqabeaacaqGmb% GaaeyoaiaabsdacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabcca% caqGGaGaaeiiaiaab2cacaqGTaGaaeiiaiaabccacaqGGaGaaeiiai% aabccacaqGGaGaaeiiaiaabccacaqGTaGaaeylaiaabccacaqGGaGa% aeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaaeylaiaab2caca% qGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaa% b2cacaqGTaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaae% iiaiaabccacaqGTaGaaeylaiaabccacaqGGaGaaeiiaiaabccacaqG% GaGaaeiiaiaabccacaqGGaGaaeylaiaab2cacaqGGaGaaeiiaiaabc% cacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGTaGaaeyl% aiaabccaaeaacaqGtbGaaeyDaiaabccacaqGGaGaaeiiaiaabccaca% qGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGRaGaae4kaiaa% bccacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGRaGaae% 4kaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabUcacaqG% RaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaab2% cacaqGTaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaaeii% aiaabccacaqGTaGaaeylaiaabccacaqGGaGaaeiiaiaabccacaqGGa% GaaeiiaiaabccacaqGGaGaaeylaiaab2cacaqGGaGaaeiiaiaabcca% caqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGTaGaaeylaa% qaaiaabAfacaqGSbGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqG% GaGaaeiiaiaabccacaqGGaGaaeiiaiaabUcacaqGRaGaaeiiaiaabc% cacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabUcacaqGRaGaaeii% aiaabccacaqGGaGaaeiiaiaabccacaqGGaGaaeylaiaab2cacaqGGa% GaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabUca% caqGRaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaaeiiai% aab2cacaqGTaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGa% aeiiaiaabccacaqGTaGaaeylaiaabccacaqGGaGaaeiiaiaabccaca% qGGaGaaeiiaiaabccacaqGGaGaaeiiaiaab2cacaqGTaaabaGaaeOs% aiaabwhacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGa% GaaeiiaiaabccacaqGGaGaae4kaiaabUcacaqGGaGaaeiiaiaabcca% caqGGaGaaeiiaiaabccacaqGGaGaae4kaiaabUcacaqGGaGaaeiiai% aabccacaqGGaGaaeiiaiaabccacaqGRaGaae4kaiaabccacaqGGaGa% aeiiaiaabccacaqGGaGaaeiiaiaabccacaqGRaGaae4kaiaabccaca% qGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGRaGaae4kaiaa% bccacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaab2cacaqGTaGaae% iiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqG% GaGaaeylaiaab2caaeaacaqGwbGaaeyyaiaabccacaqGGaGaaeiiai% aabccacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabUcacaqGRaGa% aeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabUcaca% qGRaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaae4kaiaa% bUcacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqGGaGaae% 4kaiaabUcacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabccacaqG% GaGaaeylaiaab2cacaqGGaGaaeiiaiaabccacaqGGaGaaeiiaiaabc% cacaqGGaGaae4kaiaabUcacaqGGaGaaeiiaiaabccacaqGGaGaaeii% aiaabccacaqGGaGaaeiiaiaabUcacaqGRaaaaaa!1BBA!\[\begin{gathered} {\text{L94 - - - - - - - - - - - - - - }} \hfill \\ {\text{Su + + + + + + - - - - - - - - }} \hfill \\ {\text{Vl + + + + - - + + - - - - - - }} \hfill \\ {\text{Ju + + + + + + + + + + - - - - }} \hfill \\ {\text{Va + + + + + + + + - - + + + + }} \hfill \\ \end{gathered} \]The genes are supposed to act additively (intermediate inheritance) with the exception of one locus (the 6th or 7th locus) which shows dominance for the shorter LP (for the-alleles). The effect of this locus on LP seems considerably larger than that of the other loci. There are indications of physiological barriers, which means that LP's shorter than the one of L94 or much longer than that of Va are not possible.The effect of + genes in genotypes governing LP's close to these barriers (with very few or very many + alleles respectively) is smaller than in genotypes governing intermediate LP's.     

An iron chelating compound released by barley roots in response to Fe‐deficiency stress

The excretion of phytosiderophores by barley (Hordeum vulgare L.) has recently been documented and a major difference in the Fe‐stress response of gramineous species and dicotyledonous species proposed. However, currently used methods of quantifying and measuring phytosiderophore are tedious or require specialized equipment and a cultivar easily accessible to U.S. scientists is needed. The objectives of this study were (a) to determine if “Steptoe”; and “Europa”; (used as a control cultivar) barleys would release Fe³⁺ solubilizing compounds in response to Fe‐deficiency stress and (b) to develop a technique to determine the efficiency of solubilization of Fe(OH)₃ by the released chelating substances. Two cultivars of barley were place under Fe‐stressed (‐Fe) and nonstressed (+Fe) conditions in modified Hoagland solutions (14 L). The solutions were periodically monitored for H⁺ and reductant release from the roots and plants were rated daily for chlorosis development. Periodic (6 or 7 harvests) evaluation of the release of Fe³⁺ solubilizing substances was performed as herein described. Neither H⁺ nor reductant extrusion occurred with either cultivar during Fe stress. However, Fe³⁺ solubilizing substances were released by both cultivars at relatively high levels under Fe‐stress conditions compared to the nonstressed plants. A convenient technique was developed to measure the release of Fe solubilizing substances released by barley roots.     

Comparative effect of drought duration on growth,photosynthesis, water relations,and solute accumulation in wild and cultivated barley species

Drought is a major factor limiting crop production worldwide. Barley is a well‐adapted cereal that is largely grown on dry marginal land where water and salinity are the most prevalent environmental stresses. This study was carried out to investigate the effects of drought stress and subsequent recovery on growth, photosynthetic activity, water relations, osmotic adjustment (OA), and solute accumulation of wild (Hordeum maritimum) and cultivated barley (H. vulgare L.). In a pot experiment, 60 d old seedlings were subjected to drought stress for 0, 7, 14, 21, or 28 d, and then re‐watered to recover for up to 21 d. Plants were harvested at the end of each of these drought/recovery treatments. Drought significantly reduced fresh and dry weights at the whole‐plant level, photosynthetic activities, and solute and water potentials, while increasing leaf Na⁺ and K⁺ concentrations. The adverse effects of drought on growth were more marked in cultivated barley than in wild barley and the reverse was true for photosynthetic activities. During recovery, all wild barley seedlings completely recovered. For cultivated barley seedlings, rehydration had a beneficial effect on growth and photosynthesis, independent of treatment duration, but complete recovery did not occur. The reduction in leaf solute potential at full turgor in drought‐stressed barley, relative to the control, suggests active OA which was more significant in wild barley than in cultivated barley. OA was mainly due to the accumulation of inorganic (K⁺ in cultivated barley and Na⁺ in wild barley) and organic (soluble sugars and proline) solutes. The results suggest that OA is an important component of the drought‐stress adaptation mechanism in wild barley, but is not sufficient to contribute to drought tolerance in cultivated barley. In the latter species, the results show that even short periods (as little as 7 d) of water deficit stress had considerable long‐term effects on plant growth.     

Powdery Mildew Resistance in a Collection of Chinese Barley Varieties

One hundred and forty-seven Chinese barley varieties maintained at the Gene Bank of the National Barley Improvement Centre, Zhejiang, and 84 progenies from these varieties were tested at the seedling stage for their reaction to 32 selected pathotypes of Blumeria graminis f. sp. hordei. Eighteen resistance spectra were identified comprising single or combined resistances from eight known (Ml(Bw), Ml(Ch), Mla7, Mla8, Mla9, Mla13, MlaRu4 and Mlg) and six unknown resistance genes. The most frequent gene was Ml(Bw), which was found in 69 varieties and previously detected in only a few European winter barley varieties. The genes Mla8 and Ml(Ch) were also often present, but other resistance genes were rare. After inoculation, considerably fewer pathogen colonies were observed in ‘Aiganqi’ and one line of ‘Tong5’. Twenty varieties were composed of lines with different resistance genes. China is likely to be a region of origin of the genes Ml(Bw), Mla7, as well as three unknown genes found in original landraces and perhaps another three unknown genes detected in cultivars bred in China. The resistances of varieties from the Zhejiang province and those originating from 11 other Chinese provinces were quite different. Unfortunately, none of the varieties are promising sources of resistance to powdery mildew and China does not seem to be a region suitable for identifying such sources.     

Genetic variability evaluation in a Moroccan collection of barley, Hordeum vulgare L., by means of storage proteins and RAPDs

The genetic variation existing in a set of barley (Hordeum vulgare L.) landrace samples recently collected in Morocco was estimated. Two kinds of genetic markers, seed storage proteins (hordeins) and random amplified polymorphic DNA (RAPD), were used. Only six out of 31 landraces were subjected to RAPD analysis. Both kinds of markers, RAPD and storage proteins, yielded similar results, showing that the level of variation observed in Moroccan barley was high: all landraces showed variability; 808 different storage protein patterns (multilocus associations) were observed among 1897 individuals (2.32 seeds per association, on average) with an average of 43 multilocus associations per accession. In general, genetic variation within accessions was higher than between accessions. The 100 polymorphic RAPD bands generated by 21 effective primers were able to generate enough patterns to differentiate between uniform cultivars and even between individuals in variable accessions. One of the aims of this work was to compare the effectiveness of RAPD versus storage protein techniques in assessing the variability of genetic resource collections. On average hordeins were more polymorphic than RAPDs: they showed more alternatives per band on gels and a higher percentage of polymorphic bands, although RAPDs supply a higher number of bands. Although RAPD is an easy and standard technique, storage protein analysis is technically easier, cheaper and needs less sophisticated equipment. Thus, when resources are a limiting factor and considering the cost of consumables and work time, seed storage proteins must be the technique of choice for a first estimation of genetic variation in plant genetic resource collections.     

The influence of nitrogen fertilisation on bacterial activity in the rhizosphere of barley

The effect of nitrogen addition on the activity of rhizosphere bacteria was studied using barley seedlings. Three different nitrogen sources were added to the soil (nitrate, ammonium and ammonium+nitrate) at four different concentrations (0, 100, 300 and 500 mg N kg⁻¹ soil) and the plants were allowed to grow for 6 weeks. The bacterial activity was estimated by measuring thymidine and leucine incorporation into bacteria extracted using homogenisation-centrifugation. Bulk soil bacterial activity was low compared with that of rhizosphere bacteria. Nitrogen addition did not affect the activity of the bulk soil bacteria, indicating that the activity was not nitrogen limited. The thymidine and leucine incorporation rates of rhizosphere bacteria decreased when ammonium or ammonium+nitrate was applied compared with the non-amended controls. No effect on bacterial activity was found following nitrate addition. There was a significant positive correlation between rhizosphere bacterial activity and rhizosphere pH. Shoot length following ammonium treatment was significant lower than in the non-amended control, while nitrate and ammonium+nitrate addition had no effect. This indicates that the varying effects due to nitrogen sources on rhizosphere bacterial activity were not due to effects on plant growth.     

Evaluation of floral characteristics of barley in the semi-arid climate of north Syria

Previous studies have revealed that, in areas affected by severe drought, yielding ability and stability of barley (Hordeum vulgare L.) could be improved by increasing the level of heterozygosity. One possible approach to increase the heterozygosity level is to improve the outcrossing rate. In view of that, the genetic variability and phenotypic correlation coefficients involving four floral traits related to outcrossing behaviour were investigated in 274 drought adapted barley lines from the International Center for Agricultural Research in the Dry Areas (ICARDA's) current breeding programme. A significant quantitative variation existed in anther length and width, stigma length, and anther extrusion, which indicate that selection for these traits may be effective. Anther extrusion showed weak but highly significant positive correlations with anther and stigma length. Somewhat higher positive correlations were found between anther and stigma length. The results indicate that selection for high anther extrusion and longer anthers and stigmata may increase outcrossing in cultivated barley and consequently improve yielding ability through enhanced heterozygosity.     

不同酸碱条件下内生真菌对三种禾草种子萌发的影响

内生真菌可以通过提高禾草种子在干旱、盐胁迫等逆境条件下的萌发和促进植株生长而提高宿主的竞争力。本试验在pH为4~11的8个酸碱梯度下,对带菌(E+)与不带菌(E-) Neotyphodium属内生真菌的醉马草、中华羊茅和野大麦的萌发进行了研究,旨在探讨不同酸碱条件下内生真菌对禾草种子萌发的影响。结果表明,弱酸至强碱胁迫(pH为6~11)下,内生真菌显著提高醉马草种子发芽势;酸胁迫(pH为4~5)下,显著提高种子发芽率(P<0.05);中性至强碱胁迫(pH为7~11)下,显著提高种子的胚芽长及胚根长(P<0.05);酸性至中性条件(pH为4~7)下,显著提高幼苗干重;强酸(pH=4)及强碱(pH=11)胁迫下,E+异状发芽率显著低于E- (P<0.05)。弱酸至碱胁迫(pH为6~10)下,内生真菌显著提高中华羊茅种子发芽势(P<0.05);强酸(pH=4)及碱胁迫(pH为8~11)下,显著提高种子发芽率(P<0.05);pH为4~11,显著促进中华羊茅胚芽生长(P<0.05); pH=10时对胚根的增益作用显著(P<0.05);酸胁迫(pH为4~6)下,显著提高幼苗干重;强碱胁迫(pH=11)下,异状发芽率显著增加(P<0.05)。pH为4~11,内生真菌显著提高野大麦种子发芽势(P<0.05);碱胁迫(pH为9~11)下,显著促进种子胚芽生长(P<0.05);碱胁迫(pH为8~11)下,显著提高胚根长(P<0.05);pH为4~11,显著提高野大麦幼苗干重(P<0.05);强酸胁迫(pH=4)下,E+异状发芽率显著低于E- (P<0.05)。相对于胚芽,胚根对酸碱条件变化更敏感。综合考虑,最适萌发pH条件分别为:醉马草pH为6~9;中华羊茅pH为6~7;野大麦pH为7~9。     

Analysis of Genetic Diversity in Barley Cultivars Reveals Incongruence Between S-SAP,SNP and Pedigree Data

Accurate assessment of genetic similarity is important for plant breeding, germplasm enhancement and conservation of plant genetic resources. A comparative analysis of genome diversity among a group of six-rowed spring barley (Hordeum vulgare L.) cultivars was carried out using sequence-specific amplified polymorphism (S-SAP) and single nucleotide polymorphism (SNP), with the results compared to the kinship coefficients derived from the pedigree data. Mean pair-wise GS values were estimated to be 0.0957 ± 0.144 (Kinship), 0.491 ± 0.189 (SNPs), and 0.602 ± 0.098 (S-SAPs). S-SAP and SNP-based genetic similarity (GS) values were normally distributed but kinship values had a non-normal and skewed distribution. Pair-wise correlation of GS values were lowest for the S-SAP and the SNP matrices (r =; 0.040, p<0.230) and highest for the SNP and pedigree matrices (r =; 0.240, p < 0.001). Analysis of molecular variance (AMOVA) attributed about 90.4% of observed variation to the cultivars within each of the malting and feed groups. Variance component between malting and feed groups was 6.6% for both SNP and S-SAP data suggesting lack of a significant genetic differentiation along this agronomic division. The remaining 3% of variation was attributed to genetic diversity within cultivars. Although both DNA-based marker systems were able to differentiate all barley cultivars, significant difference were observed in the pattern of genetic relationships obtained by the two marker systems and the pedigree data.